: ■

APRIL 2003

BOARD OF EDITORS

Editor

J.C. DANIEL

M.R. ALMEIDA

M.K. CHANDRASHEKARAN

B.F. CHHAPGAR

R. GADAGKAR

INDRANEIL DAS

A.J.T. JOHNSINGH

AJITH KUMAR

T.C. NARENDRAN

A.R. RAHMANI

J.S. SINGH

R. WHITAKER

Assistant Editor

GAYATRI WATTAL UGRA

INSTRUCTIONS TO CONTRIBUTORS

1 . Papers which have been published or have been offered for publication elsewhere

should not be submitted.

2. Papers should be submitted in duplicate, typed double space. Preferably an additional

copy should be submitted on a floppy diskette (3.5") using MS Word.

3. Trinomials referring to subspecies should only be used where identification has been

authentically established by comparison of specimens actually collected.

4. Photographs for reproduction must be clear, with good contrast. Prints should be at

least 9 x 12 cm and on glossy glazed paper. Text-figures, line drawings and maps

should be in Indian ink, preferably on tracing paper.

5. References to literature should be placed at the end of the paper, alphabetically arranged

under author’s name, with the abridged titles of journals or periodicals in italics and

titles of books or papers in roman type, thus:

Aluri, Raju J.S. & C. Subha Reddi (1995): Ecology of the pollination in two cat-mint

species. J. Bombay nat. Hist. Soc. 92(1): 63-66.

Prater, S.H. (1948): The Book of Indian Animals. Bombay Natural History Society,

Mumbai, pp. 35-48.

6. Each paper should be accompanied by an abstract, normally not exceeding 200 words,

and 6-8 key words. Key words should include the scientific names of important species

discussed.

7. 25 reprints will be supplied free of cost to authors of main articles and new descriptions.

Authors of miscellaneous notes, will be sent a free copy of the Journal.

8. The editors reserve the right, other things being equal, to publish a member’s contribution

earlier than a non-member’s.

9. For the standardised common and scientific names of the birds of the Indian subcontinent

refer to Buceros\/ ol. 6, No. 1 (2001).

Hornbill House,

Shaheed Bhagat Singh Road,

Mumbai 400 023.

Editors,

Journal of the Bombay

Natural History Society

VOLUME 100 (1): APRIL 2003

CONTENTS

EDITORIAL : 1

BREEDING BEHAVIOUR OF THE GREATER ADJUTANT-STORK LEPTOPTILOS

DUBIUS IN ASSAM, INDIA

( With one text-figure and two plates)

By Hillaljyoti Singha, Asad R. Rahmani, Malcolm C. Coulter and Salim laved 9

CURRENT STATUS OF THE GANGES RIVER DOLPHIN, PLATANISTA GANGET1CA IN

THE RTVERS KOSI AND SON, BIHAR, INDIA

( With one text-figure)

By R.K. Sinha and Gopal Sharma 27

CROP DEPREDATION BY WILDLIFE ALONG THE EASTERN BOUNDARY OF THE

KALAKAD-MUNDANTHURAI TIGER RESERVE, SOUTHERN INDIA

(With two text-figures)

By Punidan D. Jeyasingh and Priya Davidar 38

SMALL MAMMALS IN MONTANE ECOSYSTEMS OF THE NILGIR1S, SOUTHERN

INDIA: THEIR ECOLOGY AND NATURAL HISTORY

By Kartik Shanker 46

THE EFFECTS OF CATTLE GRAZING AND HABITAT ON HELMINTH LOADS OF

CHITAL (AXIS AXIS) IN THE MUDUMALAI WILDLIFE SANCTUARY,

SOUTHERN INDIA

By Guha Dharmarajan, M. Raman and Mathew C. John 58

DEMOGRAPHY OF LIONTAILED MACAQUE (MAC AC A SILENUS) IN AN

UNDISTURBED RAINFOREST OF SILENT VALLEY NATIONAL PARK,

KERALA, INDIA

( With two text-figures)

By Gigi K. Joseph and K.K. Ramachandran 65

NEW DESCRIPTIONS

A NEW HUMAN BLOOD FEEDING BITING MIDGE FROM INDIA, DIPTERA:

CERATOPOGONIDAE: FORCIPOMYIA MANASI

(With three texU figures)

By G Irish Maheshwari 72

A NEW SPECIES OF EUGENIA L., MYRTACEAE, FROM SEITHUR HILLS,

TAMIL NADU, INDIA

( With one text-figure)

By R. Gopalan and S.R. Srinivasan 78

SONERILA LONGIPETIOLATA MANICKAM ET AL A NEW SPECIES OF

MELASTOMACEAE FROM TAMIL NADU, INDIA

( With one text-figure)

By M.M. Josephine, V.S. Manickam, C. Murugan, V. Sundaresan and G.J. Jothi 81

A NEW SPECIES OF SPIDER OF THE GENUS TIB ELL US SIMON (ARANEAE:

THOMISIDAE) FROM JHENIDAH, BANGLADESH

( With six text-figures)

By V. Biswas and D. Raychaudhuri 84

FURTHER CONTRIBUTION TO BIOSYSTEMATICS OF CHEN OP ODIUM, REPORTING

THREE NEW SPECIES FROM NORTH INDIAN PLAINS

( With three text-figures)

By S.C. Pandeya and Amita Pandeya 87

OBITUARY

PROF. M.S. MANI (1908-2003)

REVIEWS

94

1 . BEAUTIFUL ORCHIDS OF NEPAL

Reviewed by M.R. Almeida 96

2. A BIBLIOGRAPHY OF THE PLANT SCIENCE OF NEPAL

Reviewed by M.R. Almeida 96

3. AROMATIC AND MEDICINAL PLANTS — YIELDING ESSENTIAL OIL FOR

PHARMACEUTICAL, PERFUMERY, COSMETIC INDUSTRIES AND TRADE

Reviewed by M.R. Almeida 96

4. RAPTOR WATCH: A GLOBAL DIRECTORY OF RAPTOR MIGRATION SITES

Reviewed by Rishad Naoroji 98

MISCELLANEOUS NOTES

MAMMALS

1. The Harvard collection of South Asian

mammals

By Kristofer M. Helgen, John Mathew

and Christine A. Monta 100

BIRDS

2. Little grebe Tachybaptus ruficollis\ An

addition to the avifauna of Ladakh

By Harkirat Singh Sangha,

Rishad Naoroji and Maan Barua 104

3. Sighting of lesser frigatebird Fregata ariel

Gray in the Kole Wetlands of Thrissur, Kerala

By E.A. Jayson and C. Sivaperuman 106

4. Indian pond-herons Ardeola gray 'd feeding on

dragonflies

By V. Santharam 108

5. Variation recorded in the iris, bill, leg and foot

colouration in cattle egret Bubulcus ibis

By Tushar M. Sanghani 109

6. Record of a nesting colony of painted stork

Mycteria leucocephala at Man-Marodi Island

in the Gulf of Kutch

By Abdul Jamil Urfi 109

7. First sighting of lesser adjutant-stork

Leptoptilos javanicus from Sanjay Gandhi

National Park, Mumbai

By Anish Andheria 1 1 1

8 . Black ibis Pseudibis papillosa feeding on frogs

from crab holes

By J. Mangalraj Johnson Ill

9. Note on breeding of Andaman teal Anas

gibberifrons in south Andaman Islands, India

By Sarang Kulkami and Manish Chandi 112

10. Some observations on nesting of Bonelli’s

eagle Hieraaetus fasciatus (Vieillot)

By Deepali Kulkarni and Banda Pednekar .... 113

11. Status of white-bellied sea-eagle Haliaeetus

leucogaster in Ratnagiri district, Maharashtra

By Vishwas Katdare and Ram Mone 1 13

12. Cannibalism in Indian white-backed vulture

Gyps bengalensis in Keoladeo National Park,

Bharatpur, Rajasthan

By Gargi Rana and Vibhu Prakash 116

13. Wintering site fidelity in western marsh-harrier

Circus aeruginosus (Linn.), in Keoladeo

National Park, Bharatpur, Rajasthan

By Ashok Verma 1 1 7

14. Southernmost record of eastern calandra-lark

Melanocorypha bimaculata and sighting of

lesser kestrel Falco naumanni from Matheran,

a hill station near Mumbai

By Anish Andheria 118

15. Unusual feeding association between Siberian

crane Grus leucogeranus and wild boar Sus

scrofa in Keoladeo National Park, Bharatpur,

Rajasthan

By Gargi Rana and Vibhu Prakash 120

16. Common coot Fulica atra from Kyongnosla

in East Sikkim

By Usha Ganguli-Lachungpa and

Bishnu Lai Sharma 121

u

17. Sighting of the lesser florican Sypheotides

indica in Gir interpretation zone, Devaliya, Gir

Sanctuary

By B.P. Pati

1 8. Grey-headed lapwings Vanellus cinereus seen

around Machilipatnam, Krishna District,

Andhra Pradesh, India

By Colin Conroy

19. The grey-headed lapwing in Tamil Nadu - A

rejoinder

By V. Santharam

20. A sight record of buff-breasted sandpiper

Tryngites suhruficollis in Goa

By Paul I. Holt and Rick Heil

21. Group fishing of house crows ( Corvus

splendens) with river terns ( Sterna aurantia)

By Sattyasheel N. Naik

22. Terns of the Vengurla Rocks, a review and

update

By Heinz Lainer

23. Range extension of Alexandrine parakeet

Psittacula eupatria in Gujarat

By Aeshita Mukherjee, C.K. Borad and

B.M. Parasharya

24. Mimicry of a crow chick by an Asian koel

Eudynamys scolopacea as a defence against

attack by house crows Corvus splendens

By Colin Ryall

25. Buffy fish-owl ( Ketupa ketupu) in Sundarbans,

Bangladesh

By Gertrud Neumann-Denzau and

Helmut Denzau

26. A note on the circumorbital skin colour of

Indian grey hornbill Ocyceros birostris

By Aasheesh Pittie

27. Sighting of Malabar pied hornbill

Anthracoceros coronatus in Sanjay Gandhi

National Park, Mumbai

By Anish Andheria, Supriya Jhunjhunwala

and Paritosh Khanvilkar

28. Nocturnal feeding by white-bellied drongo

Dicrurus caerulescens

By Satish Kumar Sharma

29. Total albinism in large grey babbler Turdoides

malcolmi

By Satish Kumar Sharma

30. Yellow-rumped flycatcher Ficedula

zanthopygia in Kerala

By Paul I. Holt

31. Breeding records of the Asian brown

flycatcher Muscicapa dauurica in southern

India

By V. Santharam

32. Crimson sunbird Aethopyga siparcija seheriae

in Madhya Pradesh

By E.P. Eric D’Cunha 147

33. Incubation period of crimson sunbird

Aethopyga siparaja

By Vishwas Katdare, Rohan Lovalekar and

Ameya Modak 148

34. The Spanish sparrow Passer hispaniolensis

found nesting in Hanumangarh district,

Rajasthan

By Manoj Kulshreshtha and

Harkirat Singh Sangha 149

35. Termite removal from nest material and repair

of damaged nest by white-rumped munia

Lonchura striata

By R. Shyama Prasad Rao 151

36. Birds of Goa — Some supplementary notes

By V. Santharam 151

37. Birds of Tabo: A lesser known cold desert in

the western Himalaya

By Arun P. Singh 152

OTHER INVERTEBRATES

38. Occurrence of Triops granarius (Lucas),

Crustacea: Notostraca, from Madurai, Tamil

Nadu

By C.S. Velu and N. Munuswamy 154

39. Occurrence of rare jumping spider

Harmochirus brachiatus (Thorell) (Family:

Salticidae) in the banana agro-ecosystem of

Vadodara, Gujarat

By Siliwal Manju and Dolly Kumar 157

40. Description of female Amyciaea forticeps

(Cambridge), Araneae: Thomisidae, with a

redescription of its male from Kerala, India

By K. Sunil Jose, Samson Davis,

A.V. Sudhikumar and P.A. Sebastian 157

41. Rare sighting of ogre-faced spider Dinopis

goalparaensis , Araneae: Dinopidae, in the

banana agro-ecosystem of Vadodara, Gujarat

By Siliwal Manju and Dolly Kumar 160

BOTANY

42. Observations on Bauhinia malabarica Roxb,

Leguminosae: Caesalpinioideae. shape of

calyx is not correlated with sexual nature of

flowers

By S. Bandyopadhyay 161

43. Rediscovery of Ceropegia evansii McCann,

Asclepiadaceae, from Maharashtra

By P. Tetali, Sujata Tetali,

P. Lakshminarasimhan, P.V. Prasanna

and B.G. Kulkarni 162

122

123

126

135

136

138

141

142

144

145

146

Cover Photograph: Waterfall by Varad Giri

ACKNOWLEDGEMENT

We are grateful to the Ministry of Science and Technology,

Govt of India,

FOR ENHANCED FINANCIAL SUPPORT FOR THE PUBLICATION OF THE JOURNAL.

CITATION OF I C/EC NUMBERS FOR GENETIC MATERIALS

It is brought to our notice by the National Bureau of Plant Genetic Resources (NBPGR),

Pusa Campus, New Delhi 110 012, India, that authors writing papers on particular plant

materials (genetic materials) should indicate IC numbers for Indigenous Collections and

EC numbers for Exotic Collections. Authors can directly procure these single accession

numbers for each genetic material from NBPGR. In the present Intellectual Property

Rights regime, it is in our national interest that all the germplasm material possess a

single national accession number.

Authors are therefore requested to procure IC/EC numbers from NBPGR and state them

on the manuscript, without which papers will not be accepted for publication.

Editors

Editorial

A Web of Rivers

The idea of linking the rivers of India to provide water security to a region dependent

on the vagaries of the monsoon has so far been a dormant pipe dream. Beset by legal and

political problems over the waters of the Cauvery river, the highest judicial and administrative

authorities of the country seem to have clutched at a nebulous last straw, a project to link

the rivers of the country, as a palliative to a persistent problem.

In a memorandum to the Prime Minister, signed by 58 concerned conservationists,

including three retired bureaucrats who had been involved in the administration of the

water resources and environment of the country as Secretaries to the Govt of India, state :

“The idea of the ‘linking of rivers’, dormant for a long time, has acquired new

prominence now, particularly in the context of the acute form that the Cauvery dispute took

in the course of the year 2002, as well as the drought that afflicted several parts of the

country in that year. In response to a public interest writ petition, the Supreme Court has

desired that the project for the linking of the rivers of India be accelerated. The Prime

Minister has announced the setting up of a task force to consider the modalities of

implementing the project, and declared that it would be taken up “on a war footing”. The

Leader of the Opposition in the Lok Sabha has welcomed this undertaking.

The project has been the subject of much reporting and comment in the media in

recent months. It has been presented by the Government as a major initiative and the definitive

answer to the future water problems of the country, and it has been so hailed by some.

However, some others have expressed apprehensions. We, the signatories to this

memorandum, feel that this decision is fraught with serious consequences, and that the

Government should carefully reconsider it before proceeding further. Without commenting

on the Supreme Court’s observations in this case, we shall set forth our reasons for urging a

reconsideration of the decision by the Government.

Outline of Proposal

Our understanding of the project, derived partly from the Report (September 1999) of

the National Commission for Integrated Water Resources Development Plan (NCIWRDP)

and partly from presentations currently being made by governmental agencies, is briefly

outlined here for confirmation or correction.

Without going into the history of the idea of the linking of rivers of India, we note that

the ‘Ganga-Cauvery Link’ proposal mooted by Dr. K.L. Rao and the ‘Garland Canal’ idea

put forward by Captain Dinshaw Dastur were examined and found impractical, the former

on the grounds of the very large financial and energy costs involved, and the latter because

it was technically unsound; and that the proposal now taken up is based on the work that the

National Water Development Agency has been doing during the last two decades after its

establishment in 1982 in pursuance of the ‘National Water Perspectives’ brought out by the

Ministry of Irrigation in 1980. There are two main components in it, namely the Himalayan

Rivers component and the Peninsular Rivers component. The Himalayan component

envisages a number of links, including some within the Ganga system (Kosi-Ghagra, Gandak-

Ganga, Ghagra- Yamuna Sarda- Yamuna, etc); some between neighbouring rivers in the

Brahmaputra system (Manas-Sankosh-Teesta); a couple between those two systems (Teesta-

Ganga, and an alternative Brahmaputra-Ganga link); one long link from Sarda to Sabarmati

through the Yamuna and Rajasthan; one from the Ganga to Subamarekha via Damodar and

then on to Mahanadi; and a few others. The general idea is to transfer waters from ‘surplus’

eastern rivers to ‘deficit’ central, western and southern regions. The Peninsular Rivers

component again involves a number of links, of which the most important would be those

connecting Mahanadi, Godavari, Krishna, Pennar and Cauvery. The idea is to transfer the

surpluses estimated to exist in the Mahanadi and the Godavari to the deficit southern basins

(Cauvery, Vaigai). Other links in the Peninsular component would include Ken-Betwa,

Parbati-Kalisindh-Chambal, Par-Tapi-Narmada, Damanganga-Pinjal, etc. Another idea is

the partial diversion of certain rivers flowing into the Arabian Sea eastwards to link with

rivers flowing into the Bay of Bengal (Bedti - Varda, Netravati-Hemavati, Pamba - Achankovil

- Vaippar).

Mandate of the Task Force

We note that the Task Force has been asked to examine not the soundness or viability

of this project but the modalities of its implementation. Three main difficulties have been

recognized: the formidable challenge presented by the accelerated time-frame indicated by

the Supreme Court; the magnitude of the financial resources needed (roughly and tentatively

estimated at Rs. 5,60,000 crores); and the problem of bringing about the necessary political

consensus on the transfers involved. The Task Force appears to be concentrating on these

three tasks at present. However, there are some prior questions that need to be asked: Why

has this project been proposed? How did it emerge? How does it fit in with the national

planning process? Is it necessary and feasible, and is it likely to be beneficial on the whole?

As these questions seem beyond the mandate of the Task Force, we propose to raise them

here.

Sudden Emergence

The project appears to have suddenly emerged into prominence. If the Government

had been contemplating a monumental project of this kind, there would have been some

indications. There were none. The Ninth Plan made no reference to it. Even the Tenth Plan

(which lays special emphasis on water and wishes to be regarded as a ‘Water Plan’) refers to

many important approaches, policies, programmatic initiatives, and so on, but says nothing

about any river-linking project. The Prime Minister’s important Address to the National

Water Resources Council ( 1 April 2002) did not mention it. It seems clear that the Government

were not seriously thinking of any river-linking project. The NWDA’s proposals were non-

starters for various reasons. The Government’s own initial submissions to the Supreme

Court were very cautious and lukewarm. The Supreme Court’s direction (if its observations

can be so regarded) and the Government’s enthusiastic response to it have changed all that.

A project that was not on the anvil has suddenly become the most important undertaking of

the Government. This seems to us to be a bypassing of the planning process.

2

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

National Commission’s Observations

Not very long ago the high-level National Commission for Integrated Water Resources

Development Plan (NCIWRDP), the first national commission on water, set up in 1996,

submitted its Report (September 1999). Its Terms of Reference specifically included ‘Inter-

Basin Transfers’ as an item. It reviewed the NWDA’s studies. It did not discuss the proposed

Himalayan links in detail because the data are classified as confidential, but did observe that

the costs involved and the environmental problems would be enormous; that the further

expansion of irrigation in the desert areas of Rajasthan would need examination from all

angles; that the NWDA’s Himalayan component would require more detailed study; and

that the actual implementation was unlikely to be undertaken in the immediate coming

decades. On the Peninsular component, after a careful examination of the water balances of

the various basins, the Commission observed: “Thus there seems to be no imperative necessity

for massive water transfers. The assessed needs of the basins could be met from full

development and efficient utilization of intra-basin resources except in the case of Cauvery

and Vaigai basins. Therefore, it is felt that limited water transfer from Godavari at Ichampalli

and Polavaram towards the south would take care of the deficit in Cauvery and Vaigai

basins... Though surplus is available in Mahanadi also, the transfer from that river would

require much longer link and is in any case not required for the immediate future....” (The

Commission then takes note of some uncertainties that may affect the above judgment and

says that further studies as to the future possibilities of inter-basin transfers need to be

continued.) The decision to embark on this massive project “on a war footing” seems difficult

to understand in the light of those observations of the National Commission.

Rationale of Project

However, there is now a project, and we must consider its rationale. The project is

claimed to be the answer to the country’s problems of recurring floods and drought in

different areas; the generation of hydroelectric power is also put forward as a justification.

Neither flood control nor hydroelectric power calls for a linking of rivers. In the case

of hydroelectric power, the usual practice is to postulate a ‘potential’ in some rivers or areas

(for instance, Narmada, Brahmaputra, the Northeast of India, Nepal) and propose large

projects (Sardar Sarovar, Dihang, Subansiri, Tipaimukh, Karnali, Pancheswar, and so on)

to exploit that potential. Each such project will have to be looked at carefully, but what

needs to be noted in the present context is that while the need for hydroelectric power may

lead to the formulation of particular projects in specific locations, it would not by itself take

us to the idea of linking rivers. (Incidentally, the linking of rivers or inter-basin transfers

would in the generality of cases require much energy - normally in excess of what the

project might generate - but in this case we are told that the project will be a net generator

of large quantities of power: a figure of 30,000 MW has been mentioned. That strains our

credulity and will need careful examination with reference to each link.)

Similarly, the problem of recurring floods in certain rivers or areas may lead (rightly

or wrongly) to the formulation of specific projects with flood control as one of the objectives

(or a primary objective) — for instance, the DVC projects, a high dam on the Kosi, and so

on — and will not by itself call for a linking of rivers. It must also be noted that opinion on

JOURNAL , BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

3

flood control has changed over the years. It is now generally recognized that big dams play

only a modest role in flood-moderation; that even in those projects (not many) where flood

cushions have been built in, that cushion tends to get eaten into partly by excessive silting

and partly by the more powerful demands of irrigation and power generation; that

considerations of the safety of structures sometimes necessitate the release of waters causing

‘man-made’ floods downstream; that by and large, the old notion of ‘flood control’ has to

change to the newer ideas of learning to live with floods and minimizing damage; and that

this requires a relatively greater reliance on non-structural than on structural measures. By

now, this has almost become conventional wisdom. Even if all the river-linking proposals

are implemented, the contribution that this will make to the mitigation of the flood problem

will not be substantial. Dr. Bharat Singh, a doyen among engineers and the former Vice-

Chancellor of the Rourkee University, has observed: “Any water resources engineer will

immediately discard inter-linking of rivers as a flood control measure”.

As regards drought, we have the answers already. Rajendra Singh has shown in Alwar

District in Rajasthan that rainwater-harvesting can be practised successfully even in low-

rainfall areas. Earlier, Anna Hazare had brought about a transformation through water-

harvesting (along with other measures) in Ralegan Siddhi (which is also a low-rainfall

area). The Madhya Pradesh Government has initiated large Statewide programmes of water-

harvesting and conservation. In the water-scarce parts of Gujarat, some good NGOs have

remarkable achievements in this regard to their credit. Dhan Foundation has been doing

good work in the southern States. The large numbers of tanks in Tamil Nadu, Karnataka

and Andhra Pradesh were remarkable water-management systems that have gone into decline,

and efforts are on to restore and rehabilitate them. Similar efforts are also needed, and are in

progress, in respect of other traditional systems such as ahars and pynes in Bihar, johads in

Rajasthan, and so on.

In brief, the primary answer to drought has to be local; it is only thereafter, and in

some very unpromising places, that the bringing in of some external water may need to be

considered. Besides, the river-linking project, if implemented, will take water only to a

small part of the arid or drought-prone areas; large parts of such areas will remain unserved

and will have to meet their needs through the local augmentation of water availability. It

was in recognition of the importance of such local, community-led initiatives of rainwater-

harvesting and watershed-development that the Prime Minister strongly urged the promotion

of such initiatives on a nationwide basis in his Address to the National Water Resources

Council on 1 April 2002.

(Incidentally, the project as now outlined essentially envisages the addition of waters

to certain existing rivers. The additional waters will thus go to areas that are already being

served to some extent by that river or by a canal from a reservoir on that river. How will this

benefit the uplands and plateaux that are unserved by the existing rivers or are drastically

water-short? A glance at the two maps showing the proposed links does not provide a clear

answer to this question. However, it is being claimed that irrigation will be extended to

additional areas. This may well be true in the sense that areas unreached earlier in the

vicinity of a river or within the command area of a project may now receive some irrigation,

but will the waters reach the country’s drylands?)

4

JOURNAL , BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

A further point to be kept in mind is that it is not primarily drinking water needs but

the large demands of irrigation that lead to proposals for long-distance water transfers,

though the waters so transferred may also be used to meet drinking water requirements.

Water transfers for irrigation may be proposed either for providing additional water to areas

already under irrigation or for extending irrigation to arid or ‘rainfed’ areas. In both cases,

difficult questions arise.

In irrigated areas (for instance, the Cauvery basin), the question is whether large

demands for additional irrigation water should be unquestioningly accepted and met through

supply-side solutions such as large dams or inter-basin transfers, or a serious attempt made

to improve water-use efficiency in irrigated agriculture, get more value out of a given quantum

of water, reduce the water-demand, and minimize the need for supply-side projects. In the

context of the prevailing low efficiency of water-conveyance in canal systems and water-use

in irrigated agriculture, bringing in more water from another basin would really amount to

the provision of more water for being wasted. It would also mean that there would be no

motivation at all for changing cropping patterns and shifting from water-intensive crops to

crops that need less water; on the contrary, the tendency to grow water-consuming crops

would receive strong encouragement. (It may be added that cropping patterns and water-use

practices that lead to or aggravate water-scarcity are often the results of government policies

relating to agriculture and water, and what is called for is the rectification of those policies

rather than the importation of water.)

In arid or drought-prone areas, the introduction of irrigated agriculture of a kind

appropriate to wet areas may be unwise. ‘Development’ in arid areas should perhaps take

other, less water-intensive forms. The slogan of ‘making the desert bloom’ is not necessarily

a sound one. It can be argued that the Rajasthan Canal project was not a good idea but a

misconceived one. These are difficult but important questions that need careful consideration.

In both irrigated and rainfed areas, the bringing in of external water may also have

other secondary consequences: the need to bring in farmers from elsewhere and the resulting

social tensions (as in Rajasthan); increased incidence of conditions of water-logging and

salinity (a concomitant of irrigated agriculture in many places); the possibility of the repetition

of the ‘Green Revolution’ patterns of agricultural development and the related phenomena

of monoculture, loss of biodiversity (disappearance of indigenous varieties of seeds of plants

and grains), the problems arising from chemical fertilizers and pesticides, the loss of micro-

nutrients from soils, and the replacement of healthy indigenous varieties of food crops by

high-yielding, commercially viable, but nutritionally deficient crops; social inequities of

diverse kinds; and so on. These are not unavoidable consequences, but they are dangers that

have to be kept in mind.

Subject to all those caveats, the idea of taking water from ‘surplus’ to ‘deficit’ basins

may seem prima facie a good one. That indeed is the principal driving force behind

the project, and that is also what gives it its popular appeal in water-scarce States.

However, there are many serious difficulties with that plausible proposition, which need to

be noted.

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

5

Some Difficulties

Gigantism / Altering Nature

To start with, there is the fundamental objection, not to the idea of ‘inter-basin transfer’

per se (though that aspect does need consideration), but to the grandiose nature — the

gigantism — of the undertaking. This will be a massive intervention in nature, an ambitious

attempt to alter nature. That it is to be compressed into a short span of time may aggravate

the intervention but that is a secondary point, the main one being that it amounts to nothing

less than the redrawing of the geography of the country. It appears to us that this is a severe

case of technological hubris of a kind that (we thought) had been discredited and was a

thing of the past.

Criticisms of gigantism are sometimes responded to with the answer that no gigantism

is intended; that the project will proceed carefully and slowly, in a piecemeal manner, from

the minor and relatively less problematic links to the more difficult and ambitious ones. Is

such a careful, exploratory, step-by-step approach in fact intended? This seems inconsistent

with what we have been seeing and hearing in recent months: the Supreme Court’s desire

that the project be accelerated and the time-frame compressed; the Prime Minister’s

announcement that the project will be taken up on a war-footing; the setting up of a Task

Force; the references to the order of investments involved; the publicity surrounding the

project; and so on. It appears that the Government wants to make dramatic announcements,

and at the same time claim that it is adopting a slow, careful, modest, exploratory approach.

The general impression in the country is certainly that a massive project has been undertaken.

If that is not the case, the Government should make the position clear.

Strange Idea

There is in fact an oddity about the proposition that we have tended not to notice. One

can understand if the planners start from an identification of the needs of particular areas,

proceed through a consideration of options and alternatives, and finally arrive at a decision

to link two or more rivers as the only or the best option in a given case. Instead, the present

project starts with the proposition that the rivers of India must be linked, and then proceeds

to consider possibilities of storages, links, transfers, etc. What is the basis for that a priori

proposition (even if it is an old one)? How did we arrive at this strange idea that all the

rivers of India — or the major ones — must be linked? The analogy sometimes put forward

with the linking of highways or with a national power grid is inapt and misleading. Human

creations or productions such as highways or power can be manipulated by humans. That

does not necessarily apply to rivers. Rivers are not human artefacts; they are not pipelines to

be cut, turned around, welded and re-joined. They are natural phenomena, integral

components of ecological systems, and inextricable parts of the cultural, social, economic,

spiritual lives of the communities concerned. (So too are related features, both natural and

man-made, such as lakes, wetlands, tanks, beels , ahars and pynes, and so on.)

Serious Consequences

The project is potentially fraught with serious consequences. It will necessarily involve

dams, reservoirs, diversion of waters, canal systems, and so on. By now there is adequate

6

JOURNAL BOMBAY NATURAL HISTORY SOCIETY. 1 00(1). APR. 2003

knowledge of what all this entails: violent disturbance of pristine areas and of the lives of

(tribal) communities living there, disruption of the habitats and movement routes of wildlife,

loss of bio-diversity (flora and fauna), changes in river morphology and water quality (arising

from the stilling of flowing waters), submergence of forests and agricultural lands, changes

in the micro-climate, public health consequences, displacement of people and their livestock

and the related problems of resettlement and rehabilitation, reduction of downstream flows,

the consequent alteration of the river regime (reduction of the capacity of the river to cope

with pollutants and regenerate itself; reduction in nutrient content in downstream flows;

diminution of groundwater-recharging, reduction in freshwater outflows into the sea), and

the impacts of these on aquatic life, riparian communities and their livelihoods such as

agriculture or boat-plying, and on estuarine conditions (including estuarine fish populations)

and possible salinity incursions; and so on. These impacts and consequences have been

observed in many projects, and will need to be studied carefully in the case of each of the

proposed links.

(Incidentally, much harm has been done in the past by the tendency to regard only

water abstracted from the stream as ‘used’ and water flowing in the stream and particularly

into the sea as ‘wasted’. To minds so conditioned, the fact that floods occur in some areas

and drought is experienced elsewhere immediately suggests that water must be transferred

from the former to the latter places. Behind this lies an ignorance of the multiple purposes

served by flowing water — even floods — and the importance of water flowing into the sea,

and a failure to recognize the consequences of a diversion of flows. Rivers must flow if silt

is to move and nutrients are to reach the plains, the deltaic region, and mangrove areas such

as the Sunderbans. Such flows and nutrients also enter the coastal waters and contribute to

the increase of marine wealth, whether it be shoals of fish or algae and other organisms

which hold the key to the future nutritional, medicinal and other needs of our country and

even of humanity at large. Before diverting waters and reducing downstream flows, we

must make sure that the alluvial deltas will not die, forcing the migration of populations

and causing distress in the coming generations. Rivers must have enough water to support

riverports, inland navigation and riverine fauna and flora, and to check the incursion of

salinity in coastal areas. The concept that no water is to be allowed to go waste into the sea

needs to be seriously challenged on hydrological and meteorological grounds.)

It has been argued that similar projects have been undertaken elsewhere without

catastrophic consequences, but that is a questionable statement. Water-resource projects are

part of the kind of ‘development’ that the world has been pursuing, which has in fact had

many catastrophic consequences. But leaving that aside and confining ourselves to projects

on rivers, it is well-known that old-style planning in the former Soviet Union led to the

diversion of two rivers that were flowing into the Aral Sea, resulting in the virtual death of

that sea. That is now recognized as a great environmental disaster, perhaps the greatest

ever, and desperate attempts are being made to reverse it. With the ‘linking of rivers’ project

we may be headed for other unforeseen disasters and may discover this too late. A degree of

caution seems warranted before the Government embarks on this enterprise. (It may be

added that there is a move in some countries away from the past history of interference with

the natural flows of rivers towards a restoration of the original flows to some extent.)

JOURNAL , BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

7

Those who advocate caution are apt to be accused of timidity and exhorted to look at

China which has embarked on the massive Three Gorges Project. That is not necessarily a

good project; the disasters that it will bring will be seen in the future. The opposition to

Three Gorges in China is muted because dissent is not easy in that country. Those who are

envious of China’s ability to ‘get things done’ must reflect on how far they are prepared to

go in emulating that system.

Announcement in Advance of Examination Clearance

This is a ‘concept’ that consists of some twenty or thirty projects. For each project,

some small and some big, a proper feasibility study will have to be prepared as an

interdisciplinary exercise, fully internalizing economic, social, sociological, human,

environmental and other aspects ab initio. Thereafter, the projects will have to be examined

and evaluated, again in an inter-disciplinary manner, and cleared by the appropriate agencies.

Thorough Environmental Impact Assessments, comprehensive Cost-Benefit Analyses

covering direct and indirect financial, economic, environmental, ecological, social and human

costs and benefits (quantifying these wherever possible), qualitative assessments of non-

quantifiable considerations, and based on these, rigorous investment appraisals, will need

to be undertaken. We do not know what the outcome of that process will be: all projects may

pass the test; all may fail; or some may survive a stringent scrutiny while others may not. In

advance of that process, a project has been announced and expectations raised in the general

public. The presumption is that the project or projects will be found acceptable and cleared.

We fear that this may reduce the whole process of examination, evaluation and clearance to

a mere formality, a mockery. With the conclusions already presumed and announced at the

highest level, it seems difficult to believe that the governmental agencies concerned (the

CWC, the Technical Advisory Committee, the Ministry of Environment and Forests and its

Committees, the Task Force that has now been set up) will be able to undertake a serious

and objective examination. The pressure on them to be ‘positive’ will be very great.

Incidentally, we are told that NWDA has prepared feasibility studies for some five or

six links, and that these have been “ratified by engineers, sociologists and economists”. If

indeed there are feasibility studies of some of the proposed links, we would strongly urge

that they should be put into the public domain for engineers, geographers, environmentalists,

economists, agronomists, soil scientists, sociologists, social anthropologists, financial

analysts, and others outside the Government to examine and offer their comments. This

massive undertaking is too important a matter to be left entirely to the internal processes of

the Government.”

Such a massive environmental intervention will result in substantial human

displacement, and it is the poor, the invisible people who will harvest the Grapes of Wrath.

— J.C. DANIEL

8

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY 100(1), APR. 2003

JOURNAL

OF THE

BOMBAY NATURAL HISTORY SOCIETY

April 2003

Vol. 100

No. 1

BREEDING BEHAVIOUR OF THE GREATER ADJUTANT-STORK

LEPTOPTILOS DUBIUS IN ASSAM, INDIA1

( With one text-figure and two plates)

Hillaljyoti Singha2-3 4, Asad R. Rahmani24, Malcolm C. Coulter5 and Salim Javed2-6

Key words: Greater adjutant-stork, Leptoptilos dubius , breeding behaviour,

incubation period, fledgling

We studied the breeding behaviour of the greater adjutant-stork Leptoptilos dubius (Gmelin), the

rarest stork in the world at north Haibargaon — its traditional breeding colony in Nagaon

(26° 21' N, 92° 45' E), Assam, India — during 1995-1997. Focal nests of early (September-

October) and late (November-December) breeding storks in both the seasons, from nest building

to fledging of the last young, were observed from dawn to dusk. Development of young from

hatchling to fledgling and the associated parental behaviour were recorded in detail.

The incubation period was 35 days while the fledging period was 142 days. Parents left unguarded

chicks at the age of 4 weeks; at 6 weeks the chicks could defend themselves, and at 9 weeks they

attained juvenile stage. The overall breeding behaviour of the greater adjutant-stork was found to

resemble its congeneric the Marabou stork Leptoptilos crumeniferus.

Introduction

Of the 20 storks found in the world, the

greater adjutant-stork Leptoptilos dubius is

perhaps the most endangered. Earlier widely

'Accepted February, 2001

^Department of Wildlife Science, Aligarh Muslim University,

Aligarh 202 002, Uttar Pradesh, India.

'Present Address: Department of Zoology,

Birjhora Mahavidyalaya, Post Office and District Bongaigaon,

Pin 783 380, Assam, India.

Email: [email protected]

4 Present Address: Bombay Natural History Society,

S.B. Singh Road, Mumbai 400 023, Maharashtra, India.

Email: [email protected]

5P.O. Box 48, Chocorua, NH. 038 1 7, USA.

Email: [email protected]

distributed in Nepal, Bangladesh, Myanmar,

Thailand, Cambodia and South Vietnam (Baker

1929, Flemming et al. 1979, Ali and Ripley

1987, Hancock et al. 1992), this largest of Asian

storks has drastically reduced in number, and is

confined to the Brahmaputra Valley of Assam,

India (Saikia and Bhattacharjee 1989, Rahmani

et al. 1990) with a small breeding population of

100-150 birds in Cambodia (Mundkur et al

1995).

The greater adjutant-stork is a colonial

breeder. Very little information was available on

its breeding biology (Kahl 1966, 1970, 1971;

Baker 1935; Hume and Oates 1890; Saikia and

Bhattacharjee 1990) prior to our study. We

studied the breeding biology of the stork

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

9

BREEDING BEHA VIOUR OF GREA TER A DJUTANT-STORK

intensively in the north Haibargaon breeding

colony at Nagaon for two successive breeding

seasons from 1995-1997, right from its arrival

at the breeding colony, including the

development of the chick(s) to fledgling, up to

its departure. The results were compared with

those of other stork species.

Study Area

The study site, north Haibargaon, is a

traditional breeding site of the greater adjutant-

stork at Nagaon (26° 2T N, 92° 45' E). It is a

small, semi-urban town situated in middle

Assam, on the southern bank of the River

Brahmaputra. New buildings are being made

very near the nesting trees, which are on private

property, near human habitation. A few busy

public roads intersect the nesting colony. Behind

the houses, there are isolated patches of forest

and mixed plantations. Shrubs are present under

the tall bamboo Bambusa sp. and betelnut Areca

catechu dominates the area. Kolong a small river,

flows about 1 00 m from the colony.

The Brahmaputra Valley has four seasons:

winter (December-February), pre-monsoon

(March-May), monsoon (June-September) and

the retreating monsoon (October-November).

The average rainfall during monsoon is 286 cm.

Methods

Chronology: The greater adjutant-storks

aggregate at the colony in early September

(Saikia and Bhattacharjee 1996a). Eggs are laid

from September in Assam (Hancock et al. 1992).

Storks were monitored daily from the first week

of September, when they arrived at the site, till

the end of May when all the nests were vacated.

The adult storks on nests, and on nesting and

non-nesting trees were monitored twice (morning

and evening) daily.

Analysis: Monthly average of storks was

calculated from the data collected. For analysis,

data of only those days when both morning and

evening counts were taken was considered. Each

nesting tree and nest(s) on it were assigned a

code number to avoid counting error.

General Breeding Biology: Each year a

tall watchtower ( machan ) was built to observe

the colony. The height of the machan was equal

to or just above the nests. Care was taken not to

disturb the birds while building the machan.

Once the birds started incubating, they did not

desert the nest. They were observed till the focal

nests were either vacated or abandoned. A nest

was considered abandoned when the stork pair

stopped visiting before laying the eggs, or if the

eggs did not hatch and the pair left the nest, or if

all nestlings died and the parents stopped

attending the nest. A nest was considered vacated

when the last nestling of the clutch fledged.

During the first breeding season, from

November 1 , 1 995 to April 21,1 996, we observed

five breeding pairs from a 20.5 m high machan.

There were two trees with one nest each and one

with three nests. From the last week of November

to the first week of December 1995, three more

nests were built on a tree where there was only

one nest earlier. Out of the eight nests, five early

ones (categorised as early breeders) were

observed from the incubation stage, while the

later nests (categorised as late breeders) were

observed from the nest building stage. Since two

nests were abandoned before the eggs hatched,

and all the nestlings in one nest died, only five

nests were observed till the fledging of juveniles.

The average distance between the machan and

the nests was 13 m.

During the second breeding season, from

Novembers, 1996 to May 12, 1997, we observed

breeding pairs on three nests on three trees from

a 23 m high machan from incubation till the

juveniles fledged. The distances between the

machan and nests were 12 m, 30 m and 60 m.

The breeding behaviour of the focal pairs

was monitored continuously from dawn to dusk,

0500 to 1700 hrs, six days a week. However, the

10

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

BREEDING BEHA VIOUR OF GREATER ADJUTANT-STORK

duration of observation decreased to 1 1 hrs (0545

to 1645 hrs) a day during the shorter days of

winter. It was assumed that the individuals seen

in the late evening stayed through the night, as

they were seen there again the next morning.

Individuals of the focal pairs were

identified by their natural markings, facial and

bill patterns and individual variations. Gender

was determined by the copulation position. We

found that the males were slightly larger than

the females, which also helped to identify a male

and a female of a pair (see also Kahl 1972a). We

sketched facial patterns and noted the

identification characters on ‘face cards’ (Coulter

and Bryan Jr. 1988, Coulter 1989). The sketches

were redrawn as the facial pattern of the stork

changed.

We observed the breeding behaviour of

other storks in the colony as far as possible. The

morphological and behavioural changes from

chick to fledgling stage were recorded. A chick

was distinguished from a juvenile when blackish-

brown feathers replaced its snowy white down

feathers.

Incubation and Fledging Period:

Incubation was estimated as the period from the

initiation of incubation till the hatching of the

first chick. The initiation of incubation was

considered when, soon after nest building, the

parents began behaving as if they were brooding

eggs (Coulter 1989). It was difficult to know the

exact date of hatching despite daily monitoring.

The hatching at each nest was assumed on

hearing the first call of the chick, or finding fresh

broken egg shells below the tree, or observing

the parent’s posture and behaviour in the nest.

In the focal nest, just after hatching, the parent

stork would stand with its legs apart, point the

bill towards the floor of the nest (to the new

hatchling) and regurgitate or re-ingest food.

Otherwise, when the chick was first seen, its

probable age was subtracted to get the hatching

date ( Kahl 1966 and Pomeroy 1978a). In three

cases where incubation period exceeded 55 days,

it was considered as probable re-laying of egg(s)

and such nests were excluded from the analyses.

The mean incubation period was calculated from

five nests of both seasons, where egg laying and

hatching was observed from the machan.

The fledging period was considered as the

duration from the hatching to fledging of the

juvenile, assuming that the same juvenile was

the first chick. Before finally leaving the nest,

juvenile(s) make exploratory flights and remain

away from the nest for some time, and could be

missed during a daily census. It was logistically

difficult to see the first flight of all juveniles in

the whole colony. Therefore, if a juvenile was

not seen continuously for a week, it was regarded

as fledged, and the fledging date recorded as

the date it was last seen. We recorded the date

of first flight in 12 nests from both the seasons,

to calculate the fledging period.

Results

Chronology: Data were collected over

seven and nine months (November 1995 through

May 1996 and September 1996 through May

1997) in the first and second breeding seasons,

respectively. Storks were found to arrive at the

breeding site from the first week of September.

In both the seasons, the number of storks

increased to a peak in the early part of the season

and gradually declined to almost zero in May. In

the first season (1995-96), the highest

aggregation of storks was found in December,

while in the second breeding season (1996-97),

it was in October (Figs la & b). But the overall

population trend in the study area was similar in

both the breeding seasons. In all the months, in

both breeding seasons, fewer number of storks

were counted in the morning than in the evening.

General Breeding Biology

Pre-Chick Hatching Period: (a) External

morphological changes: At the onset of the

1 1

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

BREEDING BEHA VIOUR OF GREATER ADJUTANT-STORK

60

50

o

«*■»

CO

4=1

° 30

Ut

X

g

3

£ 20

10

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NOV DEC JAN FEB MAR APR MAY

Months

Fig. la: Population trend of greater adjutant-stork in North Haibargaon Nesting Colony (1995-96)

40

30

«/>

M

O

+-»

CO

4-*

° 20

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SEP OCT NOV DEC JAN FEB MAR APR MAY

Months

Fig. lb: Population trend of greater adjutant-stork in North Haibargaon Nesting Colony (1996-97)

12

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY / 00(1). APR 2003

BREEDING BEHA VIOUR OF GREATER ADJUTANT-STORK

breeding season, the stork underwent

conspicuous morphological changes to become

a brilliantly coloured bird (Plate 1, Fig. 1). The

pale whitish wing band of the greater secondary

coverts of the non-breeding season became bright

silvery white. The upper parts of the body

including back, wing, tail and mantle turned to

bluish-grey from slaty grey or blackish-grey. The

under-tail coverts, which were white at their base

and dark smoky grey at the tip, became fluffier.

The head and nape became bright red. Black

pigment appeared on the pale red part between

the head and forehead. The skin of the face and

forehead appeared rough and encrusted with dark

spots. The base of the bill, just below the dark

skin, turned pinkish. The neck turned bright

yellow with a pinkish tinge. The dorsal air sac

became bright red, encircled by a ruff of white

feathers. The gular pouch appeared bright red.

We found that the female was brighter than

the male. All breeding pairs wore the breeding

plumage except one dull coloured male. There

were some storks in the colony which, despite

being brilliantly coloured, were smaller than an

adult in size, and had a few erect ‘hairs’ on their

head, and black ‘hairs’ hanging from the tip of

the gular pouch. They were most probably

younger storks or first time breeders.

As breeding progressed, the breeding pairs

gradually regained their non-breeding colour. By

mid January, the upper part turned ashy grey;

head, neck and forehead became paler from

bright red and black respectively. The silvery

white wing band also faded. In April, they looked

like non-breeding storks with blackish upper

parts and a dirty wing band.

(b) Flocking: Breeding storks flocked in

the beginning of the breeding season. There were

some aggressive interactions, including chasing,

fighting, squealing, bill clapping and frequent

flying from branch to branch and tree to tree.

Flocking occurred due to competition for pair

formation and nesting sites. One stork would

follow another, and within a short time many

storks would gather on a tree. At this time, a

stork perched on a branch would stretch its

wings, with a forward curve, bend its neck down

and bring the open bill between its feet. At the

same time, the tail would be cocked and the stork

would make a long squeal ‘qui-e-e-i’, similar to

the call of the black kite Milvus migrans , or a

loud booming nasal sound ‘we-i-nh’. The storks

were also found to make loud and deep ‘woom’

sounds like the “lowing of a cow” (Hume and

Oates 1890). The female was more vocal than

the male, generally squealing and making nasal

‘we-i-nh’ noises, which gradually became faint

later in the breeding season. This high-pitched

vocalisation would be accompanied by occasional

bill clapping with the bill pointed upward. A dull

coloured stork and some subadult storks also

visited the colony. However, they did not take

part in the flocking.

Flocking would occur on one or two trees

in the colony at the same time. Each stork made

a small territory of its own and refused entry to

other storks, whose approach resulted in

ferocious fights. Sometimes flocking (crowding)

occurred continuously for ten to fifteen days on

a particular tree. These activities started in the

morning, gradually declined at noon and again

increased towards evening. While some storks

formed pairs and selected nesting sites, the

remaining moved to other trees for flocking.

These activities increased sharply to a peak

in the beginning of the breeding season, and

gradually ended in the latter part of the breeding

season. While many storks settled down for

nesting, some that were unable to find a partner

still crowded around to disperse the pairs and

invade the nests. We termed these as ‘floating

storks’, most of which were probably first time

breeders. Crowding was seen as late as the last

week of January.

(c) Pair formation and courtship: During

the process of crowding, a male stork would

create a territory on a potential nest site, and

chase other storks that came near him. He

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

13

BREEDING BEHA VIOUR OF GREATER ADJUTANT-STORK

announced his territory by frequent, loud bill

clattering, pointing the bill upward, downward,

or horizontally, and prodded the air with frontal

arched body and half folded wings towards its

breast. When he accepted a female stork, both

perched closely to form a pair. A third stork,

usually a female, was often found trying to break

the pair bond. The female of the pair was more

aggressive than the male in chasing away her

rival. In two cases in the first breeding season

1995-96, we found that the third stork

temporarily replaced the female of the pair, but

was finally driven away. The male easily accepted

the new partner.

In some cases, the new pair was found to

perch close together continuously for two to three

days, without foraging, before building the nest.

We saw the pair take initiative in the courtship

display, which varied only slightly among

different pairs. A male would pluck a fresh twig

from a nearby branch and put it near the feet of

the female, or drop the twig after holding it for a

few minutes. He would touch and gently grasp

one of the female’s tarsi with his bill. Sometimes

the female also touched her own legs. She would

withdraw her foot whenever a male grasped it.

Once, a male pulled at one of her primaries. He

gently clasped her bill, touched her breast in a

preening gesture and gradually came closer to

her. During this process they leapt to another

branch or made a short flight to another tree with

the second partner following behind.

When a female approached a male, she

would lower her head to touch the male’s breast

with both her bill and head. He would cross his

neck over hers and she adopted a submissive

posture. Typical ‘swaying twig-grasping’ and

‘up-down’ displays (Kahl 1971, 1972a) were

shown by both; however, the former was mostly

shown by the male. The stork perched close to

its partner bent its neck downward, pointing the

bill down about 45°, and oscillated gently from

side to side four to five times at half an oscillation

per second. Sometimes, at the extreme ends of

an oscillation, it would pull or touch a twig and

give a mild jerk. Both the partners perched side

by side, sometimes preened themselves and

showed an up-down display simultaneously or

one by one. The bill was then sharply brought

down almost touching the breast, or abdomen or

feet with a sudden loud nasal ‘we-i-nh’

immediately followed by throwing it upwards

pointing vertically towards the sky. Thus, they

clattered their bills, stopping only after the bill

was again brought down below the horizontal

position. The clattering ceased before the bill was

brought to a horizontal position in brief bill-

clattering sequences.

(d) Mating and nesting: Usually, the

courtship display was followed by mating which

occurred on the nest-branch or in the nest. Only

twice was mating seen outside the nesting tree.

Just before mating, in most cases, both the

partners stood side by side in close contact. The

male lifted one of his legs onto the back of the

female and mounted (Plate 1, Fig. 2). He either

mounted from the side, or from the front or rear,

standing on her shoulder for a few seconds and

then positioning himself for copulation. As soon

as the male mounted, the female bent her ‘knees’

in a submissive posture, both the wings spread

for balancing. During copulation, the male

flapped his wings vigorously and sometimes

snapped his bill frequently. The bill clattering

usually started as soon as he copulated, at times

simultaneously with mounting. On a few

occasions, the sound of the bill clapping was not

heard. During copulation, the female lowered her

open bill 45°, swayed it from side to side and

made a moaning sound. The male usually clasped

the female’s bill and clattered against her bill.

After copulation, he stood on her shoulder for

some time. Mating was sometimes followed by

preening.

The whole process of mating ranged from

1 1 to 68 seconds (average 25.56 ±10.38 sec, n =

52). Mating occurred 1-6 times a day, between

0545 and 1635 hrs. Out of 141 matings observed,

14

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

BREEDING BEHAVIOUR OF GREATER ADJUTANT-STORK

Singha, Hillaljyoti et al.\ Leptoptilos dubius

Plate 1

Figs 1-4: Greater adjutant-stork Leptoptilos dubius : 1. Adults in breeding plumage, 2. Adult male mounting

on a female, 3. Parent tending one week old chick, 4. Adult feeding two week old chicks

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

15

BREEDING BEHAVIOUR OF GREATER ADJUTANT-STORK

Singha, Hillaljyoti et al.\ Leptoptilos dubius

Plate 2

7

Figs 5-7: Greater adjutant-stork Leptoptilos dubius: 5. Parent with four week old chick, 6. Parent tending

six week old chick, 7. Nine week old juvenile

16

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

BREEDING BEHA VIOUR OF GREATER ADJUTANT-STORK

81 (57.45%) occurred in the forenoon, and

59 (41.84%) in the afternoon. One was seen at

mid-day. Mating was more frequent during nest

building and early incubation. However, late

mating was seen in four pairs whose first chick

was 26, 32, 43 and 45 days old, respectively.

Simultaneously with mating, nest

construction began. The male would place a

green leafy twig on a flat horizontal branch, with

or without an erect limb. The female stepped on

it to hold it down, and soon the base was formed

by piling one twig upon another. During the

initial stages of nest construction, the female

stood guard at the nest, while the male brought

most of the nest material from the same or other

trees in the colony. The stork did not go beyond

200 m to bring nest materials. Usually, mating

occurred after frequent nest material trips by the

male. During nest building, the female became

more vocal, making a loud booming sound,

accompanied by bill snapping. The male

responded by clattering his bill when he reached

the nest. Both sexes arranged the nest material.

They would remove leaves from the nest and drop

them as if cleaning the nest. In the early stages

of nest building, the male generally roosted with

his partner in the nest or near the nest. The nest

construction took 2 to 4 days. Soon after that,

the female spent more time on the nest, until she

laid the eggs. Then the male took care of the

nest, relieving the female for longer periods.

During nest construction, the male and

female never left the nest together, but on two

occasions the storks were absent from the nest

(11 and 36 minutes) to collect nest material.

Incubation: The eggs were laid

asynchronously and the incubation started as

soon as the first egg was laid. The newly laid

eggs were chalky white, with a green and blue

tinge, which became soiled as the incubation

proceeded. Both male and female incubated.

During incubation (34.6 ±2.70 s.d. days, n = 5),

the incubating stork preens, flaps and stretches

its wings, it rearranges and repairs the nest,

rotates the egg(s) and protects the nest from other

storks and birds. When the bird stands up after a

long bout of incubation, it shakes its body,

stretches its neck forward with an open bill

slightly downwards and changes its orientation

for the next phase of incubation. It rotates the

eggs to different positions by grasping them with

the bill or shovelling them towards its feet with

the tip of the beak. It then tilts the nest slightly

with its feet, and the eggs roll back to the

depression in the middle of the nest.

Post-hatching period: a) Growth of chicks

and associated behaviour of parents: The growth

rate of the chick varied in different nests. A

general pattern of growth and behaviour of the

chick is described below.

The chicks hatch at intervals of one or two

days. The newly born chick is about 10 cm tall,

and has a pale yellow, slightly curved down beak

and a large blackish head, and neck with yellow

patches. Dorsally, the body has bluish-black down

feathers, and pink underparts. The eyes are large

and black; the swollen eyelids are blackish, while

the forehead is sky blue. The chick grows fast

and becomes doubie its size within a week, with

white down feathers almost covering the back.

During the first week of their life, the chicks

spend most of the time sleeping.

When a chick hatched, the parents stood

with legs apart and tended the baby frequently.

The behaviour and posture of the parent indicated

egg hatching (Plate 1, Fig. 3). The chick made

feeble sounds, which became louder with time,

into chittering. Both parents fed the young. The

parents regurgitated food on the floor of the nest

and the chicks picked it up. Soon after hatching,

the parents started regurgitating black, granular

half-digested food matter, so that the hatchlings

could feed easily. The feeding frequency by

regurgitation was higher in the early stages.

Initially, the chicks could not consume all the

food brought to them; the excess was re-ingested

by the parent, and regurgitated iater. While re-

ingesting, the parent pressed large portions of

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17

BREEDING BE HA VIOUR OF GREA TER A DJUTANT-STORK

food between its mandibles to soften and crush

it, so that the chicks could devour it easily. We

found that although the parents would not feed

the young bill to bill, it would help by holding

the food such that it could be easily swallowed.

The chick would jerk the food, toss its head

forward and backward quickly and swallow the

food. The chicks were found to be voracious

feeders right from hatching. Even a day old chick

tried to swallow a frog, much larger than its head!

In one case, a three-day old chick gulped a fish

that was larger than its body length; for some

time the tail of the fish protruded outside its bill.

At two weeks (Plate 1, Fig. 4), the white

down feathers completely covered the body of

the chick, however, the underparts were still

pinkish. The yellow patches on the head almost

disappeared. The eyelids became whitish. The

forehead or shield turned white with black spots.

Dark black pigment spreads over the cheek. The

ventral side of the throat was covered with dark

black pigment in longitudinal bands. Dorsally,

the neck was greyish and the shoulders black.

Erect grey and white ‘hairs’ grow on the head

and neck. The ill-developed sky-blue pouch could

be distinguished. Beneath the wings and near

the flanks, black feathers developed. The legs

and toes became pinkish in colour.

Generally the parents relieved each other

after more than 24 hrs. The reliever brought food

for the chick(s) once a day. For about two weeks,

most of the parents’ time was spent in brooding

the chicks and after that, in standing guard.

However, in the morning and evening, and

during rain they brooded and sometimes shaded

the youngs. The parents also provided shade to

the chick(s) by stretching their wings and

standing against the sun.

The 3-week old chick is covered all over

with snow white foamy down feathers, except

ventrally. The bill is pale yellow with black marks

at the base. Black ‘hairs’ hang from the tip of

the pouch and on the shoulder. Pupils black, iris

brown. Legs pinkish-white.

The 4-week old chick is about 30 cm in

height. The shield is still white with black spots.

The grey and black crest of thick ‘hairs’ grows

longer. Down the throat, and ventrally,

longitudinal black patches are prominent. Black

primaries and secondaries start to grow. Black

tail feathers also appear.

At 4 weeks, a chick can stand on its feet

and flap its wings 4-5 times at a stretch (Plate 2,

Fig. 5). The wing span is approximately 60 cm.

The chick walks in the nest. The parents also

gradually start leaving the nest, initially for short

durations only. When the parents return with

food, the young ones perform typical begging

displays. They drop down to their tarsi opposite

each other (if more than one young), wings half

folded, slightly lifted upward, body bent forward,

tail cocked and toss their heads up and down

rhythmically with open bill and nasal ‘honk-

honk’ sound.

In the 5-week old chick, the primary

feathers of the wings start developing. Initially

they have a black tip and bluish rachis. The

primaries at the tip of the wings are now larger,

about one and half inches. The tail feathers

become more prominent. The pinkish white legs

turn whitish. The longitudinal black patches of

the throat become concentrated only on the

pouch. The ear opening is surrounded by a white

mark. At this stage, a chick tries to leap in the

air, frequently flapping its wings. It defecates over

the edge of the nest and swallows food very fast.

A 6-week old chick is fully covered by thick

down, which has changed from snowy white to

dirty white (Plate 2, Fig. 6). The black spots on

the shield are almost gone. The sky blue throat

pouch, with pink tinge at the base, and sparse

black pigments, becomes elongated. The black

‘hairs’ hanging from the pouch become longer.

Black feathers on the shoulder around either side

of the base of the neck turn brownish and grow

to meet at the breast, forming a necklace-like

band. The wings develop four layers of feathers:

the lowermost black primaries are concealed by

18

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BREEDING BEHA VI OUR OF GREATER A DJUTANT-STORK

a broad brownish band of greater secondary

coverts, and two more black layers above the

second layer. Twelve broom-shaped fan feathers

with blue rachises grow at the tail, black horse-

shoe shaped flat feathers at the tip, followed by

thin grey ‘hairs’ hanging from it.

At this stage, the parents even stay away

from the nest at night as the chick is able to

defend itself. It would even threaten a crow or a

kite hovering overhead by poking its head

towards the intruder, with a harsh ‘khll-o-ck’

sound. When other storks came near the nest, it

sat down on its tarsi, faced the intruder with half

folded wings, and tossed its head up and down,

emitting a nasal ‘khll-i-ck khll-i-ck’ sound

rhythmically with movement of the head till the

intruder flew off. At this stage, the chicks also

clean the nest like adults.

By 7 weeks, another row of black feathers

developed just above the fourth layer on the

wings. Two longitudinal rows of black feathers

were growing on either side of the midline of

the mantle. When the wings were folded, three-

fourths of the upper part was covered by black

feathers. At this stage, the parents came to the

nest only to feed the chick(s). The chick flapped

its wings more frequently, 10 to 12 times at a

stretch.

At 8 weeks, six rows of black feathers

appeared on the wing. The tail fan became

elongated and expanded. While flapping the

wings, the chick tried to lift itself up in the air.

Throughout the fledgling period, the

parents occasionally nursed the chick. Up to

3 weeks, they would gently touch the head and

body of the chick, but later they also preened the

chick.

(b) Growth of Juvenile and associated

parental behaviour: At nine weeks, the chick

attained juvenile stage (Plate 2, Fig. 7). It almost

equalled its mother’s height. The upper parts

became completely black. The broad brown layer

of the secondary coverts became more distinct.

The shield was white with very few black spots.

The bill was pale yellow. The ear opening

surrounded by white mark became more

prominent. The pupil was black and the iris

brown. Near the eyes and on the cheek, black

spots were sporadically present. The thick crest

of black and grey ‘hairs’ on the crown and neck

were longer, and so were the ‘hairs’ at the tip of

the whitish pouch. The legs became creamish-

white.

The juvenile’s food was supplemented with

pieces of meat and intestinal parts of mammals

in addition to fish, amphibians, reptiles and birds.

Gradually, it can leap straight upward 1-2 in in

the air with vigorous flapping of wings. This

leaping is more frequent when the wind blows.

At about four months, the juvenile flies from one

branch to another and then to the other trees in

the colony. It can chase other intruders from its

nest. Sometimes it flies outside the colony and

comes back to the nest to be fed by the parents. It

can also be seen on other nests in search of food

and nest material. The parents feed the juvenile

till it fledges. The fledging period was about 142

days (141.94 ±22.45 days, n=35). The first flight

of the juvenile was seen at 126.25 ±9.35 s.d. days

(n=12). The fledging period ranged from 96 to

173 days in the first breeding season, and 1 10 to

197 days in the second breeding season in

different nests. The juvenile fledeed

asynchronously; not necessarily the older sibling

leaving earlier.

Other Breeding Behaviour: Greeting

display: Greeting display between a pair

commences soon after pair formation. It gradually

declines by the time the chicks reach juvenile

stage, because the parents hardly come to the nest

at the same time. Among the greeting display,

the up-down display and bill clattering were

common. Usually when a partner arrives at the

nest, the other partner greets it by upward bill

clattering. Then both demonstrate an up-down

display. Each stork could recognise its partner

from a long distance; even about 100 m. When

nest material was brought by the male, he

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19

BREEDING BEHA VJOUR OF GREATER ADJUTANT-STORK

generally uttered a low pitched ‘kis-kis’ sound

with one second pause after each ‘kis\ The

female brought nest material with nasal ‘we-i-

nh we-i-nh’ sound. In the early part of the

breeding season, a female greets the male with

her long typical squeal, swaying the bill from

side to side. When a female arrives on the nest,

the male sways his bill from side to side, bowing

down his head with a typical ‘kis-kis’ sound.

When a male reaches the nest, the female sways

her down-pointed bill from side to side, and

places the nest sticks in accordance with the

movement of the bill. Downward short

duration bill clattering was also observed to greet

each other. Sometimes there was no greeting

display.

Aggression: Agonistic behaviour was seen

more in the beginning of the breeding season.

Aggressiveness also varied according to the

individual and the situation. Generally, the

female appeared to be more aggressive. Storks

were found to chase other storks, which

approached their nest, sometimes leaving behind

an unprotected nest with egg(s) or chick(s). A

stork could chase another stork up to 100 m,

flying at it with a stretched neck and loudly

snapping bill. Loud bill clattering was not only

a greeting display, but also a threat to the other

storks. Sometimes a breeding stork was found to

tolerate the presence of other storks and even

crows very near its nest.

In the early part of the breeding season,

during nest building, and even during incubation,

some storks try to invade other nests for

occupation or to replace a partner from an

established pair. A case of nest invasion occurred

in the first breeding season when one pair

occupied the nest of another pair. Once a stork

was also found to snatch food from another

stork’s nest. While fighting they poked their bills

at each other’s head in quick succession. Post

flying juveniles were also found to be aggressive

towards other adult storks that tried to enter their

nest.

Nest arrangement: Storks repaired and

cleaned their nest throughout _the breeding

period, but this activity progressively decreased

as the young grew up. The storks cleaned the

nest by removing egg shells and decomposed old

leaves from the interior of the nest. They

frequently picked up leaves by inserting one third

of the bill into the huge nest and threw rotten

leaves outside the nest. They grasped sticks and

placed them, pulling and pushing them all along

the nest rim and also at the bottom. Standing on

one leg at the centre of the nest, the bird slowly

placed the sticks along the periphery, so that a

shallow depression developed at the centre of the

nest. Sometimes this would continue for more

than half an hour.

Nest material stealing: On finding an

unguarded or vacated nest, storks stole nest

materials. Even a five-month old juvenile was

seen stealing sticks from empty nests. Sometimes

they would rob nest material in the presence of

its owner.

Father-offspring mating: What appeared

to be father-offspring mating was seen in one

nest in both the breeding seasons. The adult male

suddenly placed one leg on the back of one of

his young and mounted it. He stood on the back

for 12 to 20 seconds and came down. Sometimes

he appeared to copulate with the young, with or

without mild bill clattering. The young uttered

harsh chittering. In the first breeding season, this

was observed ten times when the young were

9 to 1 1 weeks old. In the second breeding season,

it was observed twice when the young ones were

5 and 9 weeks old.

Discussion

Chronology: The greater adjutant-storks

arrive at the North Haibargaon breeding colony

just after the monsoon, when the dry season

begins. In both the breeding seasons, the time of

arrival was identical suggesting that the storks

follow a definite time frame for breeding. The

20

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BREEDING BEHA VIOUR OF GREATER ADJUTANT-STORK

nesting period synchronises with the reducing

water level in the lakes and ponds (Saikia and

Bhattacharjee 1996a, Bhattacharjee and Saikia

1 996). All other breeding storks in India, painted

stork Mycteria leucocephala, white-necked stork

Ciconia episcopus , black-necked stork

Ephippiorhynchus asiaticus, Asian openbill stork

Anastomus oscitans and lesser adjutant-stork

Leptoptilos javanicus breed just after monsoon

(Ali 1996). In Assam, from September onwards

water level gradually recedes with the decreasing

rainfall. This period also coincides with the

increase in availability of prey species, as many

species of fish and frog breed during monsoon

in the inundated low-lying areas. When water

level drops, they concentrate in drying pools and

puddles and become easy prey for the stork.

Most bird species breed around the time

when food supplies are readily available

(Thompson 1950). In the congeneric Marabou

stork Leptoptilos crumeniferus , breeding usually

begins in dry season and ends in rain (Brown

et al. 1982). Pomeroy (1978b) also states the

possibility of an intrinsic (circannual) rhythm

for nesting period in the Marabou stork. The

American wood stork Mycteria americana

initiates nesting when water level goes down

(Kahl 1964). Seasonal rains strongly influences

the beginning of nesting in the Maguari stork

Ciconia maguari (Thomas 1985). It seems that

food supply and seasonal change with shorter

days correspond to initiate breeding of the greater

adjutant-stork in North Haibargaon breeding

colony.

The population trend throughout the

breeding season in both years of our study

suggests that as the breeding season progresses,

the greater adjutant-storks start leaving the

colony. The storks that did not build nests leave

the colony; the parent storks also spend less time

in the colony when the young grow up. In the

second breeding season it was seen that the

number of storks reached a peak in October

(Fig. lb) when almost all breeding storks had

arrived in the colony. In both the seasons there

was a small population peak in December

(Figs la & b). This could be due to the late

breeders who returned to the colony. A similar

second small peak was seen in a painted stork

colony by Urfi (1993), which he speculated could

be due to a second breeding attempt or prolonged

breeding effort. In Maguari stork also, late-

arriving individuals begin to build nests well after

the beginning of the breeding season (Thomas

1986).

The reason for smaller numbers of storks

in the morning count is probably because some

storks leave very early for foraging, much before

sunrise. In the evening, all the storks returned to

roost, so more birds were seen.

General Breeding Biology

External morphological changes: The

external morphological changes in the greater

adjutant-stork during the breeding season were

similar to the observation of Saikia and

Bhattacharjee (1996a) and, Bhattacharjee and

Saikia (1996). Similar changes occur in the

Marabou stork also (Brown et al. 1982). Pomeroy

(1977b) found that males of the Marabou stork

were considerably larger than females, which we

found in greater adjutant-stork also. Kahl (1972a)

has reported this earlier. In painted stork (Desai

el al. 1977) and American wood stork Mycteria

americana (Kahl 1962), males are larger than

females. The brighter coloured female approaches

the male to initiate courtship. Competition for pair

formation was between females for the male who

had selected a potential nest site.

Flocking and aggressive behaviour: Lack

(1968) also observed that in some colonial

species, the sub-adults also come to the breeding

sites, form pairs, occupy nesting sites and build

nests, but do not proceed further. We found some

sub-adults and some probable fresh adults

arriving in the colony. Bhattacharjee and Saikia

(1996) also observed that both sub-adults and

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21

BREEDING BEHA VIOUR OF GREATER A DJUTANT-STORK

non-breeding greater adjutant-stork also came

to the nesting colony, but Pomeroy (1977a)

observed that it does not necessarily follow that

all storks in breeding plumage are sexually

mature. The minimum age at first breeding is

4 years in Marabou stork (Pomeroy 1977b), and

3 years in male and 4 years in female in Maguari

stork (Thomas 1984). According to

Bhattacharjee and Saikia (1996), the greater and

lesser adjutant-stork become sexually mature at

3-4 years. Lack (1968) has explained the arrival

of the sub-adults at the breeding colony as the

preparatory year to learn the best feeding areas

around their nest without the strain of finding

food for a brood.

Bhattacharjee and Saikia (1996) reported

rigorous vocalization and 14 to 16 storks flying

together from tree to tree in the early part of the

breeding (September), what we have called

‘flocking1. Coulter (1989) has termed the

aggressive interactions in American wood stork

as ‘mobbings’ and Kahl (1972b) has referred to

them as “bachelor parties”. Although aggression

was always associated with crowding, fight was

not between two storks; instead a flock of five to

fifteen greater adjutant-storks took part together

in this activity. The aggressive interactions of

‘mobbings’ in American wood storks were

moderately frequent in the early part of the

breeding season, and rose to a peak in the

following weeks correlating with an increase in

nest building, and declined gradually (Coulter

1989). We observed the same in the greater

adjutant-stork.

We observed that in greater adjutant-storks

aggressiveness varied individually, and female

storks seemed to be more aggressive. While

Thomas (1986) noted that attacking Maguari

storks were mostly males. Though female

American wood storks are generally less able to

defend themselves (Coulter 1989), we found that

greater adjutant-stork females even invaded other

nests. As Coulter (1989) recorded nest invasion

in American wood stork, we found a similar case

in greater adjutant-storks too. The replacement

of one incubating female American wood stork

by another pair and throwing out the former’s

egg was observed in greater adjutant-storks also,

but in the case of greater adjutant-storks the

aggressor was a female. Moreover, while

guarding a nest with eggs or chicks, a female

sometimes chased away others near the nest,

leaving the nest unguarded. Such behaviour was

not seen in American wood storks by Coulter

(1989) who found that unguarded chicks were

thrown out by neighbouring storks. Similarly, in

greater adjutant-storks, while pulling nest sticks

from an unattended nest, the “robber” stork

attacked the defending chicks, of which, two

3-week old chicks fell out.

The ‘aerial clattering threat’ described by

Kahl (1972a) was also observed in the greater

adjutant-stork. According to him, lesser adjutant

remains silent while chasing another stork in the

air, but Marabou clatters loudly as the opponent

is approached closely. Similar behaviour was

observed in the greater adjutant-stork.

The other hostile display described by Kahl

(1972a) such as snap display, pre-flight snap,

erect-gape and anxiety stretch were noticed in

the greater adjutant-stork. However, forward

poking, which is similar to ‘forward display’

behaviour in herons (Meyerriecks 1960;

Tomlinson 1976), was not mentioned by him. In

this behaviour, two greater adjutant-storks stand

erect on their nest or branch face to face and

poke at each other, sometimes with a bill

snapping sound, frequently but hardly any

physical contact. Sometimes a greater adjutant-

stork was also seen shooting its bill sharply in

horizontal direction in the air when the opponent

was at a distance.

Pair-formation, courtship and greeting

display, mating and nesting; In many aspects

of morphology and behaviour, the greater

adjutant resembles its African cousin, the

Marabou stork (Kahl 1974), but it is closer to

the lesser adjutant (Kahl 1 970). We saw the same

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BREEDING BEHA VI OUR OF GREATER ADJUTANT-STORK

‘swaying twig-grasping’ as described by Kahl

(1972a) for the greater adjutant and Marabou

stork. For the first time we observed side to side

swaying of the bill by a male greater adjutant to

greet a female partner at her arrival to the nest

with “kis-kis” sound. This behaviour can be

compared with the ‘advertising sway’ of the

Asian openbill stork described by Kahl (1971).

The differences are; (a) greater adjutant-stork did

not lift its feet with each oscillation; and

(b) greater adjutant-stork makes a typical sound.

The female greater adjutant also greeted the male

in the nest by swaying downward pointing bill

and arranging nest sticks. Similar ‘twig passing

display’ is seen in the great white egret Egretta

alba (Tomlinson 1976), where the female greets

the male on his arrival to the nest by ‘stretch

display’.

The most common greeting display in the

greater adjutant-stork is the up-down display that

Kahl (1971) has seen in all except the Saddlebill

stork. He has mentioned that all the three species

of Leptoptilos give frequent up-downs whenever

a member of a pair returns to the nest. But we

found that the greater adjutant did not give an

‘up-down’ display every time its partner arrived

at the nest.

Kahl (1970, 1971, 1972a and 1974)

emphasised on the difference between Marabou

storks and both the adjutants in up-down display.

In the former, the bill is first thrown upwards

and vocalizations are made, then the bill is

pointed downwards and clattered. In the latter

the bill is directed upward during both

vocalizations and clattering. We also observed

the upward bill clattering, but would like to stress

that the clattering did not always cease before

the bill reached horizontal position; sometimes

clattering stopped only after the bill pointed

downwards. On some occasions we observed one

member of the pair greeting the other by short

downward bill clatterings. Vocalization of the

lesser adjutant is hoarser and more rasping than

the greater adjutant (Kahl 1970).

There is no literature regarding the mating

behaviour of greater adjutant-storks. However,

it was found that the loud copulation clattering

of greater adjutant-storks resembles that of other

storks, except Asian openbill storks where the

male does not clatter his mandibles loudly (Kahl

1970). The balancing posture of the female

during copulation is similar to that of other storks

except the white stork where the female does not

spread her wings wide (Kahl 1971) and the Jabiru

stork where the female usually opens her wings

fully at first and then closes them partially after

the male is in position (Kahl 1973). The

occasional post copulation preening in greater

adjutant-storks is also found in other storks, e.g.

painted storks (Desai et a/. 1977) and Maguari

stork (Thomas 1986). The copulation duration

of greater adjutant-storks (25.56 ±10.38 sec) is

found to be the longest among storks: 10 sec in

painted stork (Desai et al. 1977), 8.77 ±1.15 sec

in Maguari stork (Thomas 1986), 24 sec in black-

necked stork and 15 sec in Jabiru stork (Kahl

1973). Late mating, as we found in greater

adjutant-storks, is also reported in painted storks

where copulation occurred even after the fledging

of the young (Desai et al. 1977). However,

attempted father-offspring mating behaviour is

not reported in other storks. It was, however, not

known if the young was a male or female.

Development - egg to fledgling: Our

description of freshly laid eggs of greater

adjutant-storks is similar to what was described

by Hume and Oates (1890). The asynchronous

laying of eggs is also reported by Saikia and

Bhattacharjee (1996a). Lack (1968) states that

in Ciconiiformes, the successive eggs in a clutch

are laid two or more days apart, incubation starts

with the first egg, and the young hatch one or

more days apart.

The development of the chick to fledgling

stage and associated parental behaviour are more

or less similar with other storks. The stork

nestlings spend most of their time between meals

sleeping and thereby reduce energy demands

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BREEDING BEHAVIOUR OF GREATER ADJUTANT-STORK

(Kahl 1962). The rapid early growth pattern

exhibited by altricial birds, aids to survival in

several ways. A four- week old greater adjutant

tried to leap in the air with flapping wings. The

Marabou chick can stand and flap wings in

17 days (Brown et al. 1982) and Maguari chick

can stand in 22 days (Thomas 1984). The

postures, movements, and vocalizations in the

begging display and nesting defence display of

both Adjutant species were quite similar to the

Marabou stork (Kahl 1972a). However, in

solitary breeding storks, vocalization of the

chicks are not as loud as those in colonial species

(Kahl 1973).

Incubation and Fledging Period:

Pomeroy (1978a) estimated the age of the young

“ fairly accurately” from their appearance to get

the date of hatching, but our observations on

different nests indicated that it would definitely

vary. We found that individual chick growth was

quite varied, i.e., a particular stage could not be

always assigned to a particular age. Some chicks

grew faster than the others.

The incubation period of the greater

adjutant is around 35 days as observed in the

five focal nests. Except for the five nests,

prolonged incubation period exceeding 35 days

was noticed in some other nests. In one case,

even when the first chick had grown to 30 days,

an egg was still being incubated. Finally, it was

dropped from the nest. Prolonged incubation has

been reported in a number of bird species (Skiitch

1962, Afik and Ward 1989). Drent (1975)

suggested that prolonged incubation was a

functional response caused by the inherent

variability in incubation period. Marks (1983)

thinks that prolonged incubation behaviour is

related to the time interval in which the entire

clutch would normally hatch. This does not seem

to be the case in greater adjutants whose clutch

size is small and eggs hatch simultaneously.

Prolonged incubation is also reported in painted

stork (Desai et al. 977). Pomeroy ( 1 978a) reports

that one pair of Marabous’ incubated for

1 50 days, and eventually one young fledged from

that nest. He regarded it as re-laying of eggs in

the same nest, which he found was 8% in

Marabou stork. We also found 7.5% cases of

probable re-laying in greater adjutant-storks. In

three nests, we also noted a prolonged incubation

period of 93, 97 and 107 days, which we suspect

was re-laying.

According to Lack (1968), the incubation

period in Ciconiidae varies between 30-33 days.

Incubation period in greater adjutants reported

earlier is 30 days (Saikia and Bhattacharjee

1996b). It seems that our estimate of incubation

period of 35 days is longer than in other storks:

32 days in American wood stork (Heinzman and

Heinzman 1965), 32 days in white stork (Schuz

1972), 29-32 days in Maguari stork (Thomas

1984, 1986), 32 days in painted stork (Desai

et al. 1977), 30-31 days in white-necked stork

(Scott 1975), 28-30 days in Abdim’s stork

Ciconia abdimii (Farnell and Shannon 1987) and

29-3 1 days in Marabou stork (Brown et al. 1 982,

Kahl 1966, Pomeroy 1978a).

The fledging period of greater adjutant-

storks was about 142 days. The fledging period

in Marabou stork was 1 32-135 days (Kahl 1 966,

Pomeroy 1 978a). There was a gap of 67-74 days

between first and last fledging in the greater

adjutant-stork colony. We also found that young

which hatched earlier did not necessarily fledge

earlier. Even siblings of the same nest fledged

asynchronously. However, we could not find out

the exact fledging period according to the first

flight of the nestling, as it was impossible to

monitor all the nests of the colony continuously.

However, we were able to calculate the duration

of hatching to first flight in a few nests (c. 120

days, n=12) which was closer to the fledging

period in Marabou stork.

Comparing its biology with other storks,

it appears that the greater adjutant-stork

resembles, in most of its behaviour and biology,

its congeneric the Marabou stork. The long

breeding season of this colonial bird is an

24

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY 100(1). APR. 2003

BREEDING BEHA HOUR OF GREATER ADJUTANT-STORK

important phase in its life cycle, during which

the crucial need seems to be food.

Acknowledgements

We are grateful to the US Fish & Wildlife

Service, especially to Mr. David Ferguson, SFC

coordinator, for funding the project and the

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the reproductive pattern of the painted stork Ibis

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We thank Mr. J.C. Daniel, Honorary Secretary,

and Dr. Ranjit Manakadan, Senior Scientist,

Bombay Natural History Society for their

suggestions.

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( Mycteria americana) in Florida. Ecological

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reproductive biology of the Greater Adjutant-Stork

Leptopti/os dubius (Gmelin) and L. javanicus

(Horsfield). Ardea 60: 97- 111.

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in India and Ceylon. J. Bombay Nat. Hist. Soc. 67:

453-461.

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relationships of the storks. The Living Bird 10: 151-170.

Kahl. M.P. (1972a): Comparative ethology of the

Ciconiidae. Part 2. The adjutant storks Leptoptilos

dubius (Gmelin) and L. javanicus (Florsfield). Ardea

60: 97-111.

Kahl, M.P. (1972b): Comparative ethology of the

Ciconiidae. The Wood Storks (Genera Mycteria and

Ibis). Ibis 114: 1 5-29.

Kahl, M.P. (1973): Comparative ethology of the

Ciconiidae. Part 6. The Black-necked, Saddlebill and

Jabiru Storks (Genera Xenorhvnchus,

Ephippiorhynchus and Jabiru). Condor 75: 17-27.

Kahl, M.P. (1974): Comparative behaviour and ecology

of Asian Storks. National Geographic Society Research

Reports. Pp. 173-176.

Lack, D. (1968): Ecological adaptations for breeding in

birds. Methuen and Co. Ltd. London, 409 pp.

Marks, J.S. (1983): Prolonged incubation by a long-eared

Owl. J. Field Ornithol. 54(2): 199-200.

Meyerriecks, A.J. (1960): Comparative breeding

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BREEDING BEHA VJOUR OF GREATER ADJUTANT-STORK

behaviour of four species of North American Herons.

Nuttak Ornith. Club 2: 1-158.

Mundkur, T., P. Carr, Sun Hean & Chhim Somean (1995):

Survey for large waterbirds in Cambodia March-April

1994. Gland: IUCN.

Pomeroy, D.E. (1977a): Marabou Stork Leptoptilos

cntmeniferus breeding colonies in Uganda../ East Africa

Nat. His. Soc. and National Museum. 31(161): 1-11.

Pomeroy, D.E. (1977b): The biology of Marabou Storks

in Uganda. I. Some characteristics of the species and

the population structure. Ardea 65: 1-24.

Pomeroy, D.E. (1978a): The Biology of Marabou Storks

in Uganda 11. Breeding biology and general review.

Ardea 66: 1-23.

Pomeroy, D.E. (1978b): Seasonality of Marabou Storks

Leptoptilos cntmeniferus in Eastern Africa. Ibis 120:

313-321.

Rahmani, A.R.. G. Narayan & L. Rosalind ( 1 990): Status

of the Greater Adjutant-Stork (Leptoptilos dubius )

in the Indian Subcontinent. Colonial Waterbirds 13:

138-142.

Saikia, P & P.C. Bhattacharjee (1989): A Preliminary

Survey of adjutant storks in Assam. Asian Wetlands

News 2(2): 14-15.

Saikia, P & P.C. Bhattacharjee (1990): Nesting records

of greater adjutant-storks in Assam, India. Specialist

group on Storks, Ibises and Spoonbills Newsletter

3(1 &2): 2-3.

Saikia, P & P.C. Bhattacharjee ( 1 996a): Studies on some

aspects of the breeding biology of greater adjutant-

stork, Leptoptilos dubius from the Brahmaputra Valley,

Assam. Tropical Zoology 1(1): 57-64.

Saikia, P & P.C. Bhattacharjee (1996b): Some aspects of

the breeding biology of greater adjutant-stork

Leptoptilos dubius in Brahmaputra Valley, Assam. In:

Abstracts: Salim Ali Centenary Seminar on

Conservation of Avifauna of Wetlands and Grasslands.

Bombay Natural History Society, Mumbai.

Schuz, E. (1972): Other stork species. In: Grzimek’s

Animal Life Encyclopedia, (Ed. Grzimek, H.C.). Van

Nostrand Reinhold Co.

Scott, J.A. (1975): Observations on the breeding of the

Woolly-necked Stork. Ostrich 46(3&4): 201-207.

Skutch, A.F. (1962): The constancy of incubation. Wilson

Bull. 74: 115-152.

Thomas, B.T. (1984): Maguari Stork nesting: juvenile

growth and behaviour. Auk 101: 812-823.

Thomas, B.T. (1985): Coexistence and behaviour difference

among the three western hemisphere storks. Pp. 921-

93 1 . In: Neotropical Ornithology. (Eds: Buckley, P. A..

M.S. Foster. E.S. Morton, R.S. Ridgely and F.G.

Buckley) Ornithology Monogr. No. 36.

Thomas, B.T (1986): The behaviour and breeding of adult

Magauri Storks. Condor 88: 26-34.

Thompson. A.L. (1950): Factors determining the breeding

seasons of birds: an introductory review. Ibis 92: 173-

184.

Tomlinson, D.N.S. (1976): Breeding behaviour of the great

white egret. Ostrich 47: 161-178.

Urfi, A.. I. (1993): Breeding pattern of painted storks

(Mycteria leucocephala Pennant) at Delhi Zoo, India.

Colonial Waterbirds 16(1): 95-97.

26

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

CURRENT STATUS OF THE GANGES RIVER DOLPHIN,

PLATANISTA GANGETICA IN THE RIVERS KOSI AND SON, BIHAR, INDIA1

( With one text-figure )

2 3

R.K. SlNHA ' AND GOPAL SHARMA2

Key words: Platanista gangetica , population, threats, conservation,

River Kosi, River Son, Bihar

Surveys were conducted in February and March, 2001 to assess the current status of the Ganges

river dolphin in the Rivers Son and the Kosi. No dolphin was sighted in the entire stretch of

about 300 km of the Son, in Bihar. The local fishermen reported total elimination of dolphin

population in c. 100 km stretch of the Son from the Uttar Pradesh - Bihar border to the Son

Barrage at Indrapuri. During monsoon, the dolphins migrate for about 200 km from the mainstem

of the Ganges into the Son up to the barrage. Dolphins were sighted in the entire stretch of about

300 km of Kosi between the Kosi Barrage at the Indo-Nepal border and its mouth at Kursela in

Bihar. A total of 87 dolphins were sighted in the Kosi during the survey, however, many must

have been missed due to the highly braided channel of the river. In both the rivers, no apparent

source of pollution was found. Siltation and construction of the barrage were observed to be the

main cause of habitat degradation in both the rivers.

Introduction

The Ganges river dolphin Platanista

gangetica , commonly known as susu, is

distributed in the Ganga-Brahmaputra-Meghna

and Karnaphuli-Sangu river systems of India,

Nepal and Bangladesh, between the foothills of

the Himalaya and the estuarine zone.

All the three Asian species of freshwater

dolphins are classified as Endangered or

Critically Endangered. Listed in order of most

to least threatened, are baiji ( Lipotes vexillifer)

in River Yangtze of China (population: a few

tens), bhulan ( Platanista minor ) in River Indus

of Pakistan (population: a few hundreds), and

susu ( Platanista gangetica ), population about

2,500. The fourth freshwater species, boto ( Inia

geoffrensis) is found in the Amazon River System

in South America, population of which is

estimated to be about 5,000. These four are the

'Accepted August. 2001

Environmental Biology Laboratory,

Department of Zoology, Patna University,

Patna 800 005, Bihar, India.

Email : rksinha@mail .girija.net. in

only freshwater dolphin species found in the

world. Obligate river dolphins live only in fresh

water, their physiological and ecological

requirements apparently make it impossible for

them to live in marine waters.

Other small cetaceans are normally

associated with the marine environment, but they

do range far upstream in large Asian rivers.

These include, the finless porpoise

( Neophocaena phocaenoides) in the Yangtze

river of China, and Irrawaddy river dolphin

Orcaella brevirostris in the Ayeyarwady

(formerly Irrawaddy) river of Myanmar,

Mahakam river of Indonesia, and Mekong River

of Lao P.D.R., Cambodia, and Vietnam. Sotalia

fluviatilis is another such species found in the

Amazon-Orinoco river systems of South

America.

The freshwater dolphins have a longer

snout than marine species, which probably help

them in collecting their food in the mud bottom

of rivers. The two species of genus Platanista

found in the Ganga and Indus system are

practically blind, as they have eyes without

crystalline lenses (Herald et al. 1969) and the

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

27

CURRENT STATUS OF THE GANGES RIVER DOLPHIN IN THE RIVERS KOSI AND SON

transparency of their cornea is limited as it is

vascularised (Dawson 1980). While visual acuity

is reduced in these species, they may be able to

form crude images using the narrow aperture of

the pupil in a manner analogous to a pinhole

camera. Amazon and Yangtze river dolphins

have very limited vision.

Evolutionary adaptation to a fluviatile

environment has resulted in a regression of the

eye and the development of a sophisticated

echolocation system, which allows Odontocetes,

the suborder of toothed whales, to ‘see’ their

environment through sound. Pulsed vocalization

produced in specialized air sinuses in the nasal

passages is focused by the ‘melon’ (forehead),

which functions as an acoustic lens. The reflected

pulsed sounds are received back through the

jawbone, transmitted to the middle ear, and then

analysed by the comparatively large brain.

Although the meta-population of the susu

totals over two thousand, isolated subpopulations,

especially in Nepal and in the Karnaphuli-Sangu

Pviver System of Bangladesh, have become extinct

or critically reduced by the barrier effects of dams

and barrages (Haque 1976, Smith et al. 1994).

The distribution range of susu is shrinking, as

evinced by their elimination from many of the

smaller tributaries and upper reaches of the

Ganga, where they were found earlier. Their

population in the mainstem of the larger rivers

is declining as they are being killed both

incidentally as well as directly. Also, they

compete unsuccessfully with humans for

shrinking water and prey resources. The IUCN

recently changed the status of the species from

Vulnerable to Endangered (Baillie and

Groombridge 1996).

Dolphins swim almost constantly on their

side. Shortly after a dive, they spin 90° on their

lateral axis and 180° on their longitudinal axis,

to swim on their side in the direction opposite to

their surfacing direction. The head sweeps up

and down in a scanning motion and the deeper

pectoral fin, or flipper, trails along or slightly

above the bottom. The flippers are thought to

have an important tactile function. Shortly before

surfacing, the dolphins reverse the spin back to

the direction in which the dive began.

In the Karnali river of Nepal, in their far

upstream range as well as in the mainstem of

the Ganga, susu are found most often in ‘primary

habitats’ where convergent streams create an

eddy counter-current system in the mainstream

flow (Smith 1993). Less often, the dolphins are

found in “marginal habitats” where the river

meanders and creates similar eddy counter-

current systems, which are also areas of high

human use, making them particularly vulnerable

to local environmental disturbances.

The river dolphin often takes advantage

of the ecotone created by the transition between

scour pools and running waters, visible as eddy

turbulence. They prey on species migrating along

the mainstem, while monitoring foraging

opportunities from within the hydraulic refuge

of counter-currents.

Current Status of the Ganges river dolphin

The total population of the susu was

roughly estimated to be only 4,000-5,000 (Jones

1982). Dolphins are sighted throughout the

Ganga from the Middle Ganga Barrage, Bijnor

(129 km downstream of Haridwar) to its mouth

at Sagar Island in the Bay of Bengal. About

35 susus have been isolated between the two

barrages at Bijnor and Narora (166 km) in Uttar

Pradesh (Sinha et al. 2000). The population

between Narora and Allahabad (about 500 km)

in low water season (January-March) is very

sparse (a few tens) (Sinha 1999). In the lower

reaches of the Ganga in West Bengal, only

152 susus were sighted in the Bhagirathi-

Hooghly river system below Farakka Barrage

(Sinha 1997). Maximum dolphins survive in the

Ganga mainstem between Allahabad and

Farakka. Less than 100 have been estimated in

Chambal river, a tributary of the Yamuna. A total

of about 2,000 dolphins have been estimated in

28

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

CURRENT STA TUS OF THE GANGES RIVER DOLPHIN IN THE RIVERS KOSi AND SON

the Ganga-Brahmaputra river systems in the

Indian territory (Sinha 1999). A few hundreds

are present in Bangladesh. A comprehensive

review of the susu’s status in the entire Ganga

system including tributaries has recently been

documented (Sinha et at. 2000).

In the nineteenth century, dolphins were

plentiful in the entire distribution range, though

no actual data on populations is available. They

were found in the Yamuna as far as Delhi, even

in May when water was very low (Anderson

1879). In the last couple of decades, no dolphin

has been sighted in the Yamuna at Delhi. Their

current distribution in the Yamuna is mainly

beiow the confluence of the Chambal and

Yamuna near Etawah. In most of the small

tributaries, dolphins have become locally extinct

or are sighted only in the rainy season.

Platanista gangetica is legally protected,

being included in Schedule I of the Indian

Wildlife Protection Act (1972) and in Appendix I

of Convention of international Trade in

Endangered Species of Wild Fauna and Flora

(CITES), which prohibits trade in dolphin

products by signatory countries.

Past Distribution in Bihar

Anderson (1879) has mapped the

distribution of dolphins in the entire stretch of

the Ganga, all its tributaries both large and small,

in the state of Bihar. No detailed study, especially

in the Ganga and some of its tributaries was

conducted until more than 100 years later (Sinha

1996). Flowever, even this study was far from

complete, as it was mainly conducted in the

mainstem of the Ganga. When interviewed, the

locals in North Bihar reported that about 40-50

years ago during monsoon, the susu were

frequently sighted in all the rivers, including very

small rivers, and connected water bodies. But,

unfortunately, no baseline data on the status are

available.

Though a few papers on the Ganges river

dolphin were published from Bihar earlier (Nath

1974, Gupta 1986, Ali 1992, Singh and Ahmed

1994, Kumar 1996) nothing specific is

mentioned about its status and the reports carry

only a general account of the species. Moreover,

the papers record only casual observations on the

animal in the River Ganga. Based on a systematic

study, Sinha (1996, 1997 and 1999), Sinha

et al. (2000) reported the current status and

distribution of the susu in the Ganga and many

of its tributaries. Though the overall estimate of

susu abundance in the entire distribution range

is not known, the largest sub-population occurs

in the mainstem of the Ganga and its tributaries

in Bihar. Most of the tributaries or parts thereof

are yet to be surveyed thoroughly and these

surveys were conducted in an effort to bridge this

gap-

Study Area

(Fig. 1)

River Son: The River Son originates from

Amarkantak Hills at Sonabhadra, in Madhya

Pradesh, at an elevation of 600 m. It flows

northwards through Madhya Pradesh, Uttar

Pradesh and Bihar before it discharges into the

Ganga at Haldi-Chhapra village near Maner,

about 35 km upstream of Patna. In Uttar Pradesh,

it receives the Rihand tributary across which the

Rihand dam was constructed in 1963. Though

the Son is a perennial river, the main source of

water is rain. Its total length is 784 km of which

about 300 km are in Bihar. It enters the state of

Bihar near the village Domarkhoha in Rohtas

district. After flowing for about 35 km in Bihar,

it receives the River North Koel from Chhota

Nagpur Plateau of South Bihar (now Jharkhand

State). About 65 km downstream of the

confluence, the Son Barrage was constructed in

1965 at Indrapuri, about 15 km upstream of

Dehri, to divert the river water through three

irrigation canals — Patna Canal on the right side

and Western Canal, which is divided into Buxar

and Ara Canals on the left side. A weir

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

29

CURRENT STA TUS OF THE GANGES RIVER DOLPHIN IN THE RIVERS KOSI A ND SON

30

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

Fig, 1 : Map showing course of the Rivers Kosi and Son in Bihar

CURRENT STATUS OF THE GANGES RIVER DOLPHIN IN THE RIVERS KOSIAND SON

constructed at Dehri in 1869-79 provided water

for irrigation of 0.35 million ha of land. As the

weir became old, the new barrage at Indrapuri

was constructed which created a physical barrier

for the migratory aquatic animals including

dolphins of the river. The irrigation canals have

converted the entire command area into a ‘Grain

Bowl’ in Aurangabad, Jehanabad, Patna, Rohtas,

Bhojpur, Kaimur and Buxar districts of Bihar.

But this left almost no water downstream of the

barrage to maintain its status as a river. The bed

of the river consists mainly of coarse sand, which

can retain little organic detritus. Extraction of

sand as building material throughout this stretch

of the river has added to the degradation and

destruction of the river habitat.

River Kosi: The River Kosi (965 km)

originates in Tibet at an altitude of 5,490 m and

flows through Nepal; after running for c. 285 km

in Bihar it joins the Ganga near Kursela in

Katihar district. The Kosi is formed by the

convergence of three rivers, the Sun Kosi, Arun

Kosi and Tamur Kosi in Nepal. After the

confluence, the river flows through a narrow

gorge for 10 km and enters the plains at Chatra,

traverses another 25 km and enters India near

Hanuman Nagar. The total drainage area is

74,500 sq. km of which 1 1 ,000 sq. km lies within

India.

The Kosi basin is the third largest in area

in India. Its waters have the highest rate of

siltation among the rivers of the state (average

annual suspended load is 2,774 tonnes/sq. km)

and it has a steep gradient. The Kosi is a

torrential river of the mountains that has a

catchment area too large for its relatively short

course. Due to these topographical and

meteorological features it is rated as one of the

most problematic rivers of the world and is noted

for its rapidity and unstable banks. Thus, in about

200 years, the river has moved 1 12 km laterally

from Purnea to its present position.

Average discharge in normal years for the

Kosi is estimated to be 1,75,000 cusecs. The

average run off during monsoon (June-

September) is about 83% and only 17% in the

rest of the year. July-August is the period of peak

flow, whereas January-February is the leanest

period (Datta Munshi and Datta Munshi 2000).

The Kosi Barrage at Hanuman Nagar was

commissioned in 1965 from which two canals

take off on either side to irrigate nearly one

million hectares of land in Nepal and India. The

barrage is intended to prevent the river from

moving sideways. The important tributaries of

the Kosi are rivers Bagmati and Kamala Balan.

Survey Methodology

Downstream survey was conducted using

country boats along the entire stretch of the rivers

Son and Kosi. Following the recommendation

of international experts (Perrin et al. 1989),

direct count method was used to record the

number of dolphins. Although searches were

conducted continuously along the total length of

the river under study, areas of confluence,

meandering, downstream of sandbars etc.

received greater attention. Search effort in these

areas was maintained for at least thirty minutes

to avoid missing extremely quiescent or long-

diving animals. During survey, best, high and

low estimates of the number of animals in the

groups were recorded. The high and low estimate

was used to reflect confidence in the accuracy of

the best estimate. The low estimate was

considered to be an absolute minimum count and

the high estimate as maximum count. Identical

best, high and low estimates were used to indicate

a high level of confidence in our best estimate.

Occasionally the dolphin appears to follow the

boat, which adds uncertainty to whether the

subsequent sighting is of a new or the same

animal. In this case, a low estimate of zero was

used to reflect the possibility of making double

counts. The dolphin’s long dive time,

unpredictable movements and quiescent

behaviour also make single counts unreliable.

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

31

CURRENT STATUS OF THE GANGES RIVER DOLPHIN IN THE RIVERS KOSIAND SON

During quiescent behaviour, the dolphin surfaces

without an audible blow exposing only the

uppermost dorsal surface of the melon. If

subsequent surfacing, or confirmation by a second

member of the team did not substantiate such a

sighting it was given a best and low estimate of

zero and high estimate of one. Estimates were

arrived by concensus among the team of observers

that initially sighted the dolphin.

When a sighting was made the size, sex

and colour among other features were

ascertained. Diagnostic characters for individual

animals (visible scarring and deformities, ratio

of rostrum length to the height of melon, etc.)

were noted and sketched. Photographs were taken

by a 35 mm SLR camera, equipped with a

300 mm telephoto lens and motor drive.

Data on the frequency of dolphin sightings

per visit to habitat locations was recorded

throughout the study and later processed.

Results and Discussion

River Son: A field survey was conducted

in River Son between February 22 and 28, 2001.

The survey started from a small tribal village

Domarkhoha located at the Uttar Pradesh - Bihar

border. Markings of large soft-shell turtles on

the sand bar were seen, but no dolphin was

sighted. Farther 5 km downstream at Belduria

ghat / Newaria ghat, the locals informed that no

aquatic wild animals except large soft-shell

turtles were found in the area. However, a good

number of dolphins and crocodiles had been

reported in the river about 40-50 years ago. The

Kaimur hill range is on the left bank (Rohtas

district, Bihar) of the river between Domarkhoha

and Newaria ghat. After this ghat, the river

channel meanders towards the right bank

(Garhwa district, Jharkhand) and a large island

of about 100-150 sq. km has been formed which

is used for farming. On the left bank, a 1-3 km

wide plain is heavily cultivated for wheat, potato

and vegetables. The river flows close to the left

bank again at Teura ghat. No dolphin was sighted

in this stretch. As reported by a fisherman, there

used to be many deep pools of water in the river,

but a high rate of siltation had filled up all these

pools resulting in loss of habitats of dolphins and

crocodiles. About 10 km downstream of Teura

ghat is Uli ghat on the left bank. Opposite Uli,

the River North Koel from Daltongunj and

Garhwa districts discharges into the Son.

However, the main channel of the Son flows close

to the left bank. In spite of good habitats for

dolphins near Uli ghat, no dolphin was sighted.

It was learnt that about 10 years ago there used

to be some dolphins in the area, but after the

construction of the Rihand Dam in Uttar Pradesh

the flow of the river had reduced which had

affected the dolphin population. Another local

fisherman informed about the killing of one

dolphin about 15 years ago at village Jhitikia

opposite Amjhore, about 30 km downstream of

Uli. Earlier, the fishermen of village Deuri,

located near Jhitikia, practised dolphin oil

fishing. Frequent sightings of otters by the locals

were reported near this ghat (river bank).

The river channel is highly braided and

several small as well as large sand bars have been

formed in the river downstream of Uli. The bed

consists of coarse sand, mainly quartzite. The

flow in the river increases after the North Koel

joins the Son at village Tilothu, about 10 km

upstream of Son barrage. It was reported that

after construction of the barrage at Indrapuri in

1965, dolphin migration stopped and no more

dolphins are sighted now. However, during the

flood every year juvenile gharials occasionally

drift into this area. Locals reported the killings

of soft-shell turtles and presence of otters in the

river in this area. About 200 fishermen fish in

the river at Tilothu. The depth of river water was

only about 50 cm to a metre at most places and

many times it was difficult for a boat to float

down in the highly braided river.

A large number of sandbars occupied by

grasses and other thick vegetation have formed

32

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

CURRENT STA TUS OF THE GANGES RIVER DOLPHIN IN THE RIVERS KOSI A ND SON

behind the barrage. They provide good habitat

for otters and many species of migratory birds.

The local fishermen informed us of sightings of

dolphins below the barrage during the high floods

every year. They migrate up to the barrage from

the Ganga river about 200 km downstream.

About 18 km downstream of the barrage

is the 3.06 km long Dehri Road Bridge,

commissioned in February 1965. The river

channel is highly braided and the depth of the

river flow is insufficient to float even a country

boat. Farther 30 km downstream of Dehri — a

ferry ghat, is Mahadeva ghat on the left bank

and Daud Nagar on the right bank. The river

bed is more than 3 km wide and is widely used

for vegetable farming, both at Dehri and

Mahadeva ghats. Most of the fishermen were

busy in farming activities. A few of them were

found doing subsistence fishing using small drag

nets. A local fisherman informed us of the

sighting of dolphins occasionally during flood

season only. Otters are also sighted during the

rainy season. The soft-shell turtles are found and

killed during summer season. About 40 km

downstream of Mahadeva ghat is Malhi Patti

ghat on the right bank near Baiderabad. No

dolphin could be sighted in this stretch; however,

locals informed that during monsoon 3-7

dolphins were sighted every year. Heavy siltation

in the river has destroyed the habitat of the susu.

Otters are reportedly sighted occasionally near

the Malhi Patti ghat. Turtles are found mainly

during the monsoon. Farther 5 km downstream

near Arwal is Ahiyapur ghat where a very good

habitat for dolphin with eddy counter-current

exists, but no dolphin was sighted. However,

locals informed that during monsoon 10-15

dolphins are sighted here. Reportedly otters and

turtles are also found. It is an important fish

spawn cpllection centre in the monsoon. Sighting

of five adult dolphins during the last monsoon

of 2000 at Mohammadpur ghat, 20 km

downstream of Arwal was reported by a local

fisherman. During the survey, water flow in the

River Son at this ghat was not enough to sustain

a dolphin population in this season. Similarly,

at Udaipur ghat near Pali, otters were found in

the boulder pitched river bank, but no dolphin

was sighted. It was reported that dolphins could

be seen only during monsoon. The killing of a

dolphin, three years ago, was reported by the

locals. At Koilwar, the biggest site of sand mining

in the River Son, 1 5-20 susu were reported during

the monsoon. Earlier the dolphins used to be

killed here by harpooning. The local fishermen

consume both the meat and oil of the dolphin.

The drift gillnets with large mesh size are

dangerous for dolphins during monsoon as they

get entangled in these net.

River Kosi: A survey in the River Kosi

was conducted between March 2 and 10, 2001.

Sighting records of dolphins are given in Table 1.

The survey was started from the Kosi Barrage at

Birpur. Water depth in the reservoir of the

barrage was only about 5 m. The river water was

being diverted to irrigation canals and almost

no water was allowed to flow into the river

downstream of the barrage. Hence, the river

water below the barrage was very shallow, and

not enough to sustain dolphin population below

the barrage. When the gates of the barrage were

opened in April, the dolphins from farther

downstream moved to the barrage site. In April

1994, dolphins were sighted here (Sinha et al.

2000). One or two dolphins are reportedly killed

here every year. The locals reported that the

dolphins cross the barrage gates both ways in

the monsoon season. This needs to be confirmed,

but if true it is good for the survival of the

dolphins not only in India but also in Nepal.

Further 42 km downstream of the barrage at

Bhaptiahi, two dolphins were sighted. Here the

water depth was about 5 m. As water in the river

in this stretch was quite low no dolphin was

sighted between the barrage and Bhaptiahi.

Poison-fishing in the floodplain wetlands was

reported by the local fishermen, as also killing

of soft-shell turtles. The river channel was highly

JOURNAL , BOMBAY NATURAL HISTORY SOCIETY 100(1), APR. 2003

33

CURRENT STA TUS OF THE GANGES RIVER DOLPHIN IN THE RIVERS KOSI AND SON

Table 1 : Status of the Ganges dolphin in the River Kosi during March, 2001

H - Highest, B - Best, and L - Lowest; Total No. of the dolphins: H-87, B-78, L-73; * - Reported by locals

braided and shallow in this stretch. Farther

downstream, four aduits, two sub adults and one

calf dolphin were sighted at Sujanpur, a village

on the left bank near Thirbitia. The locals were

expecting the number of dolphin to increase up

to 20-25 after the gates of the barrage would be

opened in April. At this site, two channels of the

Kosi join resulting in increase in water flow. The

water depth near the village was 6 to 8 m. Locals

here do not kill dolphins, though turtles and

avifauna were poached. Otters were reported in

the area. About 9 km downstream, a very good

dolphin habitat was found near the spur, but no

dolphins were sighted. However, just 3 km

further downstream at Situharghat three adults,

two sub-adults and one calf dolphin were sighted

(Highest 6 - Best 5 - Lowest 5) in one group.

The locals reported killing of a dolphin at this

site. Farther downstream, a highest of 9 and 8

each of best and lowest count of dolphins

34

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY / 00(1), APR. 2003

CURRENT STA TUS OF THE GANGES RIVER DOLPHIN IN THE RIVERS KOSI AND SON

(4 adults, 3 sub-adults and one calf) were sighted

at E2 Ghat (93.2 km from the barrage). About

20-25 hard-shell turtles were also sighted.

Erosion of the left embankment was noticed here.

The locals reported degradation of dolphin

habitat due to the erosion. Incidental killings of

dolphin by gillnets were reported, however, no

intentional killing was reported. Soft-shell turtles

of 20-40 kg were reported at this site. Poison-

fishing in floodplain wetlands was reported by

the locals. The fishing activity was found to be

very low. Fourteen kilometres further

downstream is Baluaha ghat, another Ferry ghat.

The river flows close to the left bank in this

stretch. One dolphin was sighted here, however,

the locals reported sighting of a good number of

dolphins around the year. A large number of

small turtles were also sighted. Ten kilometres

further downstream is Dengrahi ghat where a

highest of 9 and 8 each of best and lowest counts

of dolphins were sighted in two groups including

6 adults and 3 sub adults. It was a very good

habitat for the dolphins and the river was flowing

mainly along the left bank. Gharials were

reportedly sighted here. Soft-shell turtles up to

50 kg are killed regularly, however, dolphins are

not killed here according to the locals. About

23 km further downstream is Koparia, 125 km

downstream of the barrage. The east embankment

of the Kosi terminates here. The river channel was

highly braided and it was impossible for a boat to

float downstream from Koparia.

The survey was resumed from Bad la ghat

in River Bagmati, an important tributary of Kosi.

In this area, the highly braided channel of River

Kosi flows parallel to Bagmati being separated

only by 7 km. A channel of Kosi discharges into

Bagmati upstream of Badla ghat. Near the Badla

ghat railway bridge, a highest of 10 and 9 each

of best and lowest counts of dolphins, including

6 adults, 3 sub-adults and one neonate were

sighted in two groups. The river water was quite

deep and meandering, and with erosion, eddy

counter-currents were creating a good habitat for

the dolphins. The water current in the river was

very slow but the deeper zone sustained a good

population of the dolphins. At Chautham ghat,

2 1 km farther downstream, 4 adult dolphins were

sighted. Ten kilometres downstream of the

Chautham ghat, the Bagmati joins the Kosi near

Dumri Road Bridge. The Bagmati is deeper than

Kosi near the confluence. Absence of dolphins,

at the confluence of Bagmati and Kosi, was

disappointing. The local fishermen reported non-

availability of fish in the area as a reason for the

absence of the dolphins.

Three adult dolphins were sighted about

2 km downstream the Dumri bridge. Dolphins

are reportedly neither killed nor eaten by the

locals in this area. The dolphin oil is used only

as folk medicine. Otters are occasionally sighted

farther 35 km downstream at Vijay ghat,

Naugachhia where a highest of 20, best of 17

and lowest of 16 dolphins in three groups were

sighted. The villagers here killed dolphins and

reportedly over 30 containers of dolphin oil were

available on that day in the village. Here the river

channel was very wide (about 3-4 km) and water

near the right bank was about 15 m deep.

Downstream from Vijay ghat and about 1 km

upstream of Kursela Bridge a dry channel of

Kosi, Chhoti Kosi joins the main channel — the

Bari Kosi. At the confluence of the two, a highest

of 16 and 14 each of best and lowest counts of

dolphins including 8 adults, 4 sub adults, one

calf and neonate each were sighted. Four

dolphins were sighted in the dead channel.

Subsistence fishing with lift-net and cast-net was

observed here. A local businessman accompanying

our team up to this point informed us that a large

(20 1 .5 kg) dolphin was caught at Kursela in 1 995

and he had transported it to Siliguri fish market

himself. Otters were reported to be abundant

here. The Kosi discharges into the Ganges 3 km

downstream of Kursela Bridge.

Thus, in the Kosi a highest of 87 and best

of 78 dolphins were sighted distributed almost

along the entire stretch of the river. As the river

JOURNAL BOMBAY NATURAL HISTORY SOCIETY 100(1), APR. 2003

35

CURRENT STATUS OF THE GANGES RIVER DOLPHIN IN THE RIVERS KOSI AND SON

channel is highly braided many more must have

been missed.

Threats to the Survival of Dolphins

The main threats to dolphin survival are

the construction of barrages, heavy siltation,

farming in the river bed using chemical fertilisers

and organochlorine pesticides, use of detrimental

fishing gill nets, shortage of fish in the rivers,

and incidental as well as intentional killings of

the dolphin for oil and meat. The barrages have

created a physical barrier for all the migratory

species of the rivers, including dolphins. As there

is no industry and urban settlement on the banks

of the two rivers, the possibility of organic

pollution is remote. However, non-point sources

of pollution, namely chemical fertilisers,

organochlorine pesticides and heavy metals

cannot be ruled out.

Conservation Measures

The heavy siltation in the rivers has

degraded the dolphin habitat. There is a serious

need for mass scale plantation in the catchment

area of the rivers to reduce the silt load entering

the river system. For this, international efforts

are required as the Kosi flows through Nepal and

the greater part of the silt load originates there.

A minimal flow of water in the rivers must be

allowed even during the lean season, so that the

Refe

Ale Mohammed S. (1992): The Gangetic Dolphin. Myforest

28(3): 245-250.

Anderson, J. (1879): Anatomical and Zoological

researches: comprising an account of zoological

results of the two expeditions to western Yunnan in

1 868 and 1 875; and a monograph of the two cetacean

genera Plcitanista and Orcaella. B. Quaritch, London,

Vol.I&lI.

Baillie, J. & B. Groombridge (Eds) (1996): IUCN Red

List of Threatened Animals. IUCN, Gland,

Switzerland, and Conservation International,

Washington DC, pp. 70+368 and 10 annexure.

Datta Munshi, J.S. & J. Datta Munshi (2000): The

dolphin population can survive and migrate.

Another serious need is fishery regulation in the

rivers. No effective regulation exists. The

detrimental gill-nets must be banned and only

dolphin friendly nets should be allowed. The

efficacy of legal measures to protect dolphins is

non-existant. Hence, there is a need to motivate

wildlife and other administrative officials to

implement the Wildlife (Protection) Act

efficiently. An alternative fish attractant, other

than dolphin oil that is used throughout the state,

should be identified.

Recently, fish scrap oil has been field-tested

for three years and found to be a good alternative

to dolphin oil as fish attractant (Sinha 2002).

However, there is again a serious need of a long

term extension programme for educating the

fishermen to use the alternative fish attractant.

Education and awareness programmes to

educate the target group, i.e. fishermen, officials,

school and college children, and the common

man will be helpful in conserving the dolphin.

Periodical monitoring of the dolphin

population will help in evaluating the effects of

Dolphin Conservation Programmes, if any.

Acknowledgements

Financial help by WWF-India is duly

acknowledged. Help rendered by the local

fishermen was valuable and praiseworthy.

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sustainability of hydrological cycle of wetlands of

Kosi river basin on North Bihar, India. In: Water

Recycling and Resource Management in the

Developing World (Eds: Jana, B.B., R.D. Banerjee,

B. Gusterstam & J. Heeb). University of Kalyani,

India and International Ecological Engineering

Society, Switzerland. Pp. 665-674.

Dawson, W.W. (1980): The cetacean eye.. In: Cetacean

Behavior: Mechanisms and Functions (Ed.: Hannan,

L.M.). Wiley-Interscience, New York.

Gupta, D. (1986): The Gangetic dolphin Platanista

gangetica (Lebeck, 1801 ). Pp. 553-562. In: Wildlife

Wealth of India (Resources and Managements)

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JOURNAL, BOMBAY NATURAL HISTORY SOCIETY 100(1), APR. 2003

CURRENT STATUS OF THE GANGES RIVER DOLPHIN IN THE RIVERS KOSI AND SON

(Ed.: Majupuria, T.C.). Teepress Service, L.P.

Bangkok.

Haque, A.K.M. Aminul (1976): Comments on the

abundance and distribution of the Ganges susu

Platanista gangetica , and the effects of the Farakka

Barrage on its population. FAO ACMRRR, Scientific

Consultation on Marine Mammals, AGMRR/MM/SC

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Herald, E.S., R.L. Brownell Jr., F.L. Frye, E.J. Morris,

W.E. Evans & A.B. Scott (1969): Blind river

dolphins: first side - swimming cetacean. Science 1 66:

1408-1410.

Jones, S. (1982): The present status of the Gangetic susu

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the Indus susu P. minor Owen. FAO Advisory

Committee on Marine Resources Research Working

Party on Marine Mammals. FAO Fish Ser. (5)4: 97-

115.

Kumar, Arvind (1996): Impact of industrial pollution on

the population status of endangered Gangetic dolphin

{Platanista gangetica) in the river Ganga in Bihar,

India. Pol. Arch. Hydrobiol. 43(4): 469-476.

Nath, Bhola ( 1 974): On some aspects of habit and habitat

of the Gangetic dolphin {Platanista gangetica)

(Lebeck) in the river Ganges at Patna. The Naturalist,

Bull, of the Bihar Natural History Society Vol. 1 , pp.

6-7.

Perrin, W.F., R.L. Brownell Jr., Zhou Kaiya & Liu

Jiankang (Eds) (1989): Biology and Conservation

of the River Dolphins. IUCN Species Survival

Commission Occasional Papers 3.

Singh, A.K. & S.H. Ahmed (1994): Gangetic Dolphin.

Fishing Chimes , pp. 9-1 3.

Sinha, R.K. (1996): Bioconservation of the Gangetic

Dolphin Platanista gangetica. Final Technical

Report, Patna University, submitted to National River

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and Forests, Govt, of India, New Delhi. Pp. 69 +

42 Tables + 21 Plates + 5 Appendices.

Sinha, R.K. (1997): Status and Conservation of Ganges

River Dolphin in Bhagirathi-Hooghly river systems

in India. Int. J. ofEcol. Env. Sc. 23(4): 343-355.

Sinha, R.K. (1999): The Ganges river dolphin — a tool

for baseline assessment of biological diversity in River

Ganges, India. Final Technical Report, Patna

University. Tech. Rep. No. 1/99 submitted to

Biodiversity Support Program (BSP), a Consortium

of World Wildlife Fund, The Nature Conservancy and

the World Resources Institute, Washington, DC.

Pp. 34+16 Tables + 6 Figures.

Sinha, R.K. (2002): An alternative to dolphin oil as a fish

attractant in the Ganges river system: Conservation

of the Ganges river dolphin. Biol. Conserv. 107: 253-

257.

Sinha, R.K., B.D. Smith, G. Sharma, K. Prasad, B.C.

Choudhury, K. Sapkota, R.K. Sharma & S.K. Behera

(2000): Status and distribution of the Ganges susu

{Platanista gangetica) in the Ganges River system

of India and Nepal. In: Biology and Conservation of

Freshwater Cetaceans in Asia (Eds: Reeves, R.R.,

B.D. Smith & T. Kasuya). IUCN, Gland, Switzerland

and Cambridge, UK. viii + 1 52 pp.

Smith, B.D. (1993): 1990 Status and conservation of the

Ganges River dolphin {Platanista gangetica) in the

Karnali River, Nepal. Biol. Conserv. 66: 159-170.

Smith, B.D., R.K. Sinha, K.U. Regmi & K. Sapkota (1994):

Status of Ganges river dolphins Platanista gangetica

in the Mahakali, Karnali, Narayani and Saptakosi

rivers in Nepal and India. Marine Mamma! Science

10(3): 368-375.

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

37

CROP DEPREDATION BY WILDLIFE ALONG THE EASTERN BOUNDARY

OF THE KALAKAD-MUNDANTHURAI TIGER RESERVE, SOUTHERN INDIA1

( With two text-figures )

PUNIDAN D. JEYASINGH2'3 AND PR1YA DAVIDAR2

Key words: Sus scrofa cristatus , wild pigs, crop loss, crop raid,

human-wildlife conflict, electric fence

Crop raiding patterns by wildlife at ten villages along the eastern boundary of the Kalakad-

Mundanthurai Tiger Reserve (KMTR), southern India were studied. The Indian wild pig

(Sus scrofa cristatus) was identified as the major crop pest in this area and the crop loss caused

by it was quantified. The effectiveness of an electric fence to prevent crop raids by wild pig was

also assessed, comparing (1) mean group size of pigs, (2) extent of damage in sq. m, (3) actual

and potential loss, and (4) frequency of wild pig raids. The actual loss was estimated at

257. 19 kg ha1 accounting for approximately 7% of the actual produce. The electric fence was not

effective in preventing crop raiding by the wild pig. The number of wild pigs was not correlated

with the extent of damage. Extent of damage might be a factor of time spent in the paddy field,

suggesting that wild pigs might raid paddy fields for habitat requirements rather than for nutritional

requirements. It is vital to understand crop-raiding patterns prior to the implementation of control

strategies.

Introduction

Strategies for reconciling human needs and

conservation interests in areas abutting nature

preserves are critical to the success of

conservation plans (Gradwohl and Greenberg

1988, Western and Pearl 1989). Crop depredation

by wildlife can occur more frequently than the

highly publicized and prioritized, but sporadic

livestock raids. Over the years, farmers have

developed a variety of measures such as fencing,

culling, dogs, firecrackers, fire and drums to

chase away pest species and reduce crop loss to

wild animals. Today, when many crop raiding

species are protected by law and are focal points

for conservation, the need for effective and long-

term control methods is felt. Some of the control

measures include physical barriers, selective

culling and environmental control methods, such

'Accepted July, 2002

2Salim Ali School of Ecology, Pondicherry University,

Pondicherry 605 014, India.

3Present Address: Department of Zoology,

University of Oklahoma, 730 Van VleetOval, Room 314,

Norman, OK 730 19-0235, USA.

E-mail: [email protected]

as providing better habitat in the forest interiors,

away from human habitation (Sukumar 1992).

Recently, electric fencing has become one of the

methods widely used by both private farmers and

the government to prevent crop raids by wild

animals.

In a predominantly agricultural and

densely populated country like India, conflicts

between humans and wild animals are frequent,

and preventing conflict should be a conservation

priority (Sukumar 1992). Damage by the Asian

elephant (Elephas maximus) has been estimated

at c. $0.5 million/per year in southern India

(Sukumar 1989). Although considerable work

has been done on the crop damage patterns and

management strategies for larger wildlife, such

as the elephant and tiger (landmark studies

include Sanyal 1987, Sukumar 1991), work on

wild pigs is sparse, except for Tisdell (1982),

and Ahmed (1991).

Crop damage patterns along the eastern

boundary of the Kalakad-Mundanthurai Tiger

Reserve by different wildlife species and wild pigs

in particular were analyzed and crop loss due to

wild pigs, the major pest in the area, were

38

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

CROP DEPREDATION BY WILDLIFE

estimated. The study also attempted to

understand the possible cues for crop raiding by

the wild pig in the area, and to suggest effective

control measures. An electric fence, erected with

the aid of the World Bank, was assessed for its

effectiveness against the wild pig.

Study Area

The study was carried out from December

1998 through March 1999 in ten villages located

in the eastern boundary of the Kalakad-

Mundanthurai Tiger Reserve (KMTR), southern

India. KMTR is situated at 08° 25' - 08° 35' N

and 77° 25' - 77° 35' E and covers 795 sq. km of

the southernmost protected area in the Western

Ghats complex. Along the northeastern boundary

of the Reserve, an 8.7 km long electric fence was

erected in 1996 with partial funding from the

World Bank to control crop damage by wildlife

(Fig. 1). The fence is about 160 cm high and

consists of 7 wires running parallel to the ground

attached to granite posts at intervals of 2.5-3 m.

Wires 2 and 5 are ground wires. The rest of the

wires have an output of 36V generated from a

12V solar battery. The lowest wire is c. 10 cm

from the ground and the second wire (earth wire)

is c. 30 cm from the ground.

Ten villages located along the 26 km

eastern boundary of the KMTR were selected

for the study. Four villages were located in

the Mundanthurai area and were separated

from KMTR by the electric fence. The fifth

village abutting the Mundanthurai section

(Pudukudierrupu) was not protected by an electric

fence. The other five villages bordered the

Kalakad section of KMTR (Fig. 1). All the study

villages had lowland teak dominated deciduous

forests, thickets, and scrub jungles,

representative of the vegetation of the buffer zone

of KMTR. All the crops grown along the

boundary, such as paddy, banana, sugarcane, and

groundnut, were also cultivated in the 10 study

villages (Table 1).

Table 1: Details of the Study Villages

TDDD = Teak Dominated Dry Deciduous, Present - *,

Absent -x

Methods

Farmers’ Perceptions

A questionnaire was circulated among

farmers in the 10 study villages to estimate

wildlife raids, crop loss, and other relevant

information. Results from the questionnaire

indicated the actual or realized yield and crop

loss due to wild pigs. Potential yield and crop

loss estimates were based on quadrat data. Both

the questionnaire as well as quadrat data were

used to estimate crop loss because farmers tended

to underestimate production and overestimate

crop loss due to wildlife.

Crop Raids

Information regarding the wildlife species

involved in the raid, group size, and the time of

raid were obtained from the farmer who had

witnessed the raid or through direct observation.

The species responsible for the highest proportion

of raids and damage in the area was identified

as the major crop pest species.

Crop Damage

Crop damage was assessed within 24 hrs

of the damage. The site was visited, and the mean

plant density (MPD) was estimated to assess crop

loss. MPD measurements were determined in the

JOURNAL , BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

39

CROP DEPREDATION BY WILDLIFE

Fig. 1: Kalakad-Mundanthurai Tiger Reserve

damaged and undamaged sites in the field. MPD

was estimated by laying 12-30 quadrats (30 cm

x 60 cm) at random in the undamaged part of

the field. The field area was noted. The total

number of plants in the field (P ) was estimated

by

Pu = MPD x Area of the field in sq. m (Eq 1)

A sample of five plants was taken from

each quadrat for which the number of grains and

mean weight per grain were calculated.

The damaged area was identified as the

portion of the field where all the stalks had been

40

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

CROP DEPREDATION BY WILDLIFE

flattened to the ground and could not be

harvested. The average length and width or

radius within the damaged areas were measured

and the area of the closest resembling regular

shape, i.e., square, rectangle, or circle was

calculated in sq. m. The total number of damaged

plants (P ) was calculated using the formula

Pd = MPD x damaged area in sq. m(Eq 2)

The number of grains per damaged plant

was counted and weighed. The potential produce

(standing crop) was estimated by

P =P N W / 1000(Eq 3)

p u g g V i 7

where, Pp = Potential produce in kilograms,

Pu = total number of plants in the field,

Ng = mean number of grains per plant, and

= mean weight of one grain in grams.

The potential loss (P,) in kilograms due to

crop damage can be estimated by

P =P,N W / 1000

I d g g

where Pd = total number of damaged plants

The average actual produce (kilograms of

paddy sold by the farmer in the market) was

estimated using the farmer’s claim and a brief

survey of the buyers. The market value of the

crop was obtained from the farmers and

wholesalers to arrive at the actual loss in

kilograms and rupees. The potential loss was

the loss measured during this study based on

the yield measured by the quadrat study. Loss

per raid thus calculated was used to extrapolate

the loss for a month using the mean number

of wild pig raids in each of the ten study

villages.

Effect of the electric fence

The villages were divided into villages

protected by an electric fence and unfenced

villages. A t test or its non-parametric equivalent,

the z test, was used to test for significant

differences between the two in the following

parameters:

1 . Mean group size of pigs.

2. Extent of damage in sq. m.

3. Actual and potential loss.

4. Frequency of wild pig raids.

5. Frequency of larger wildlife sightings on

cropland.

Results

Crop raiding patterns

A total of 121 farmers were interviewed,

and 39 instances of fresh crop raids were

observed. Of these, 35 were on paddy fields,

3 on banana plantations and 1 on sugarcane. The

results show that of the 1 1 species of crop pests

reported, wild pigs were the most and accounted

for 99% of the crops damaged during the study

(Table 2). Crop raiding patterns of wild pigs on

paddy were dependent on the age of the crop

(Ahmed 1991, Jeyasingh 1999). The ears of

paddy were nipped off, chewed well, the juice

ingested and the fibre spat out. In banana

plantations, wild pigs fed on the stem, flower,

and fruit by reaching up on their hind legs and

biting the stem to bring down the canopy. On

younger plants, they nosed around the plant to

expose and feed on the tender shoot. Sugarcane

stems were consumed voraciously from the

bottom for the juice and fibre.

Other wildlife reported to stray outside the

Reserve boundary included larger herbivores like

the sambar ( Cervus unicolor ), chital (Axis axis),

Asian elephant ( Elephas maximus ) and

carnivores such as leopard ( Panthera pardus ),

wild dog (i Cuon alpinus ) and sloth bear ( Melursus

ursinus). Apart from these, smaller mammals

such as black-naped hare ( Lepus nigricollis),

common palm civet (Paradoxurus

hermaphroditus) and the jungle cat (Felis chaus)

were also sighted (Table 2).

Data on wild pigs only, the major pest on

paddy crop, was considered for analysis. The

frequency of wild pig raids in the ten study

villages was proportional to the area of land

under cultivation (Table 3). The mean number

of crop raids per month by wild pigs in the fenced

JOURNAL , BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

41

CROP DEPREDATION BY WILDLIFE

Table 2: Wildlife raids on croplands in 1998-99 in the ten study villages

IUCN = International Union for Conservation of Nature and Natural Resources, F = Fenced villages,

UF = Unfenced villages, EN = Endangered, LR/CD = Low risk/ conservation dependent, VU = Vulnerable,

DD = Data deficient

villages was 2 1 .80 and 22.54 in unfenced villages

(Table 4). The mean group size was about 11

animals in each category (Table 4). The crop loss

between the fenced and unfenced villages was

not significantly different (Table 4).

The regression between the number of

wild pigs and damaged area was not significant

(r2 = 0.09), suggesting that the extent of damage

was not dependent on the number of pigs involved

in the raid.

Economic value of crop loss

The overall crop damage in all the villages

studied was estimated to be Rs. 16,270.65, at

Rs. 4.40 per kg of paddy (Ministry of Agriculture,

Govt, of India) in all the ten villages during the

study. The approximate loss of paddy to wild pigs

was 7% of the actual produce in all the villages

(Fig. 2). Potential and actual yield were estimated

at 5270.29 kg ha*1 and 3697.93 kg ha*1

respectively. The potential loss was estimated at

366.56 kg ha-1 and the actual loss was 257.19

kg ha'1 (Fig. 2).

Effects of the electric fence on crop loss

There were frequent large mammal

sightings in the unfenced villages compared to

the fenced villages (Table 2). The mean quantity

Table 3: Sampled area and estimated crop loss per village

42

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY 1 00(1), APR. 2003

CROP DEPREDATION BY WILDLIFE

of paddy lost per pig raid in fenced villages

was 9.48 kg, with Pothigaiadi recording a

maximum of 14.376 kg/raid. In the unfenced

villages, the mean loss was estimated at

9.65 kg/raid, with Manjuvelai recording the

highest at 14.47 kg/raid (Table 3).

There was no significant difference

(P = 0.05) in the loss estimates, raiding frequency,

and wild pig group size between the fenced and

unfenced villages during the study period

(Table 4).

Discussion

The study indicated that the wild pig was

the major crop pest along the eastern boundary

of the Kalakad-Mundanthurai Tiger Reserve. The

wild pig causes significant damage to paddy, the

major crop in the study area, accounting for about

7% of the actual yield. The extent of loss might

vary, depending on the season and the year. The

crop is damaged as the wild pigs wallow in the

abundant mud and water in paddy fields. They

consume the grain at only one stage, the ‘milk

maturity stage’; otherwise they do not eat any

part of the paddy plant (Ahmed 1991, Jeyasingh

1999).

The Indian wild pig is a forest loving

omnivorous mammal. Its diet includes roots,

tubers, bulbs, fruit, insects, molluscs and remains

of tiger and wild dog kills (Prater 1980, Tisdell

1982, Seshadri 1986). As it does not have sweat

glands, the wild pig must drink regularly and

wallow to regulate body temperature (Ahmed

1991). Therefore, it requires water sources,

especially during the hot season. It prefers to

remain in the shade of reeds and shrubs, which

help in thermal regulation, and prefers open

canopy and dense undergrowth at night (Tisdell

1982). It is crepuscular, although in areas where

human interference is high, it is known to

become nocturnal (Prater 1980). The home

range of a sow tends to be 5-30 sq. km and that

of a boar about 50 sq. km. The boar is mobile

Fig. 2: Schematic representation of loss calculations

and is known to move long distances for

food and mating opportunities (Prater 1980,

Tisdell and Fadeer 1981). Group sizes vary with

climatic conditions and a sounder of 10-15

individuals is common (Brander 1923, Prater

1980).

Until the 1 960s, forests adjoining the study

villages were disturbed periodically: clear cutting

deciduous forests and planting commercially

important species, construction of dams, intense

cattle grazing and frequent fires (Joshua and

Johnsingh 1989). As a result, plant species

composition is dominated by fire resistant and

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

43

CROP DEPREDATION BY WILDLIFE

Table 4: Difference between raid frequency, group size of wild pigs and crop loss in fenced and unfenced villages

*-not significant

highly silicified species (Johnsingh 1986).

Moreover, the two major reservoirs within the

Reserve, Karaiyar and Servalar may desiccate

all potential wild pig wallows along the rivers

inside the Reserve. The non-significant

regression between number of pigs and extent

of damage suggests that the extent of damage

might be a factor of time spent in the cropland,

suggesting that wild pigs might prefer the paddy

fields mainly for the ambience rather than for

forage.

The 8.6 km long electric fence erected with

World Bank aid in 1996 does not appear to be

effective against the major crop pest. There was

no significant difference in raiding patterns of

wild pigs and crop loss between the fenced and

unfenced portions of the Kalakad-Mundanthurai

Tiger Reserve’s boundary. It is apparent that

the fence was designed to deter larger mammals

such as the elephants, sambar and chital. This

might be because the fence is designed in such

a manner that there is a gap of 40 cm

(approximate to 10 different places in the fence)

between the lowest live wire (first wire) and the

next live wire (third wire), making it possible

for smaller animals to slip through. Moreover,

the second wire at about 30 cm from the ground

is an earth wire, which might enable the larger

pigs to penetrate the fence without getting an

electric shock. We suggest that the gap between

the three lower wires be reduced to eliminate

penetration by wild pigs. The scrub dominated

hills and private irrigation canals along the

border are refuges for the sounders of wild pig

once they are outside the fence.

It is evident that the fence has not been

designed to control the wild pig. Preliminary

studies must be carried out before a control or

management project is conducted. The success

of such a venture depends on its effectiveness in

reducing crop damage by wildlife. The project

should be monitored to assess its effectiveness.

If crop damage continues despite such a project,

it may create distrust among villagers towards

the Forest Department, reducing their goodwill

towards conservation efforts in the area. Human

use of the landscape is a reality and must be

dealt with in reserve design (Kramer et al. 1 997),

the importance of people in the success of

conservation schemes has been stressed in both

developed and undeveloped countries (McNeeley

and Norgaard 1992, Kothari et al. 1996). It is

suggested that the wild pig raids reported in this

study be controlled as soon as possible, before

local villagers completely lose faith in the forest

department. It is further recommended that

suitable habitats such as wallows be created

within the Reserve for wild pigs, and wild pig

incidence in cropland be tested after such

environmental control measures.

Crop loss to wildlife in a country like India

is a bane for conservation efforts, where farmers

compete with wildlife for space and resources.

Quantifying crop loss and identifying the cues

J' y

for wildlife to raid crops are vital in developing

efficient conservation strategies. Correct

management and control measures are needed

to ameliorate the economic loss incurred due to

wildlife, and to cultivate conservation awareness

among local communities.

44

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY 100(1), APR. 2003

CROP DEPREDATION BY WILDLIFE

Acknowledgements

This study was funded by a grant from the

MacArthur Foundation to the University of

Massachusetts-Boston and Pondicherry

University. The authors wish to thank Dr. K.S.

Refer

Ahmed, B.H.M. (1991): Man and wild boar, Sus scrofa

cristatus (Wagner) interaction from the Western

Ghats region of South Maharashtra. Ph.D. thesis,

Shivaji University, Kolhapur. 183 pp.

Brander, D.A.A (1923): Wild Animals in Central India.

1st Indian edition, 1982. Natraj Publishers, Dehra

Dun, India. 296 pp.

Gradwohl, J. & R. Greenberg (1988): Saving the Tropical

Forest. Earthscan Ltu. London. 214 pp.

JeyasingFT, P.D. (1999): Crop depredation by the Indian

wild pig Sus scrofa cristatus at fenced and unfenced

areas on the eastern boundary of the Kalakad-

Mundanthurai Tiger Reserve. M.S. thesis,

Pondicherry University, Pondicherry, India. 46 pp.

Johnsingh, A.J.T. (1986): Impact of fire on wildlife ecology

in two dry deciduous forests in south India. Indian

For. 112(10 ): 933-938.

Joshua, J. & A.J.T. Johnsingh (1989): Threatened gallery

forest of river Tambiraparani, Mundanthurai

Wildlife Sanctuary, South India. Biol. Conser. 47:

273-280.

Kothari, A.,N. Singh & S. Suri (Eds) (1996): People and

Protected Areas: Toward Participatory Conservation

in India. Sage Publications, New Delhi. 276 pp.

Kramer, R., C. van Shaik & J. Johnson (Eds) ( 1 997): Last

Stand: Protected Areas and Defense of Tropical

Biodiversity. Oxford University Press, New York.

242 pp.

McNeeley, J.A. & R.B. Norgaard (1992): Developed

country policies and biological diversity in

developing countries. Arg. Ecosyst. Environ. 42(1-

Bawa for interest in the project. The Forest

Department of Tamil Nadu, in particular the

office of the Field Director of Kalakad-

Mundanthurai Tiger Reserve, which provided

facilities and logistical support and Dr. M.

Sankaran for his input and support.

ENCES

2): 194-204.

Prater, S.H. (1980): The Book of Indian Animals. Bombay

Natural History Society, Bombay. 324 pp.

Sanyal, P. (1987): Managing the man-eaters in the

Sunderbans Tiger Reserve of India — A case study.

In: Tigers of the world: The biology, biopolitics,

management and conservation of an endangered

species (Eds: Nelson, R.D. and U.S. Seal).

Noyes Publication, Park Ridge, New Jersey.

Pp. 427-434

Seshadrj, B. (1986): India’s Wildlife and Wildlife Reserves.

Sterling Publishers, New Delhi. 241 pp.

Sukumar, R. (1989): Ecology of the Asian elephant in

Southern India II. Feeding habits and crop raiding

patterns. J. Trop. Ecol. 6: 33-53.

Sukumar, R. ( 1 99 1 ): The management of large mammals

in relation to male strategies and conflict with

people. Biol. Conser. 55: 93-102.

Sukumar, R. (1992): The Asian Elephant. Its ecology and

management. Cambridge University Press, London.

Pp. 251.

Tisdell, C.A. & E.V. Fadeer (1981): Dynamics of the

northern limits of the area of distribution of the wild

boar, Sus scrofa in eastern Europe. Biol. Nauk

( Mosc ). 10(9): 56-64.

Tisdell, C.A. (1982): Wild pigs: Environmental pest or

economic resource? Pergamon Press, London.

445 pp.

Western, D. & M. Pearl (Eds) (1989): Conservation for

the Twenty-First Century. Oxford University Press,

New York. 365 pp.

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

45

SMALL MAMMALS IN MONTANE ECOSYSTEMS OF THE NILGIRIS,

SOUTHERN INDIA: THEIR ECOLOGY AND NATURAL HISTORY1

Kartik Shanker2

Key words: rodents, Rattus rattus , Millardia meltada, montane forest,

grassland, plantation

Small mammals were studied in the montane ecosystems of the Nilgiris in the Western Ghats,

southern India, from February 1994 to September 1996. A total of 35,000 trap-nights were sampled

in various habitats including montane forests, grasslands, and man-made habitats such as

Eucalyptus, Pinus and Acacia plantations. A total of nine species were trapped in the montane

forest patches and three to four species in each of the other habitats. Rattus rattus was dominant

in the montane forests, while Millardia meltada was dominant in the grasslands. Both species

were found in plantations. The occurrence of M. meltada in the high altitude grassland is

remarkable, as it is not found in such habitats elsewhere, nor is it found in intermediate habitats

in the Nilgiris. Species richness and abundance of small mammals was high compared to other

natural habitats in southern India. While various aspects of the demography, habitat utilisation

and community structure were studied, many others, regarding small mammal population dynamics,

which are pertinent to their conservation and that of their predators, still need to be addressed

with extensive field studies. Though this particular field is in its infancy in India, it is hoped that

this and other such studies will pave the way for more such work in the future.

Introduction

Small mammals have been studied

extensively, including aspects of life history (see

Fleming, 1979 for review) demography (Fleming

1975), insularity (Gliwicz 1980; Adler and

Levins 1994), desert communities (Kotler and

Brown 1988; Brown 1989), competition (Grant

1972) and habitat utilisation (August 1983).

However, they have been largely ignored in India

except in taxonomic accounts (Agrawal and

Ghose 1969; Biswas and Tiwari 1966;

Chaturvedi 1966; Ellerman 1961; Ellerman and

Morris-Scott 1951; Ghose 1964), surveys and

pest management studies in agricultural areas

(Barnett and Prakash 1975; Sood and Guraya

1976; Prakash 1976, 1988; Prakash and Ghosh

‘Accepted December, 2002

2Centre for Ecological Sciences,

Indian Institute of Science,

Bangalore 560 012, Karnataka, India.

Present Address: H-Vl/2, Habib Complex,

Durgabai Deshmukh Road, RA Puram,

Chennai 600 028, Tamil Nadu, India.

Email: [email protected]

1992; Karim 1994). The lack of concern for

murid rodents in particular is perhaps reflected

in the Indian Wildlife (Protection) Act (1972)

where this family (which includes most rodents

apart from the giant squirrels, flying squirrels

and a few others) is relegated to Schedule V

(vermin) and reduced to two entries, namely

‘rats’ and ‘mice’. There is, however, a fairly large

body of work on the distribution of rodents in

the country, largely due to numerous surveys by

the Zoological Survey of India. Notable amongst

these is the pioneering work of Dr. Ishwar

Prakash (Central Arid Zone Research Institute,

Jodhpur) which includes extensive studies on the

Indian desert gerbille Meriones hurrianae

(Prakash 1964, 1969, 1981; Prakash et al. 1965;

Prakash et al ., 1969; Prakash and Jain 1970;

Prakash and Idris 1992), ecology of rodent

communities in various ecosystems in

northwestern India (Agrawal and Prakash 1992;

Prakash 1975, 1994; Prakash and Gupta 1976;

Prakash and Rana 1973; Prakash et al. 1971;

Prakash et al. 1996, 1995) and rodents as pests

in agriculture (Barnett and Prakash 1975;

46

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

SMALL MAMMALS IN MONTANE ECOSYSTEMS OF THE NILGIR1S

Prakash 1976, 1988; Prakash and Ghosh 1992).

The Rodent Newsletter of the All India

Coordinated Research Project on Rodent Control

(Central Arid Zone Research Institute, Jodhpur)

has, since the mid 1970s, provided a forum for

issues on rodents as pests.

Apart from these efforts, however, the

available information about rodents in natural

habitats is minimal throughout most of India.

Interest in small mammal studies amongst

wildlife biologists and animal ecologists in India

gradually increased in the early 1990s. While

the latter are interested in theoretical aspects of

community ecology and population biology, the

former are more concerned about the role played

by small mammals in ecosystems, especially as

prey of small carnivores, some of which are

endangered. Chandrasekhar (1989) and

Chandrasekhar-Rao and Sunquist ( 1 996) studied

small mammal communities in various habitat

types in the Anamalais. Small mammal

communities were also examined in the context

of fragmentation of rainforests in the Western

Ghats (Mudappa, pers. comm.; Prabhakar 1998)

and in plantations in Kerala (Bhat and Sujatha

1986). Other recent ecological work includes

studies on habitat utilisation in three rodents in

sandy habitat in the Thar (Mukherjee 1 999), and

rodents as a prey base for small carnivores in

Sariska Tiger Reserve (Mukherjee 1998).

I studied small mammal (rodent and

insectivore) communities in montane (shola-

grassland) ecosystems in the Nilgiris in southern

India. The population dynamics and community

ecology of small mammals have been dealt with

extensively; I review these results briefly. Firstly,

it is clear that density estimation methods have

to be developed or modified with tropical forests

and tropical small mammal populations in mind,

especially when densities are low (Shanker

2000a). Alternately, one must focus on questions

that do not deal directly with densities, but with

other parameters that can be measured with

precision in tropical systems. In examining the

demography of small mammal populations in

shola patches, a comparison between populations

in small and large fragments showed that while

population fluctuations between fragments were

asynchronous, fluctuations within large patches

were synchronous (Shanker and Sukumar 1 999).

This requires closer examination in the context

of fragmentation in the Nilgiris and other parts

of the Western Ghats. The demography and

community structure of these small mammal

populations was related to fragment size and

habitat characteristics of the montane patches

(Shanker and Sukumar 1998). Interspecific

competition appeared to be less important than

other factors such as intraspecific competition

and predation (Shanker 2000b). Differences in

small communities in different habitats and the

significance of plantations in the landscape were

examined and metapopulation processes and

landscape effects on these populations in the

montane ecosystem were explored (Shanker, in

press).

Since small mammal community ecology

is a relatively new field of study in the tropics,

and India in particular, a great many questions

remain unanswered. In many ways, this study

threw up more questions than it answered.

I attempted to address some of these questions

with field and laboratory studies. In this paper,

I give a brief overview of the small mammal study

in the Nilgiris, with an emphasis on the natural

history and ecology of the various species. I also

present some hypotheses, and questions that

remain unanswered during the study.

Study Area

The Nilgiris are located between 11° 10* -

I I ° 30' N and 76° 25' - 77° 00’ E at the junction

of the Eastern Ghats and the Western Ghats, in

southern India. The study was conducted at

higher altitudes (1,800-2,500 m above msl) of

the Nilgiris which have a montane ecosystem,

comprising of patches of stunted evergreen forest

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

47

SMALL MAMMALS IN MONTANE ECOSYSTEMS OF THE NILGJRJS

(locally called ‘sholas’) surrounded by

grasslands. Though most grasslands have been

replaced by exotic species such as wattle (. Acacia

spp.), pine (Finns roxburghii ), tea ( Camellia

sinensis) and blue gum (Eucalyptus globulus ),

the natural ecosystem remains intact in the

southwest region of the Nilgiris plateau. The

stunted montane evergreen forest patches, usually

1-10 ha in size with few exceeding 100 ha, are

confined to depressions and folds in the

mountain. They are extremely dense, with 5000+

woody plants (>1 cm d.b.h.) per hectare, and are

dominated by the families Lauraceae, Rubiaceae

and Symplocaceae (H.S. Suresh and R. Sukumar,

unpubl. data). There is a sharply defined ecotone

between the montane forest patches and the

grasslands that are maintained by frost and fire

(Meher-Homji 1984), The climate is highly

seasonal, with a dry season extending from

December to February, a pre-monsoon season

from March to May, the southwest monsoon fr om

June to August and a second wet season from

September to November. Most of the areas

sampled receive annual rainfall of 1,500-

2,000 mm.

Methods

Sampling Procedures

Live trapping of small mammals was

carried out between February 1994 and

September 1996. Montane forest patches and

grasslands were sampled using 0.49 ha and 1 ha

plots. Traps were placed at intervals of 10 m so

that each 0.49 ha plot consisted of

49 permanently marked trap stations, in a square

grid of 7 x 7 traps (the 1 ha plots had 100 trap

stations, in a 10 x 10 configuration). Due to their

size and shape, plantations were sampled using

0.45 ha plots (15x3 trap stations). A standard

Sherman live trap (22.9 cm x 7.6 cm x 8.9 cm)

was placed on the ground at each station, close

to a tree, log, or any other appropriate runway.

The traps were baited with grated coconut and

rice. All trapped animals were identified,

uniquely marked (Ear-punch, National Band and

Tag. Co., Kentucky # 1538), sexed, weighed,

measured and released. The traps were checked

once daily between 0800 hrs and 1200 hrs. The

plots were run for five consecutive nights during

February-October 1994. Trap mortality was

found to be particularly high towards the end of

the trapping period in some seasons, and the

trapping duration was thus reduced to three

nights for the rest of the study (Shanker 1998).

Six montane evergreen forest patches,

ranging from 2 to 600 ha, were extensively

sampled using nine 0.49 ha plots and two 1 ha

plots (see Shanker 1998; Shanker and Sukumar

1998). The sampling was primarily carried out

in the Upper Bhavani region of the Mukurthi

National Park and at Thaishola, about 20 km

east of Upper Bhavani (which is about 65 km

southwest of Udhagamandalam). Thaishola is the

largest shola in the Nilgiris, (c. 600 ha) and the

other sholas sampled ranged from 2 to 60 ha.

Grasslands were also sampled using 0.49 ha

plots. Both forest patches and grasslands were

also sampled with 0.45 ha plots, which enabled

comparisons with plots in anthropogenic habitats

which had a similar design. Anthropogenic

habitats - tea, gum, wattle and pine - were

sampled using 0.45 ha plots, including wattle

plantations of three different ages (Table 1).

Additionally, a 0.90 ha plot was established in

an old wattle plantation to study the interaction

between Rattus rattus and Millardia meltada.

Each plot was sampled several times, typically

once during each season (= ‘session’).

Data Analysis

Various estimators were attempted using

capture-mark-recapture models, but these could

not be applied consistently to all species across

all seasons (Shanker 2000a). Hence, we used the

minimum number alive (Krebs 1966), which is

the total number of animals of a particular species

trapped during a trapping session.

48

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 10.0(1), APR. 2003

SMALL MAMMA LS IN MONTANE ECOSYSTEMS OF THE NILGIR1S

Table 1 : Trapping effort and habitat characteristics of the habitat types sampled

in the Upper Nilgiris and Mudumalai Wildlife Sanctuary

Results

In all, 1,310 individuals were trapped in

the sholas, grasslands and plantations with an

overall trapping success of 10.6%. A total of

9 species were trapped, there were 8 species in

montane evergreen forests, 3 species in the

grassland, 3 species in wattle, eucalyptus and

pine plantations, and 4 species in tea plantations

(Table 2). Rattus rattus was the dominant species

in the montane forest and comprised 60.9% of

the total density, while Miliardia meltada was

the dominant species in the grassland. In the

natural habitat (forest and grassland), the two

species are mutually exclusive. While the former

was trapped in the forest patches, the latter was

the only rodent trapped in the grassland. An

analysis of wattle stands of different ages shows

that Miliardia dominates young stands (grass-

like habitat), while Rattus dominates the older

Table 2: Average density (MNA/ha) of rodents and shrews in the various habitats in Upper Nilgiris

Species/ Thai- Other Grass- Wattle Wattle Wattle Blue Tea Pine

Habitat shola Sholas land young mid old Gum

Rodents

Rattus rattus

R .r. rufescens

Cremnomys blanfordi

Mus famulus

Mus platythrix

Platacanthomys lasiurus

Miliardia meltada

Vandeleuria oleracea

Funambulus sublineatus

Ratufa indica

Shrews

Suncus montan us

Suncus dayi

Feroculus feroculus

*

Thaishola is the largest shola in the Nilgiris at 600 ha. S - Single capture, R - rare species i.e. caught more than once,

but only on a few occasions, L - low density i.e. caught during many trapping sessions, but usually only one or two

captures during a session (density < 1 /ha), V - visual sighting, * - incidental capture

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

49

SMALL MAMMALS IN MONTANE ECOSYSTEMS OF THE NILGIRIS

stands (forest-like habitat). Suncus montanus, a

shrew, was the only species captured in both

habitats. The other common species in the

montane forest included Mus famulus and Suncus

dayi.

The total density and biomass of small

mammals were highest in the old and middle-

aged wattle stands. While Rattus rattus

accounted for the bulk of the density and biomass

in the old stands, Millardia meltada was the

dominant species in the middle-aged stands.

Montane forests had intermediate densities and

biomass, while grasslands had relatively low

density and biomass. The density and biomass

of the dominant species were substantially higher

in some plantations than in the natural habitats.

The density of Millardia meltada was

significantly higher in middle-aged wattle stands

than in grasslands, while the density of Rattus

rattus was significantly higher in old wattle

plantations than in montane forest patches.

R. rattus attained a maximum density of

34 animals/ha and biomass of 3.3 kg/ha in

November 1995 in an old wattle plantation.

M. meltada peaked at 31 animals/ha (1.6 kg/ha)

in June 1995 in a middle-aged wattle plantation.

Seven species of rodents (Order Rodentia)

and two shrews (Order Insectivora) were

captured during the study period (Table 2). Two

other species of rodents were sighted, but not

trapped. These included the dusky striped

squirrel Funambulus sublineatus, and the Indian

giant squirrel Ratufa indica. The latter is a much

larger species and is not relevant to the study.

Kelaarf s long-clawed shrew Feroculus feroculus

Kelaart, was caught once during preliminary

trapping. This marks an extension of its territory

from its previously known range (Pradhan et al.

1997). The small mammals considered here

ranged in size from 6 to 206 gm, and could in

theory have weighed up to about 300 gm, based

on the size of the traps used. The suborder

including rats and mice is the most widespread

and has the largest number of species. Compared

with other rodents, members of the Family

Muridae (true mice) may have evolved most

recently. One hypothesis suggests that murids

evolved in southern Asia, as the earliest fossils

of murids have been found in Pakistan. This

would have been followed by adaptive radiation

to the other continents.

Rattus rattus (Linnaeus) (Muridae)

The most common and well known of

murid rodents is the common rat Rattus rattus ,

also known as the black, roof, house or ship rat.

R. rattus is found worldwide, and owes much of

its ubiquity to accidental human introduction.

While the brown rat Rattus norvegicus , has

successfully colonised temperate countries, the

black rat has invaded and is widespread in most

tropical countries, including India. Ellerman

(1961) divides the black rats broadly into two

categories, the white-bellied form, which is

usually found in the wild and the dark-bellied

form, which is usually commensal with man; he

further sub-divides them into 16 subspecies. The

white-bellied form, which was trapped in many

habitats in our study, was identified as Rattus

rattus wroughtoni Hinton (Dr. M.S. Pradhan,

ZSI, Pune, pers. comm.). It was trapped in natural

habitats and in plantations. The dark-bellied

subspecies Rattus rattus rufescens , which is

usually found in agricultural fields, was trapped

in wattle plantations. Both subspecies were

trapped in the same area, raising questions about

their relative taxonomic status. However, in

another study, all white-bellied individuals

collected from the Nilgiris were identified as

Rattus satarae based on chromosomal,

morphological and isozyme studies (Francois

Catzeflis, pers. comm.). There are two separate

issues here: are the different forms (subspecies)

of Rattus rattus in fact separate species, as

Catzeflis suggests? Does the Rattus rattus

trapped in the Nilgiris belong to the species/

subspecies ‘ wroughtoni' or ‘satarae'l Only

extensive molecular genetic studies can provide

50

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY 100(1), APR. 2003

SMALL MAMMALS IN MONTANE ECOSYSTEMS OF THE NILGIRJS

answers to questions surrounding the taxonomy

of the various forms of Rattus rattus found all

over India.

Rattus rattus is highly arboreal in the wild.

The females are territorial, while the males

appear to be free ranging. Each female occupies

a nest on her own and has 3-4 pups. The adult is

about 15-20 cm long with a tail that is 20-25 cm

long; large males weigh about 150 gm, while

adult females weigh 100-120 gm. Though males

rarely weighed above 1 50 gm, a single individual

weighing 206 gm was trapped once during the

study. Interestingly, the white-bellied form of

Rattus rattus (wrought onil) was also dominant

in lowland evergreen forests in the Nilgiris

(Meena 1997), Anamalais (Chandrasekhar 1989;

Prabhakar 1998) and Kalakkad (Divya Mudappa,

pers. comm.).

Millardia meltada Gray (Muridae)

Millardia meltada , the soft-furred field rat,

found in cultivated fields throughout India

(Prater 1988), has adapted successfully to the

grasslands of the Upper Nilgiris. The general

colour is pale brownish-gray, grayish-white on

the underside. It has large rounded ears and a

hairy tail, has a head body length of 13-15 cm, a

tail nearly as long, and weighs 50 to 70 gm. This

rat is commonly found in agricultural land and

lowland plantations and is a major pest in many

areas in north, central and southern India.

In natural habitat that was sampled in

various studies in southern India, M. meltada

was not trapped in the mid elevation forests or

plantations in Kalakkad-Mundanthurai or

Anamalais and in the mid elevation forests and

grasslands in Mudumalai. Its occurrence in the

high altitude grasslands of the Nilgiris is

therefore a matter of some interest. It is also

notable that it was not found in high elevation

grasslands in Aravalli hills, though it was

abundant at lower elevations (Prakash et al. ,

1995). It is possible that this species colonized

the Nilgiris grassland before the advent of man

about 1 000 years ago, and subsequently occupied

the plantations about 200 years ago.

Alternatively, it may have arrived as a commensal

with man and colonized the grasslands

subsequently. While there has been a long

standing debate on the origin of the grasslands

in the Nilgiris (Bor 1938; Ranganathan 1938)

recent studies indicate that it is a climax

ecosystem (Sukumar et al. 1993; Rajagopalan

et al. 1997). It can therefore be expected that

some rodent species would have occupied this

habitat. Whether Millardia meltada is the

original colonizer or a more recent one that

arrived as a commensal can only be ascertained

by paleontological or molecular genetic evidence.

Weight of animals in captivity

Fifteen individuals of Millardia meltada

were kept in captivity. Within three to seven days

of capture, 4 individuals gained weight at an

average of 9.3% of their body weight,

5 individuals lost weight (7.8%), and 3 showed

no change in weight.

Of 1 7 individuals of Rattus rattus that were

kept in captivity, 15 (88.2%) showed an increase

in body weight within three to seven days. Eleven

of these showed an average increase of 3.14%

per day within the first week, which would lead

to an increase of about 20% within a week. The

maximum increase was 40% of the body weight

at capture in five days.

Mus spp. (Muridae)

Bonhote’s field mouse Mus famulus

Bonhote is a small animal with a brown coat

and a yellow underside. It is found at 1,507 m

and above in the Nilgiris, and a few other hills

in the southern Western Ghats. It is 5-8 cm long

and weighs about 20 gm, and is probably less

arboreal than the other rats and mice. It was

found in areas of higher tree density in the forest

patches (Shanker 2000b). The spiny field mouse

Mus platythrix Bennett a species common all

over India, is brownish above and white below.

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

51

SMA LL MAMMALS IN MONTANE ECOSYSTEMS OF THE NILGIRIS

Its fur is composed of flattened spines (Prater

1988). It was trapped in the two largest sholas

in the first year of the study and was not trapped

subsequently. Some information on the ecology

of Mus platythrix and Mus booduga (Chandrahas

1974) is available; the brood of the latter has

even been found in abandoned baya nests

(Kichtar and Tiwari 1992)

Platacanthomys lasiurus (Muscardinidae) and

other rare rodents (Muridae)

Platacanthomys lasiurus Blyth is light

rufous brown above and a dull white below. The

muzzle is pointed, the ears are thin and naked,

and the hind feet are broad and elongated. It is

13-20 cm long and weighs about 70 gm. The

spiny dormouse inhabits rocky hills and forests

at altitudes of 600 m and above. It lives mainly

in the cavities of trunks and branches, and in

clefts in rocks. The long tail, covered with hair,

serves as a balancing organ for this highly

arboreal animal. This species appeared after

several months of trapping in the two largest

sholas, Thaishola and the 60 ha shola. It was

trapped only from September to January. There

is little information on the distribution of this

species, however, a number of new locales have

recently been documented (Rajagopalan 1968;

Jayson and Christopher 1995; Prabhakar 1997).

This is the only endemic small rodent in the

Western Ghats, the other endemic rodents being

the grizzled giant squirrel ( Ratufa macroura ) and

a flying squirrel ( Petinomys fascocapillus).

The white-tailed wood rat Cremnomys

blanfordi Thomas is very similar to the common

rat in appearance, except for the tail, which is

brown for three quarters of its length, but white

towards the tip. It is highly arboreal in forests,

makes a large and untidy nest, and has a litter of

2-3 young. This species was represented by a

single capture in Thaishola and is probably not

common at higher altitudes. It was dominant in

lower elevation Deciduous Forests in Mudumalai

in the Nilgiris (Meena 1997).

The long tailed tree mouse Vandeleuria

oleracea Bennett can be distinguished by the fact

that the first and fifth toe on all four feet are

partially opposable and have a flat nail instead

of a claw. It is about 7 cm in length, with a tail

that is slightly longer. It is an extremely attractive

creature, with a reddish coat and white

underparts. It was trapped only once during two

years of trapping in the Upper Nilgiris.

Suncus spp. (Order Insectivora, Family

Soricidae)

Suncus murinus (Linnaeus), the common

or grey musk shrew, is found all over India in all

kinds of habitats, and is common in cities as well.

It is about 10-12 cm long with a slightly shorter

tail, and weighs about 20 gm. In the Nilgiris,

Suncus montanus Kelaart, a similar looking

shrew, is more common. Suncus dayi Dobson, a

much smaller shrew, weighing just 6 to 10 gm,

is also found in the Nilgiris. Shrews are highly

aggressive animals and on occasions, when two

shrews were caught in the same trap, one would

devour the other. On one occasion, a shrew and

a common rat ( Rattus rattus ) were released

simultaneously from a trap, whereupon the shrew

attacked the weakened rat and chewed off its ear.

Shrews have been known to attack rats, and

Prater (1988) suggests that presence of shrews

in houses may deter rats. Saini and Parshad

(1996) report the consumption of a gerbil by a

shrew in a multi-catch trap. Shrews tend to die

in traps as they have a high metabolic rate; also

in this case, the food in the trap was usually

vegetable matter such as coconut. The smaller

shrew, Suncus dayi , was rarely captured live in

the trap.

Discussion

Although ‘small mammals’ do not

constitute a taxonomic entity, the term generally

includes mammals from 2 gm to 5 kg (Bourliere

1975). Of the 4,200 odd mammal species, 90%

52

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

SMALL MAMMALS IN MONTANE ECOSYSTEMS OF THE NILGIR1S

weigh less than 5 kg. Ten out of sixteen

mammalian orders contain mostly small species.

In fact, both in birds and mammals, the below

1 kg class embraces the most successful order:

of some 8,600 species of birds, 5,100 belong to

the Order Passeriformes; of the 4,200 mammal

species, about 1,700 are rodents. In India, there

are about 100 species of rodents, with about

25 species in southern India. Most studies have

recorded relatively few species of small mammals

(three to five) in each habitat (Chandrasekhar

1989; Meena 1997; Divya Mudappa, pers.

comm.). Prabhakar (1998) trapped 7 species of

rodents and 1 shrew in mid-elevation rainforests.

In comparison, the species richness in the high

elevation montane forests of the Nilgiris is high

with 6 species of rodents and 3 shrews. In

addition, the grasslands had one rodent species.

Trapping success (10.6%) was also high

compared to studies in south India, where

trapping success was 5% or lower

(Chandrasekhar 1989; Meena 1997; Divya

Mudappa, pers. comm.). A pattern that seems to

have emerged from these studies is that mid to

high elevation evergreen forest fragments support

the highest species richness and abundance of

small mammals in natural habitats in southern

India. Another clear pattern is the dominance of

the white-bellied form of Rattus rattus

( wroughtoni and/or others) in evergreen habitat

at mid and high elevations (this study;

Chandrasekhar 1989; Meena 1997; Prabhakar

1998; Divya Mudappa, pers. comm.)

There has been a long debate on the

importance of competition in structuring

ecological communities (Connell 1983; Schoener

1983). The distribution of Rattus rattus and

Millardia meltada in this ecosystem is

particularly interesting. They are completely

exclusive in the natural habitat, which is

particularly notable in the shola grassland system

where there is a sharp ecotone between the two

habitat types. Rattus rattus would be found till

the edge of the shola and Millardia meltada till

the edge of the grassland, and therefore within

metres of each other, but never in the other

habitat. Though this initially indicated some

competitive exclusion, the two species coexisted

in the plantations. The distribution of the species

may thus have been influenced more by the

habitat parameters, trees in the case of Rattus

rattus and grass in the case of Millardia meltada.

This is also supported by the fact that Millardia

meltada is dominant in young wattle stands,

which have more ground cover, while Rattus

rattus is dominant in older wattle stands with

taller trees (for details, see Shanker 2000b).

Removal and introduction experiments in the

montane forests and grasslands did not show any

competition between the species (Shanker

2000b). While the evidence for competition

between the species is low, it would be interesting

to carry out reciprocal removal experiments in

the wattle stands where the species coexist. Gut

content analysis should provide some evidence

on the feeding habits of the two species. Since

one is a grassland species and the other a forest

species, another method of studying differences

in foraging would be to look at stable carbon

isotopes in bone collagen (DeNiro and Epstein

1978; Sukumar and Ramesh 1992). Food choice

experiments would also shed light on their

feeding habits.

Small mammals such as rodents are

considered to be especially important components

of the ecosystem as they serve as prey for small

and medium sized carnivores In the Upper

Nilgiris, potential predators include several birds

such as raptors, owls and crows and several

mammals such as jungle cat, leopard cat, small

Indian civet, ruddy mongoose, jackal and Nilgiri

marten (listed in Shanker and Sukumar 1999).

Study of the demography of small rodents in the

montane patches revealed some interesting

patterns. The population of Rattus rattus in the

smaller fragments was asynchronous, which may

be due to predation and demographic

stochasticity (Shanker and Sukumar 1999).

JOURNAL BOMBAY NATURAL HISTORY SOCIETY 100(1). APR. 2003

53

SMALL MAMMALS IN MONTANE ECOSYSTEMS OF THE NILGIR1S

Further, population characteristics of small

mammals were affected by patch size (Shanker

and Sukumar 1998). Apart from Rattus rattus

and Suncus montanus, the other species were not

trapped during all trapping sessions in the forest

plots. This indicates that some of these species

might periodically go extinct in the smaller

patches. These would thus be a good system for

the study of metapopulation biology, especially

in the context of the persistence of rare species

in the patch network. The shola-grassland and

shola-plantation habitats also provide different

kinds of landscapes to study the effects of edge

permeability and habitat connectivity on small

mammal populations. The results show that

plantations, especially old stands, have high

small mammal densities. This could affect the

population dynamics of small mammals in the

forest patches. These factors need to be taken

into account in the management of these areas,

and may be important factors to consider in the

conservation of predators that depend on these

small mammals for food.

The theory of island biogeography

(MacArthur and Wilson 1967) was extended to

isolated habitats on land with a study of mammals

on mountaintops (Brown 1971). Since there are

certain patterns in insular small mammal

populations in the Nilgiris, similar patterns may

exist in other montane ecosystems in the Western

Ghats. Other studies on fragmentation in southern

India have found effects of insularity and patch

size on small mammal communities in lower

elevation evergreen forests (Prabhakar 1998). It is

possible that the patterns observed in the Nilgiris

may be generalised to montane systems in the

Western Ghats. It would also be interesting to look

at the montane systems of various ranges as islands

and examine the impact of isolation on the small

mammal communities of ‘mountain tops’. One

could also compare the distribution of Rattus rattus

and Millardia meltada in the Nilgiris with other

montane ecosystems.

The relationship between population

synchrony and geographical distance was not

clear at smaller spatial scales. One of the ways

to study population structure is through

molecular genetic analysis. Populations of Rattus

rattus from several montane forest patches and

Millardia meltada from grasslands in the same

areas were studied using multi locus minisatellite

DNA fingerprinting. Patterns of inter-individual

and interspecific variation in these rodent

populations were examined. Inter-individual

variation in Rattus rattus was found, which could

be used in population genetic analysis (Kartik

Shanker, Anindya Sinha and Trupta Purohit,

unpubl. Data). More data is required before

patterns of variation and population structure can

be discerned. Currently, molecular tools such as

microsatellite analysis are widely used for

population genetic studies and can be used to

answer these questions.

It is clear that much work remains to be

done on small mammals in tropical ecosystems.

The montane ecosystems of the Western Ghats

offer a fascinating landscape where ecological

hypotheses of interest can be tested. It is hoped

that future generations of Indian ecologists will

address some of these questions.

Acknowledgements

The study was supported by the Ministry

of Environment and Forests, Govt, of India, and

the John D. and Catherine T. MacArthur

Foundation, USA. I thank the Tamil Nadu Forest

Department and Electricity Board for their

assistance. I am grateful to my Ph.D. supervisor,

Dr. R. Sukumar for his support, to my assistant,

Jalendran and particularly to P. Mayavan for his

assistance and companionship in the field.

54

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SMALL MAMMALS IN MONTANE ECOSYSTEMS OF THE NILGIRIS

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JOURNAL , BOMBAY NATURAL HISTORY SOCIETY. 100(1), APR. 2003

57

THE EFFECTS OF CATTLE GRAZING AND HABITAT ON HELMINTH LOADS

OF CHITAL {AXIS AXIS) IN THE MUDUMALAI WILDLIFE SANCTUARY,

SOUTHERN INDIA1

Guha Dharmarajan2, M. Raman3 and Mathew C. John4

Key words: cattle, chital, Axis axis, helminth loads, habitat

Coprological estimation of helminth eggs per gram of faeces (epg) was used as an index to study

how habitat and sympatric grazing by cattle affect helminth loads in chital ( Axis axis ) at the

Mudumalai Wildlife Sanctuary, Tamil Nadu. Two distinct groups of chital were sampled,

fortnightly, in Dry Deciduous forest (DDF) and Scrub forest (SF), one in areas with sympatric

cattle grazing and the other in areas without cattle. The median helminth load (MHL) of chital in

DDF without cattle (2 epg) and SF without cattle (1 epg) showed no significant difference (p>0.05).

However, the MHL of chital in DDF with cattle grazing (2 epg) and SF with cattle grazing

(4 epg) differed significantly (p<0.05). Chital in DDF with or without sympatric cattle had MHLs

of 2 epg (p>0.05). However, the MHL of chital in SF with sympatric cattle grazing (4 epg) and

without cattle grazing (1 epg) showed a significant difference (p<0.05). It is hypothesized that in

nutrient-poor scrub areas where chital are further subjected to resource competition by cattle,

nutrient levels may be lowered, making chital increasingly susceptible to either their own parasites

or to those of cattle.

Introduction

In India there is increasing human pressure

on protected areas, one manifestation is the

growing domestic-wild animal interface and its

resultant — increased chances of interspecific

transmission of disease. Endangered wildlife

populations are usually protected from virulent

pathogens because they are usually well below

the threshold population size (HT) (Lyles and

Dobson 1 993). However, this may not be the case

when wild and domestic animals share both

pasture and pathogens. In this study, we

examined the effects of both cattle and habitat

upon parasitic worm loads in chital {Axis axis).

We used a quantitative estimate of the number

of propagules (eggs or larvae) in the host faeces

as an index of parasite loads in the host. Egg

'Accepted July, 2002

2Centre for Ecological Science, Indian Institute of Science

Bangalore 560 012, Karnataka, India.

Email: [email protected]

3 Dept, of Parasitology, Madras Veterinary College,

Tamil Nadu Veterinary and Animal Sciences University,

Chennai 600 007, Tamil Nadu, India.

4 Dept, of Wildlife Science, Madras Veterinary College,

Tamil Nadu Veterinary and Animal Sciences University,

Chennai 600 007, Tamil Nadu, India.

output is considered to be an accurate indicator

of parasite biomass because of its positive

correlation with worm size and number

(Skorping et al. 1991). Though this method has

limitations (Foreyt and Trainer 1 980), being non-

invasive it is useful in wildlife.

Study Area

Mudumalai Wildlife Sanctuary and

National Park, and Sigur Reserve Forest (11° 32-

1 1° 93’ N and 76° 22'- 76° 43' E) have elevations

between 900-1000 m above msl. A high diversity

of vegetation types has been observed (Sukumar

et al. 1992). There is a rainfall gradient from the

western side (1,800 mm/year) which is

characterized by Moist and Dry Deciduous Forests

(DDF) to the eastern side (600 mm/year) composed

mainly of Dry Thorn or Scrub Forest (SF).

Material and Methods

Hosts sampled

Cattle and chital were sampled between

0700 and 0900 hrs at fortnightly intervals, from

May to late July, 1999. As the maximum linear

58

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

EFFECTS OF CA TTLE GRAZING AND HA BITA T ON HELMINTH LOA DS OF CHITA L

distance between sightings of individual chital

is only 2 km (Barette 1991) they could be divided

into non-overlapping groups:

1. Chital in Dry Deciduous Forest areas

with sympatric cattle grazing.

2. Chital in Dry Deciduous Forest areas

without sympatric cattle grazing.

3. Chital in Scrub Forest areas with

sympatric cattle grazing.

4. Chital in Scrub Forest areas without

sympatric cattle grazing.

Since the same cattle grazed in both Dry

Deciduous and Scrub Forest areas, samples

collected from cattle in different habitat types

had to be pooled.

Coproiogical study

A representative sample (approx. 2 gm)

was collected from distinct, fresh dung piles in

labeled, pre-weighed containers with 10 ml of

10% formalin. The exact weight of faeces was

calculated by subtracting the weight of the

container and formalin from the final weight.

The intensity of helminth infection was

determined by the quantitative Sedimentation-

Flotation Technique of Watve (1992).

Terminology

Helminth loads are expressed in terms of

eggs per gram of faeces (epg). We use the term

to include Sungworm larvae. Due to the

methodology used, definitions of some terms

differ from Margolis et al. (1982). ‘Prevalence’

indicates the percentage of samples positive for

helminth eggs or larvae. ‘Sympatric’ is defined

as: (of biological speciation or species) taking

place or existing in the same or overlapping

geographic areas (Hanks 1979).

Statistical Analyses

Calculation of index of dispersion,

d-statistic and fitting of the negative binomial

distribution has been done as per Ludwig and

Reynolds (1988). Green’s index was calculated

as per Green (1966). The Mann-Whitney Test

was performed as per Conover (1971).

Result and Discussion

The negative binomial distribution gave

good fits to observed data in cattle and all four

sub-populations of chital (Table 1 ). The d-statistic

being > 1.96 (Table 1), a clumped parasite

distribution is hypothesized (Ludwig and

Reynolds 1988), implying that a few hosts

harbour many helminths, while many hosts have

few or none (Waid et al. 1985).

Comparison of the median helminth loads

between the four groups of chital are reported in

Table 2. The median was used in preference to

the mean since the data are non-normal ly

distributed. The median helminth loads of chital

in Dry Deciduous Forest areas without cattle

grazing (2 epg) and Scrub Forest without cattle

grazing (1 epg) showed no significant difference

(p>0.05). Similarly comparison of chital in Dry

Deciduous areas with sympatric cattle grazing

and without cattle grazing showed that both had

median helminth loads of 2 epg (p>0.05).

However, the median helminth loads of chital in

Dry Deciduous Forest areas with cattle grazing

(2 epg) and Scrub Forest with cattle grazing (4

epg) differed significantly (p<0.05). And, the

median helminth loads of chital in scrub forest

areas with sympatric cattle grazing (4 epg) and

those without cattle grazing (1 epg) showed a

significant difference (p<0.05). It is thus clear

that neither forest type nor cattle grazing affects

helminth loads in chital if they act independent

of each other. However, helminth loads in chital

increase dramatically when they are sympatric

with cattle in scrub forest. Since immuno-

competence is affected by nutrition (Rolston

1992; Lyles and Dobson 1993), it is likely that

chital in scrub forest with sympatric cattle

grazing have high helminth loads due to

increased competition with cattle for limited

forage resources.

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

59

Table 1 : Distribution of helminth eggs in chital sub-populations and domestic cattle at

the Mudumalai Wildlife Sanctuary, Tamil Nadu

EFFECTS OF CA TTLE GRAZING AND HABITA T ON HELMINTH LOADS OF CHITA L

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JOURNAL, BOMBAY NATURAL HISTORY SOCIETY 100(1), APR. 2003

EFFECTS OF CATTLE GRAZING AND HABITAT ON HELMINTH LOADS OF CHITA L

Chital in Scrub may get less nutritional

resources than those in the Dry Deciduous for

three reasons: differences in the primary

productivity of the two areas, vegetation

differences between the habitats and the time

period of our study. In general, low-rainfall scrub

areas have a lower primary productivity than the

moderate to high rainfall Dry Deciduous Forests

(Sukumar 1992). This is because of an almost

linear relationship between net primary

productivity and precipitation (Whittaker 1970).

Secondly, while the Dry Deciduous Forests has

higher levels of browse (Sukumar, 1992), Scrub

habitat in Mudumalai is dominated by grass with

little browse available, especially for the smaller

herbivores. But most important was the time

during which we sampled the chital populations.

The first rainy season in Mudumalai usually

begins in mid-May (N. Baskaran, pers. comm.).

However, in 1999, the first rams were delayed to

mid-June. Thus over half our samples were

collected during the late dry season, when the

grass is likely to have the least nutritive value

(Sukumar 1990). However, grass does not always

have low nutritive value and Ekaya (2001) found

that the nutritive value was high during the short

period of flush during the growing season. This

growing season corresponds with the beginning

of the rains in Mudumalai, and the high nutritive

value of the grass is the reason why the more

mobile species of wild herbivores, like elephants,

prefer Scrub to Dry Deciduous Forests just after

the rains (Sukumar 1990). Similarly, and in light

of the arguments given pre viously, we found that

chital in the scrub forest areas with cattle grazing

began to show a significant downward trend in

helminth loads as the wet season progressed,

though they still remained higher than the other

sub-groups of chital. This was likely because of

improved nutritional status, due to increased

availability of grass with high nutritional value.

Similar downward trends were not observed for

the other sub-groups of chital or other herbivore

species examined. These results have been

reported elsewhere (Dharmarajan 2000;

Dharmarajan and John 2001).

Sympatric cattle are likely to further reduce

the availability of forage resources for chital. In

large numbers, cattle tend to overgraze and cause

outflow of nutrients from an ecosystem (Singer

and Boyce 1996). Cattle grazing can increase

the helminth loads in other ways too. First, due

to a decrease in nutrient levels, the deer may roam

larger areas and graze for longer periods to obtain

adequate nutrition, which would increase

chances of contact with infective stages of

parasites (Gordon 1948). Secondly, overgrazing

and trampling by cattle will reduce the height of

the grass and leave more open ground, thus

increasing contact with infective stages (Kauzal

1941; Gordon 1948). Thirdly, chital in the scrub

areas grazed by cattle are under great stress

(nutritional, heat, etc.) which reduces resistance

to disease (Fowler 1986).

The reduced nutritional levels of chital in

Scrub with sympatric cattle could increase

helminth loads either by increasing the

susceptibility of chital to their own parasites or

to cattle parasites. Our data shows that the latter

is more likely to be the case in Mudumalai

(Dharmarajan et al. in press.).

Another finding supporting the view that

cattle adversely affect chital from a

parasitological point of view is that poor nutrient

habitat on its own is not responsible for increased

helminth loads. We find that helminth loads of

chital in Dry Deciduous Forest without sympatric

cattle do not differ significantly from those

of chital in Scrub without sympatric cattle

(Table 2). This may be because under “natural

conditions” (i.e. absence of domestic cattle

grazing) There is a balance between host and

parasite (Gordon 1948) and in general, worm

burdens do not become intolerable. Additionally,

factors that contribute to poor nutrition of the

host, like low moisture and vegetation cover, will

adversely affect survivability of infective

helminth larvae (Stromberg 1997). In such an

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

61

EFFECTS OF CATTLE GRAZING AND HABITAT ON HELMINTH LOADS OF CHITAL

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JOURNAL, BOMBAY NATURAL HISTORY SOCIETY. 100(1), APR. 2003

SF = Scrub forest; DDF = Dry deciduous forest; epg = eggs per gram of faeces; n = Number of Samples from Population 1;

m = Number of Samples from Population 2; T1 = Mann-Whitney test statistic; ns = not significant at 5% level; * = significant at 5% level.

EFFECTS OF CATTLE GRAZING AND HABITAT ON HELMINTH LOADS OF CHITA L

unfavourable macro-environment, larval survival

is greater in dung pats compared to pellets,

because while pellets are generally dry, dung pats

contain high amount of moisture (Soulsby 1 968).

Additionally, due to the formation of a hard, dry

outer layer, the interior of dung pats rerain

moisture for long periods of time. Thus, larvae

in cattle dung pats are likely to have improved

survival as compared to those in chital pellets.

This in turn could adversely affect helminth loads

in chital, because chital under nutrient stress are

likely to be susceptible to cattle parasites

(Dharmarajan et al. in press).

In conclusion, the increased loads in chital

in scrub forest areas with sympatric cattle grazing

is probably due to the interactive effect of poor

habitat and inter-specific competition for limited

resources. The finding that cattle may have an

adverse effect on chital, from a parasitological

perspective, in resource-poor and/or degraded

habitats should be an issue of concern to

Refer

Barette, C. (1991): The size of axis deer fluid groups in

Wilpattu National Park, Sri Lanka. Mammalia

55:207-220.

Conover, W.J. (1971): Practical non-parametric statistics.

John Wiley and Sons, Inc., New York, 462 pp.

Dharmarajan, G. (2000): Epidemiology of helminth

parasites in wild and domestic herbivores at the

Mudumalai Wildlife Sanctuary, Tamil Nadu.

Masters thesis submitted to Tamil Nadu Veterinary

and Animal Sciences University, Chennai. 75 pp.

Dharmarajan, G. & M.C. John (2001): Effect of season

on helminth loads in wild herbivores in the

Mudumalai Wildlife Sanctuary, Tamil Nadu. XII

National Conference of Veterinary Parasitology,

Tirupati, Andhra Pradesh, India. August 25 -27,

2001.

Dharmarajan, G., M. Raman & M.C. John (in press.):

Effect of cattle grazing and habitat on the helminth

community structure in chital (Axis axis).

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grass and browse species from Kenya’s pastoral

ecosystems. J. Hum. Ecol. 12: 171-175.

Foreyt, W.J.& D.O. Trainer (1980): Seasonal parasitism

changes in two populations of white-tailed deer in

conservationists. Though the present work was

restricted to chital, cattle may have similar

adverse effects on other wild herbivores in and

around Mudumalai Sanctuary. More detailed

work is required to get a clearer picture of the

effects of cattle on wild herbivores, from a

physiological and pathological perspective,

especially in terms of diseases caused by more

serious pathogens like viruses and bacteria.

Acknowledgements

We thank the Tamil Nadu Forest

Department; Mr. A. Udhayan, 1FS; Dr. V.

Krishnamurihy (retd. Forest Veterinarian);

Mr. A. A. Desai, Dr. N. Baskaran, Mr. S.

Swaminathan (Bombay Natural Flistory Society);

Dr. R. Sukumar, Dr. N.V. Joshi (Centre for

Ecological Sciences); Dr. G. Rajavelu, Dr. S.

Ramesh, Dr. M.G. Jayathangaraj (Madras

Veterinary College) and Mr. Sachin Ranade.

E N C E S

Wisconsin. J. Wild l. Manage. 44: 758-764.

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medicine (Ed.: Fowler, M.E.). 2nd edn. W.B.

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pp. 33-35.

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diseases, with special reference to studies with

nematode parasites of sheep. Aust. Vet. J. 24: 17-

45.

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spatial distributions. Res. Pop. Ecol. 8: 1-7.

(Original not seen)

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determine the population of infective larval

nematodes on pastures. A ust. Vet. J. 17: 181-184.

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A primer on methods and computing. John Wiley

and Sons, Inc., New York. 337 pp.

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intensive management: Population dynamics,

threatened hosts and their parasites. J. Zoo Wild!

Med. 24: 315-326.

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1). APR. 2003

63

EFFECTS OF CATTLE GRAZING AND HABITAT ON HELMINTH LOADS OF CH1TAL

Margolis, L., G.W. Esch. J.C. Holmes, A.M. Kuris & G.A.

Schad (1982): The use of ecological terms in

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Singer, R.S. & W.M. Boyce (1996): Sustainable cattle

production and livestock wildlife interface in

tropical ecosystems. Ciencias Veterinaria Voi.

especial, Pp. 101-104.

Skorping, A., A.F. Reed & A.E. Keymer (1991): Life

history covariation in intestinal nematodes of

mammals. Oikos. 60: 365-672.

(Original not seen)

Soulsby, E.J.L. (1968): Helminths, arthropods & protozoa

of domesticated animals. Williams and Wilkins Co.,

Baltimore. 824 pp.

Stromberg, B.E. (1997): Environmental factors influencing

transmission. Vet. Parasitol. 72: 247-264.

Sukumar, R. (1990): Ecology of the Asian elephant in

southern India II. Feeding habits and crop raiding

patterns. J. Trap. Ecol. 6: 33-53.

Sukumar, R. (1992): Asian Elephant: Ecology and

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Sukumar, R., H.S. Dattaraja, H.S. Suresh, J.

Radhakrishnan. R. Vasudevan, S. Nirmala&N.V.

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in a tropical deciduous forest in Mudumalai,

Southern India. Curr. Sci. 62: 608-616.

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Collier-MacMillan, London. 162 pp.

Watve, M.G. ( 1 992): Ecology of nost-parasite interactions

iri a wild mammalian host community in Mudumalai,

Southern India, Ph.D. Thesis submitted to Indian

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264-273.

64

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 1 00(1), APR. 2003

DEMOGRAPHY OF LIONTAILED MACAQUE (. MACACA SILENUS) IN AN

UNDISTURBED RAINFOREST OF SILENT VALLEY NATIONAL PARK,

KERALA, INDIA1 2 * 4

(With two text-figures )

2 3 2,4

Gigi K. Joseph and K.K. Ramachandran

Key words: Macaca silenus, demography, birth rate, survival rate, growth rate, sex ratio

The demography of the liontailed macaque ( Macaca silenus) was studied in the Silent Valley

National Park and its adjacent areas from 1993 to 1996. Birth rate, survival rate and growth rate

were estimated by methods described by earlier authors (Caughley 1977, NRC 1981). Fourteen

troops with 275 individuals were observed. The adult male:female sex ratio was 1:5.6. A low

birth rate (0.22/adult female/year) and a high survival rate (0.98/individual/year) were the

noteworthy features of the population. Birth rate decreased as the troop size and number of adult

females increased. A marginal increase in growth rate was observed. The study provides estimates

of population parameters of the endangered liontailed macaque in an undisturbed and contiguous

rainforest for the first time.

Introduction

Factors such as habitat fragmentation,

reduced habitat area, isolation of populations

leading to inbreeding depression and vulnerability

to random events make the liontailed macaque a

highly endangered species (Kumar et ai 1995,

Easa et al. 1997). Therefore, demographic studies

of this species deserve utmost importance.

The Silent Valley in Kerala and Ashambu

hills in Tamil Nadu are perhaps the only two viable

habitats left for these macaques (Green and

Minkowski 1977, Ramachandran 1990, Joseph

1998, Joseph and Ramachandran 1998). The

demography of this macaque has been studied in

fragmented forests in Anamalai hills, Tamil Nadu

(Kumar 1987). Recently, Umapathy and Kumar

(2000) reported the occurrence and abundance of

liontailed macaque in 25 rain forest fragments in

the Anamalai hills in relation to several ecological

factors. However, no long-term demographic

studies have been attempted so far in any of the

’Accepted July, 2002

2Division of Wildlife Biology,

Keraia Forest Research Institute,

Peechi 680 653, Thrissur, Kerala, India.

JEmail: [email protected]. in

4Email: [email protected]

large contiguous habitats. The present study was

conducted in order to estimate the demographic

parameters such as troop composition, birth rate,

survival rate and growth rate of this primate in the

undisturbed rainforest ecosystem in Silent Valley,

and to compare them with those obtained from

Anamalai hills (Kumar 1987).

Study Area

The Silent Valley National Park is situated

in Palakkad district, Kerala State (11° 3' to

11° 13' N; 76° 21' to 76° 35' E). It is one of the

core areas of the Nilgiri Biosphere Reserve. The

total area of the Park is 90 sq. km and it is

contiguous with Attappady Reserve Forest in the

east, Mukkurthi National Park in the north,

Nilambur forest division in the west and

Mannarkkad forest division in the south (Fig. 1).

Kunthipuzha, a tributary of Bharathapuzha,

originating from the northeastern hill ranges of

the Park, drains the area. The altitude varies from

658 to 2,383 m and the terrain is quite

undulating. Silent Valley is one of the highest

rainfall areas in the Western Ghats, with an

annual rainfall of about 6,000 mm. The annual

mean temperature is around 20 °C. The major

vegetation is of Tropical Wet Evergreen type.

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

65

DEMOGRAPHY OF LIONTAILED MACAQUE IN SILENT VALLEY NATIONAL PARK

66

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

DEMOGRAPHY OF LIONTAILED MACAQUE IN SILENT VALLEY NATIONAL PARK

Methods

The demography of the liontailed macaque

was studied in Silent Valley National Park and

adjacent areas for three years from 1 993 to 1 996.

The primate population was estimated by total

count and sweep sampling methods (NRC 1981,

Whitesides et al. 1988). Census surveys were

made on foot, radiating from the four wireless

stations (Sairandri, Nilikkal, Puchappara and

Walakkad) situated inside the Park. Frequent

stops were made to get the characteristic contact

call of the liontailed macaque. The intermittent

contact calls are audible up to 100 m. The troops

located were followed until each troop was

reliably counted or till it could not be followed.

Poor visibility due to the closed canopy and

highly undulating terrain, with intermittent

inaccessible areas, were the major limitations in

population count. Moreover, the foraging sub

units or consort pairs were often far away from

the troop. Several times, the survey was

terminated due to the presence of elephant herds,

continuous rain and mist. Complete counts were

easy when the troop passed over a stream, path

or some temporary gaps in the canopy. When

minor differences occurred between successive

counts, the larger number was taken as true.

Animals were classified into four age-sex

classes based on their morphological differences:

adult male, adult female, subadult male and

immature. Adult males were identified by their

stouter body, long canines and large tail tufts.

Adult females were identified by their elongated

nipples and baggy breasts. Less developed

musculature and comparatively shorter canines

indicated subadult males. Other individuals were

classified as immature.

A total of nine troops were identified in

1993, of which seven (Sairandri, Aruvampara,

Punnamala I, Parathod, Puchappara, Chembotty 1

and Nilikkal 1) were inside the National Park

and two in the adjoining Panthenthod beat of

the Attappady Reserve Forest (Panthenthod 1 and

Panthenthod 11). Two more troops (Nilikkal 11

and Chembotty 11) were identified in 1994 and

one (Punnamala 11) in 1995. Fission occurred in

two troops (Aruvampara and Chembotty 1)

during 1995 resulting in four troops

(Aruvampara 1 and Aruvampara 11, Chembotty

IA and Chembotty IB). All the troops were

monitored once a year up to 1996.

Disappearance of an animal between

consecutive censuses was considered as a death.

Recruitment to the troop was carefully tallied,

considering the disappearances. A black eagle

( Ictinaetus malayensis) preyed upon an immature

from the Sairandri troop in 1995. An incident of

poaching by Muduga tribals occurred in one troop

(Panthenthod 11). This troop was excluded from

analysis. The newly formed troops after fission

could be counted only once and were also excluded

from analysis. Thus, data for analysis of population

parameters such as birth rate, survival rate and

growth rate, were taken only from 1 1 troops.

Birth rate is estimated as the proportion of

females giving birth in a year, out of the total

number of adult females under observation

(Caughley 1977, Kumar 1987). Survival rate was

estimated as the proportion of individuals that

survived in the observed year out of the total

number of individuals under observation. Per

capita rate of increase or finite rate of increase X

is a simple measure of population growth rate

and was calculated as:

L = Nt+ 1/N{

where N is the number of individuals in a

population at time t. When X is greater than one,

the population has increased in the period t to

t+1. When X is less than one, the population has

decreased, and when X = 1 , the population size

has remained constant (NRC 1981).

Results

Troop composition

Fourteen distinct troops with a total of 275

individuals were identified from Silent Valley and

JOURNAL BOMBAY NATURAL HISTORY SOCIETY 100(1), APR. 2003

67

Birth rate

DEMOGRAPHY OF LIONTAILED MACAQUE IN SILENT VALLEY NATIONAL PARK

Fig. 2a: Mean birth rate during 1993-96 Fig- 2b: Mean birth rate of adult females in a troop

during 1993-96

adjacent areas, when the field study ended in

1996 (Table 1). The troop size ranged from 9 to

36 individuals, with an average of 19.64

individuals (s.e. = 0.982). All troops together

constituted 8% adult males, 45% adult females,

6% subadult males and 41% immatures. The

adult male:female sex ratio ranged from 1 :3.5 to

1:8.5 with a mean of 1:5.64 (s.e. = 0.282).

Birth rate

A total of 295 adult female-years were

monitored to estimate the birth rate during 1993-

1996. A total of 64 infants were bom in 1 1 troops,

giving a birth rate of 0.22/adult female/year.

Mean birth rate was higher in 1994-95 and 1995-

96 (0.23/adult female/year) than in 1993-94

(0.18/adult female/year). Birth rate decreased

when the troop size increased (Fig. 2a). Birth

rate also showed high negative correlation with

the number of adult females (Fig. 2b), The

Chembotty I troop had the greatest number of

adult females and the lowest birth rate (0. 1 1 /adult

female/year).

Survival rate

Table 2 shows the survival rate of all age/

sex classes together in 1 1 troops giving survival

rate of 0.98/individual/year. Fourteen

disappearances were recorded in the study period

and these included eight adult females, three

immature individuals, one adult male and two

subadult males. The highest survival rate (0.99/

individual/year) was recorded for Nilikkal 1

troop, in which only one disappearance occurred

Table 1 : Status of lion-tailed macaque troops in Silent Valley National Park and adjacent areas

68

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

DEMOGRAPHY OF UONTAILED MACAQUE JN SILENT VALLEY NATIONAL PARK

Table 2: Survival rate of all the age/sex classes for each

troop

during the study period. The survival rate was

lowest (0.93/individual/year) in Chembotty II

troop, in which three deaths or disappearances

were recorded.

Growth rate

The data for the estimation of per capita

growth rate was taken from 1 1 troops. Table 3

shows the rate of growth recorded in different

troops in each year. The highest mean growth

rate (1.12/individual/year) was noticed in 1995

and the least (1 .07/individual/year) in 1996. The

mean growth rate over the study period was 1 .09/

individual/year. There was considerable variation

in the growth rate among different troops. The

highest mean growth rate was estimated in

Panthenthod I troop (1.19/individual/year). Out

of the eight troops monitored in 1994, the highest

mean growth rate was in the Puchappara troop

(1.25/individual/year). Ten troops were moni-

tored in 1995, and the highest increase was

recorded in Panthenthod troop (1,29/individual/

year). Out of the nine troops monitored in 1996,

there was no increase in four troops and the

highest rate of increase was recorded in

Punnamala I troop (1.18/individual/year).

Table 3: Per capita rate of growth in various liontaiied

macaque troops

nd = no data

Discussion

Successive monitoring of demographic

variables provides the best means of assessing

the status of a population and the effectiveness

of management (Kyes et cil. 1 998). In the Western

Ghats, the liontaiied macaque is present in small

populations due to extensive fragmentation of

the rainforest habitat. Such small populations

often undergo random shifts in size due to natural

events or human influence, and can lead even to

local extinction. Out of the total wild population

of nearly 4,000 liontaiied macaques, the Kerala

part of Western Ghats holds more than 50%,

while the rest is shared between the states of

Karnataka and Tamil Nadu (Kumar et al. 1995).

The present study indicates that the Silent Valley

National Park population, with at least 14 troops

and 275 individuals, is one of the most important

populations in its entire range of distribution.

This population is part of a larger population in

the 400 sq. km of rainforest nearby in Attappady,

Silent Valley, New Amarambalam area.

The liontaiied macaque forms relatively

small troops compared to other macaques, most

of which have a mean troop size between 20 and

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

69

DEMOGRAPHY OF LIONTAILED MACAQUE IN SILENT VALLEY NATIONAL PARK

30 individuals (Caldecott 1986). Kumar (1987)

monitored 10 liontailed macaque troops in the

Anamalai hills and estimated a mean troop size

of 19.9 individuals. The present study also

revealed an average troop size of 19.64

individuals, with a range of 9 to 36 individuals.

The adult male:female ratio is consistently

less than 3 females per male in the genus

Macaca, except for the pig-tailed macaque

( Macaca nemestrina ) and liontailed macaque

(Caldecott 1986, Kumar 1987). In M nemestrina ,

the adult sex ratio goes up to 8.0 (Caldecott

1986). Kumar (1987) reported a mean adult sex

ratio of 5.6 for liontailed macaque population in

the wild. The present study closely agrees with

the latter in having the mean adult sex ratio as

5.67 females per male.

The remarkably high age at first birth and

low birth rate as compared to other macaques, is

characteristic of the liontailed macaque. Even

though in most of the macaques the age at first

birth is between 40 to 60 months, the lion-tailed

macaque stands out with 80 months. The birth

rate is also very low (0.28) in Anamalai hills

(Kumar 1987), and 0.23 in this study. The low

birth rate in Silent Valley population may be due

to the presence of many large troops with greater

numbers of females. In large troops with more

adult females, fewer females show sexual

Refer

Caldecott, J .0. ( 1 986): Mating patterns, societies and the

ecogeography of macaques. Anim. Behav. 34: 208-

220.

Caughley, G. (1977): Analysis of Vertebrate Populations.

Wiley, Chichester. Pp. 1-234.

Easa, P.S., S. Asari & S.C. Basha (1997): Status and

distribution of the endangered liontailed macaque

( Macaca silenus ) in southern Western Ghats, India.

Biol. Conserv. 80: 33-37.

Green, S. & K. Minkowski (1977): The liontailed monkey

and its South Indian rainforest habitat. In: Primate

Conservation (Eds.: Prince Rainier III and G. H.

Bourne). Academic Press, New York. Pp. 289-337.

Joseph, G.K. (1998): Ecology of liontailed macaque

(Maccica silenus) in tropical forests of Southern

swelling due to increased competition for food

resources (Kumar 2000). When Kumar (2000)

compared two group size classes of lion-tailed

macaque, more births occurred in the small group

size classes indicating the relation between the

group size and birth rate.

According to Kumar (1987), the high

survival rate is a characteristic feature of the

liontailed macaque. The present study

corroborates his finding, in that the mean

survival rate of different troops in Silent Valley

is as high as 0.98/individual/year. The high rate

of immature survival clearly shows increased

investment, e.g. vigilance of adults over

immatures. Various birds of prey like black eagle

(Ictinaetus malayensis ) and crested serpent eagle

(Spilornis cheela ) are considered the most

important predators of immature liontailed

macaque. Tigers and leopards also occasionally

prey on them. Adults very often sense the

presence of these predators and give alarm calls.

Acknowledgements

We thank Dr. J.K. Sharma, Director KFRI

for encouragement, and the Wildlife Wing of the

Kerala Forest Department for funding the

primate research project in Silent Valley National

Park.

E N C E S

Western Ghats, India. Ph.D. Thesis, FR1 Deemed

University, Dehra Dun. Pp. 1-210.

Joseph, G.K. & K.K. Ramachandran (1998): Recent

population trends and management of liontailed

macaque ( Macaca silenus ) in Silent Valley National

Park, Kerala, India. Indian For. 124: 833-840.

Kumar, A. (1987): The ecology and population dynamics

of the lion-tailed macaque (Macaca silenus ) in

South India. Ph.D. Thesis, University of Cambridge.

Pp. 1-174.

Kumar, A. (2000): Sexual harassment among female

liontailed macaques ( Macaca silenus) in the wild.

J. Bombay nat. Hist. Soc. 97: 42-51.

Kumar, A., S. Molur & S. Walker (1995): Liontailed

macaque ( Macaca silenus) Population and Habitat

70

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

DEMOGRAPHY OF LIONTAILED MACAQUE IN SILENT VALLEY NATIONAL PARK

Viability Analysis Workshop- Report. Zoo Outreach

Organization, Coimbatore.

Kyes, R.C., D. Sajuthi, E. Iskandar, D. Iskandriati,

J. Pamungkar & C.M. Crockett (1998):

Management of a natural habitat-breeding colony of

longtailed macaques. Trop. Biodiversity 5: 127-137.

National Research Council (1981): Techniques for the

Study of Primate Population and Ecology. National

Academy Press, Washington, D.C. Pp. 1-233.

Ramachandran, K.K. (1990): Feeding and ranging

patterns of liontailed macaque in Silent Valley

National Park. //?: Ecological studies and long term

monitoring of biological processes in Silent Valley

National Park. Research Report KFRI Peechi.

Pp. 104-133.

Umapathy, G. & A. Kumar (2000): The occurrence of

arboreal mammals in the rain forest fragments in

the Anamalai Hills, south India. Biol. Corner.

92: 311-319.

Whitesides, GAD., GHEE. Oates, SUM. Green & RAP.

Kluberdang (1988): Estimating primate from

transect in a West African Rain Forest: A

composition of techniques. J. Anim. Ecol. 57:

345-367.

m m m

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

71

NEW DESCRIPTIONS

A NEW HUMAN BLOOD FEEDING BITING MIDGE FROM INDIA,

DIPTERA: CERATOPOGONIDAE: FORCIPOMYIA MANASP

( With three text-figures)

Girish Maheshwari* 2

Key words: F orcipomyia manasi sp. nov., vector, human

Forcipomyici manasi sp. nov. was collected from the body of Mr. H.R. Sou, a research scholar,

while it was feeding on his blood. Host-specilicity of the biting midge was further confirmed by

the Precipitin Method. Mouth parts and feeding behaviour were found to resemble species of

Culicoicles , which feed on human blood.

Introduction

Biting midges are vectors of numerous

viral, protozoan and helminth pathogens. A

number of viruses have been isolated from biting

midges, namely the Simbu, Orbivirus,

Rhabdovirus and Oropouche groups and

specially the Culicoides species. The proven

association of biting midges with transmission

of Oropouche virus in man has now elevated the

medical importance of biting midges.

About a hundred species of Forcipomyici

are reported from the world, of which seventeen

are found in India (Borkent and Wirth 1997).

The host-specificity of F orcipomyia is not very

well studied and only a few species are reported

feeders on frogs and birds. F orcipomyia manasi

is perhaps the first record of Forcipomyian biting

midges feeding on human blood.

Material and Methods

A wild population of F orcipomyia manasi

has been used in the present investigation. Fed

females of the species were collected from the

field, and the Precipitin Method was used to

assess the preliminary host-specificity. The

material was prepared for taxonomic studies by

’Accepted May. 2000

2School of Entomology, St. John’s College,

Agra 282 002, Uttar Pradesh, India.

the following method adopted by the School of

Entomology, St. John’s College, Agra, India.

Adults were preserved in 70% ethanol and in

4% aqueous solution of formaldehyde (preserves

coloration better than alcohol). Before dissection,

the material was cleaned in cold KOH (10%

solution in water) and 2-propanol. Volsella of

male genitalia was removed and mounted

separately under a cover slip in lateral view. The

genitalia were first mounted laterally in Canada

balsam and the shape of the apicolatera! process

was noted. The male specimen was then

reoriented to a dorsoventral position, and females

to ventrolateral position. The terminology of

Boorman (1990) and Wirth and Messersmith

(1971) have been followed.

Taxonomic Description

F orcipomyia manasi sp. nov.

Female imago: Length: 1.44 mm. Wing

length: 0.75 mm, width 0.35 mm.

Antenna: (Fig. la): Scape well-developed,

with 8-10 setae; pedicel rounded, width more

than length, with 6-7 setae. Flagellum with

13 flagellomeres; flagellomeres 1 to 8 beaded,

1st and 8th subequal, 2-7 wider than long, 9-13

elongated, ultimate flagellomere longest with

distinct pointed tip. Length, width of pedicel

(mm) 0.030, 0.050. Length, width of

72

JOURNAL , BOMBAY NATURAL HISTORY SOCIETY 100(1), APR. 2003

0.02 mm

NEW DESCRIPTIONS

0.1 mm

Fig. 1: a. antenna, b. maxilla, c. mandible, d. hypopharynx, e. maxillary palp, f. labrum-epipharynx,

g. labium, h. wing

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

73

0.04 mm

NEW DESCRIPTIONS

flagellomeres (mm) 0.024, 0.024; 0.013, 0.022;

0.015, 0.021; 0.015, 0.021; 0.015, 0.019; 0.015,

0.018; 0.015, 0.018; 0.018, 0.018; 0.044, 0.016;

0.047, 0.015; 0.052, 0.015; 0.055, 0.015; 0.079,

0.014. AR* = 2.08.

Terminal

elongated

fl age Horn ere

*AR (Antennal Ratio ) =

Rest of the

basal flagellomeres

Head: Coronal suture absent, frontal

tubercle present. Temporal setae numerous. Eyes

bare, narrowly separated by small bridge,

ommatidia moderate. Clypeus broader than long,

U-shaped, with 1 0 setae. Length, width of clypeus

(mm) 0.048, 0.079. Maxillary palp (Fig. le) with

five palpomeres, of which third and fourth are

ovoid, the former elongated with a large pit

bearing capitate sensilla. Length, width of

palpomeres (mm) 0.012, 0.009; 0.024, 0.012;

0.036, 0.020; 0.017, 0.0141, and 0.021, 0.012

and setae 2, 2, 5, 3, 5, respectively.

Proboscis with well-developed cibarial

pump and upwardly directed cornua. Mandible

(Fig. lc) strong, serrated with 20-22 small teeth;

maxilla (Fig. lb) scalpel-shaped with 15-17

backwardly directed teeth. Labrum-epipharynx

(Fig. If) sclerotized apically with pointed

hypopharynx (Fig. Id); labium (Fig. Ig) setose

and flappy. Mouth parts adapted for blood-

sucking.

Thorax: Humeral pit and parapsidal suture

absent. Antepronotum with 5 setae. Acrostichals

and dorsocentralis numerous, scattered, not

arranged in rows. Scutellum with two rows of

scutellars, mid scutellum bears 8-9 large setae.

and lateral with 7-8 small setae. Anapleural

suture present; pre-episternals absent. Postnotum

bare.

Wing (Fig. Ih): Light brown; costa large,

ending before 2/3 the wing. Radial sector large,

densely covered with macrotrichia. First radial

cell obliterated, second radial cell compact, R4 5

proximally with light pigmentation, false veins

M, and M3+4 present. Media bifurcates distal to

cross vein r-m. Wing densely covered with fine

macrotrichia; microtrichia present on wing

membrane. Brachiolum with 28-30 sensilla

campaniformia. Radial sector with 3 sensilla

campaniformia, Subcosta with two at wing base.

Alula without macrotrichia; squama with two

elongate setae. CR* = 0.64.

Length of costa

*CR (Costal Ratio) =

Total length of wing

Legs (Fig. 2i-m): Femora and tibia slightly

swollen; fore and hind tibial spurs present,

lengths (mm) 0.048, 0.032 respectively; mid

tibial spur absent; width at the apex of fore, mid

and hind tibia (mm) 0.036, 0.028, 0.028

respectively. Fore tibial comb absent; 5-7

elongated setae present at the apex; hind tibial

apex with two combs; first comb with 8 large

spines; third spine longest; second comb with

13 small spines. Pseudospurs and palisade setae

present on first four tarsomeres; ultimate

tarsomere with a pair of markedly curved claws

and an empodium. Length and proportions of

legs as in Table 1.

Genitalia (Fig. 2n & 3o): Alimentary canal

filled with blood. Spermatheca single, circular;

spermathecal neck absent. Coxasternapodeme

Table 1: Forcipomyia manasi sp. nov., female: lengths (mm) and proportions of legs

Legs Fe Ti Ta1 Ta2 Ia3 Ta4 Ta5 LR

P, 0.265 0.274 0.157 0.068 0.060 0.044 0.044 0.573

P2 0.298 0.338 0.153 0.080 0.060 0.048 0.040 0.452

P3 0.322 0.322 0.189 0.092 0.068 0.052 0.048 0.587

74

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NEW DESCRIPTIONS

0.04 mm

Fig. 2: i. hind leg, j,k,l. fore, mid and hind tibial apex respectively, m. apex of ultimate tarsomere,

n. dorsal view of female genitalia

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

75

0.1 mm

NEW DESCRIPTIONS

Fig. 3: o. ventral view of female genitalia

highly sclerotized. Gonapophysis VIII divided

into dorsal and ventral lobes. Postgen itai plate

triangular with a pair of strong setae. Cerci club-

shaped.

Holotype: 9 on slide, india: Uttar Pradesh.

Agra. St. John’s College Campus, I7.viii.1997,

Coil. H.R. Sou, Det. Maheshwari, G.

Paratype: 9 on slide, india: Rajasthan,

Camel Farm (Bikaner), 15.x. 1997, Coil. H.R.

Sou, Det. Maheshwari, G.

Systematics: F manasi sp. nov. is, perhaps,

the only known species of Forcipornyia which

feeds on human blood. According to taxonomic

characters, it comes closest to F. jhapogi

Maheshwari et al. Since the species is

haeinatophagous, the abdomen of the female is

generally inflated by the accumulation of blood.

F. manasi can be distinguished by the presence

of a single spermatheca, highly sclerotized

coxasternapoderne, large sensory pit on third

maxillary palp and U-shaped clypeus. F. jhapogi

can be separated from manasi by the presence of

spermafhecal neck and antepronotum with eight

setae.

Etymology: Since the species feeds on

human blood, it is named manasi.

Discussion

Only a few species of biting midges are

known intermediate hosts feeding on human

blood. These are Culicoides graham ii,

C. inornatipennis and C. austeni. Forcipornyia

spp. feed on a variety of hosts such as insects,

amphibia, birds and mammals. Those feeding

on invertebrates have lacinia with retrorse teeth

and coarsely toothed mandibles. Forcipornyia

manasi sp. nov. is characterised by the margin

of the apical portion of the labrum having a

continuous row of teeth and the absence of

76

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR 2003

NEW DESCRIPTIONS

sensilla coeloconica on first flagellomere of the

antenna, which indicates its adaptation to feeding

on human beings.

The fusion of the ramus with

coxasternapodeme IX, the fusion of gonocoxite

with tergite IX and absence of gonostyius in the

female genitalia is a clear synapomorphy

compared with other such as F confluens and

F. conigera. Presence of a single spermatheca in

F. manasi and jhapogi is an apomorphic

character, which isolates them from other spp.

of Forcipomyia. F. manasi also resembles

F. barbipesi but by the presence of a spermathecal

diverticulum, an apomorphic character, barbipesi

Boorman, J. ( 1 990): Two new Forcipomyia (Lasiokelea)

sp. (Diptera: Ceratopogonidae) from Oman.

Journal of Oman studies 10: 125-130.

Borkent, A. & W.W. Wirth (1997): World species of biting

midges (Diptera: Ceratopogonidae). Bull. Am. Mus.

can be isolated from manasi. The mass culture

of the species is being established in the

laboratory for further studies on vectorial

capacity.

Acknowledgements

1 thank the Department of Science and

Technology (DST), Govt, of India, New Delhi

for financial support. 1 am also thankful to

Dr. S.S. Shukla, Department of Forensic

Sciences, Institute of Forensic Sciences, Agra for

his valuable suggestion to determine host-

specificity by the Precipitin method.

N c e s

Nat. Hist. 233: 1-257.

Wirth, W.W. & D.H. Messersmith (1971): Studies on the

genus Forcipomyia. 1 . The North American midges

of the subgenus Trickohelea (Diptera:

Ceratopogonidae) Ann. Ent. Soc. Am. 64: 15-26.

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

77

A NEW SPECIES OF EUGENIA L., MYRTACEAE, FROM SEITHUR HILLS,

TAMIL NADU, INDIA1

( With one text-figure)

R. Gopalan and S.R. Srinivasan2

Key words: Eugenia seithurensis sp. nov., southern Western Ghats, Tamil Nadu

A new species of Eugenia , from southern Western Ghats Tamil Nadu, India, is described and

illustrated.

Introduction

Seithur hills in Virudhunagar district

(formerly Kamarajar district), Tamil Nadu in

southern Western Ghats is one of the richest areas

in biodiversity. Botanical explorations in this

hilly terrain were conducted in early 1970s by

one of us (SRS). During these surveys a member

of the Family Myrtaceae was collected in a shola

forest, which was misidentified as Eugenia

discifera Gamble and deposited at MH. When

E. discifera was later collected by one of us (RG)

in the Agasthiyamalai (Pothigai) hills,

Tirunelveli district, it did not match with the

earlier determined E. discifera referred to above.

On critical study of the specimen, it was found

that it was not E. discifera and that it was an

undescribed species, which is described and

illustrated herewith.

Eugenia seithurensis Gopalan &

S.R. Sriniv. sp. nov.

E. calcadensis Bedd. affinis, sed foliis

obovatis, ad apicem abrupte acuminatis (acumine

c. 3 mm longo); floribus axillaribus (solitariis

vel binatis) vel terminaliter umbellatis; pedicellis

4-8 mm longis; stylo glabro; fructibus globosis

differt.

'Accepted June, 2000

2Botanical Survey of India,

Southern Circle, TNAU Campus,

Lawley Road P.O., Coimbatore 641 003,

Tamil Nadu, India.

Allied to E. calcadensis Bedd. but differs

in leaves being obovate, abruptly acuminate

(acumen c. 3 mm long) at apex; flowers axillary

(solitary or in pairs) or in terminal umbels;

pedicels 4-8 mm long, style glabrous; fruits

globose.

Tree, up to 10 m tall; branches and

branchlets terete, glabrous, ienticellate. Leaves

opposite, simple; petioles 3-7 mm long, glabrous,

rounded beneath, canaliculate above; lamina

obovate, 3. 2-5. 4 x 2. 9-3. 5 cm, coriaceous,

glabrous, punctate on both surfaces, attenuate at

base, entire and recurved along margins, abruptly

acuminate (acumen c. 3 mm long) at apex.

Flowers axillary (solitary or in pairs) or in

terminal 4-8-flowered umbels or cymes, bisexual,

actinomorphic; pedicels stout, 4-8 mm long,

hirsute; bract 1, linear-lanceolate or lanceolate-

elliptic, c. 5 mm long, hirsute; bracteoles 2,

linear-lanceolate, c. 3 mm long, hirsute. Calyx

tube campanulate, 2-3 mm long, appressedly

brown silky hairy; lobes 4 (2 + 2), subequal; outer

sepals smaller, broadly ovate, 2. 5-3. 5 x 3-4 mm,

thick, sparsely hairy, ciliate along margins,

concave; the inner orbicular to suborbicular, 3-5

x 4-5 mm, punctate, sparsely hairy, broad at base,

ciliate along margins, truncate or rounded at

apex. Petals 4, obovate or elliptic-oblong, c. 1 1

x 7 mm, thick in middle, membranous along

periphery, punctate, ciliate along margins above

1/3, obtuse at apex. Stamens many; filaments

4-10 mm long, glabrous; anthers globose,

2-loculed, basifixed. Ovary inferior, obconic,

78

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

NEW DESCRIPTIONS

Fig. 1: Eugenia seithurensis sp. nov., A-B. Twigs with terminal inflorescence & axillary flowers,

C. Bracteole, D. Outer sepal, E. Inner sepal, F. Petal, G. Pistil, H. Fruit

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

79

NEW DESCRIPTIONS

appressedly hairy, 2-3 (-4) mm long; style

5-7 mm long, thick, glabrous; stigma simple.

Disc hairy. Fruits globose, c. 2 cm, crowned with

persistent calyx lobes; seeds globose, c. 1 .4 cm.

Holotype (S.R. Srinivasan 63532, CAL),

isotypes (S.R. Srinivasan 63532, MH — acc. no.

120279 - 120283) and paratype (S.R. Srinivasan

65986, MH — acc. no. 120285 - 120292) were

collected in between Deviar Estate and forest

border of Kerala, Seithur Hills, Virudhunagar

district, Tamil Nadu, at about 1,350 m above msl

on June 12, 1979.

Rather rare in sholas, only a few trees were seen.

FI. & Fr.: April-October.

Etymology: This species is named after the

type locality.

Acknowledgements

We thank Dr. R Daniel, Deputy Director,

BSI, Coimbatore for encouragement and

facilities, Dr. V.J. Nair, Scientist Emeritus, for

the Latin diagnosis, Shri. N.C. Rathakrishnan,

former Scientist, BSI, Coimbatore, for

confirming the novelty and Dr. A.N. Henry,

Scientist Emeritus, for suggestionss.

80

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

SONERILA L ONGIPETIOLA TA MAN1CKAM ETAL., A NEW SPECIES

OF MELASTOMACEAE FROM TAMIL NADU, INDIA'

( With one text-figure )

M.M. Josephine, V.S. Manickam, C. Murugan, V. Sundaresan and G.J. Jothi* 2

Key words: Sonerila longipetiolata sp. nov., Kanyakumari district, Tamil Nadu

Sonerila longipetiolata Manickam et al ., a new species is described below with

illustrations.

During recent floristic studies conducted

along the southern Western Ghats, we collected

specimens of an interesting species of Sonerila

Roxb. On critical study, it was found to be a new

species. Hence, it is described and illustrated

here.

Sonerila longipetiolata sp. nov.

(Fig. 1)

Sonerila longipetiolata sp. nov. est affinis

ad S. travancorica Bedd. sed est dissimilis folia

base inaequali, petiolata longa, corolla

emarginata, indumenta molle et floris numerosis.

Typus: india: Tamil Nadu, Kanyakumari

district, Muthukuzhivayal path c. 1,300 m,

22.xii.1999, Coll. V. Sundaresan and M.M.

Josephine 20104 (Holo. XCH, Xavier’s College,

Palayamkottai).

Perennial herb, c. 30 cm high. Stem

3-5 mm in diam; adpressedly villous, branched,

base terete, apex two channelled; internodes

2.5-11.0 cm long. Leaves opposite, unequal;

petiole 1. 0-7.0 cm long, villous; blade ovate-

elliptic, 1. 0-7.0 x 1. 5-5.0 cm, coriaceous, softly

adpressedly villous, base unequal, slightly

cordate, margins sub entire, ciliate, apex acute,

veins 2-3 pairs, pinkish beneath. Inflorescence

scorpioid cyme, terminal. Peduncle 3. 0-5.0 cm

long, puberulous. Pedicel 1.0-1. 5 cm long,

'Accepted June, 2000

2Centre for Biodiversity and Biotechnology,

Research Department of Botany,

St. Xavier’s College, Palayamkottai 627 002,

Tamil Nadu, India.

puberulous. Flowers 10-15 per peduncle, mauve-

deep pink. Calyx 0. 7-1.0 cm long, tube

infundibuliform, teeth 3, short, glandular hairy.

Petals 3, free, 1.0-1. 2 x 0.7-0.75 cm, glabrous

within, glandular hairy without mid nerve, apex

emarginate, obovate-elliptic, 8-9 nerves,

prominent. Stamens 3, equal, c. 1.0 cm long,

free; anther c. 0.7 mm long, yellow, oblong,

divaricated, much attenuated upwards; filament

c. 6.2 mm long. Ovary inferior, 1.0 x 0.33 cm,

glandular hairy; ovules numerous. Style simple,

filiform, 1.0-1. 2 cm long; stigma small,

capitellate. Capsule trigonous, 0.75-1.0 x 0.31-

0.33 cm, six ribbed, glandular hairy. Seeds

numerous, c. 1.0 mm long, raphe excurrent on

the side near the top.

FI. and Fr.: December-January.

Habitat: Understorey of evergreen forest,

semi-shaded, associated with Bambusa

arundinacea Willd.

Table 1 : Comparison between

S. travancorica and S. longipetiolata

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

81

lulu g z

NEW DESCRIPTIONS

Fig. 1 : Sonerila longipetiolata sp. nov., A. Habit, B. Flower, C & D. Petal (Dorsal & Ventral view),

E. Pistil with Calyx, F. & G. Stamens, H. ovary (l.s.), 1. Ovary (c.s.), J. Seed, K. Stern (c.s.)

82

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

NEW DESCRIPTIONS

Sonerila longipetiolata is allied to

S. tra\>ancorica but differs from it as shown in

Table 1.

Status: As the species is restricted to only

one locality, it must be categorized as rare; since

the number of individuals is small, vulnerable

is probably more correct.

Etymology: The species is named after its

characteristic long petiole.

Acknowledgements

We thank Dr. R. Gopalan, Botanical Survey

of India (S. Circle), Coimbatore for his critical

comments on the taxon. Our gratitude to Chief

Wildlife Warden, Field Director, KMTR for

permission to carry out the survey. We also thank

the University Grants Commission (UGC) for

financial assistance.

References

Gamble, J.S. (1957): Flora of The Presidency of Madras. Vol 1. Adlard & Son Limited, London Pp. 352.

Clarke, C.B. (1879): Melastomaceae. In: The Flora of British India (Ed.: Hooker, J.D.). Vol 2. L. Reeve & Co., London.

Pp 534.

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

83

A NEW SPECIES OF SPIDER OF THE GENUS TIBELLUS SIMON

(ARANEAE: THOMISIDAE) FROM JHENIDAH, BANGLADESH1

( With six text-figures)

V. Biswas2 and D. Raychaudhuri3

Key words: New species, spider, Tibellus, Araneae, Thomisidae, Bangladesh

A new species of spider genus Tibellus Simon, T. shikerpurensis is described and illustrated from

Jhenidah, Bangladesh.

Introduction

Spiders of the genus Tibellus Simon

(Family: Thomisidae) are poorly known in

Bangladesh although a few records, on different

species, were found (Chowdhury and Nagari

1981, Biswas et. al. 1993, Okuma et. al. 1993,

Begum and Biswas 1997). Several species on the

other hand, of the genus are known from

countries like India (Tikader 1980, Tikader and

Biswas 1981) and Pakistan (Dyal 1935).

The genus Tibellus Simon is being reported

for the first time from Bangladesh and the same

is being described as T. shikerpurensis sp. nov.

The types are at present in the collection of the

Department of Zoology, Government P.C.

College, Bagerhat, Bangladesh.

Materials and Methods

Collection and preservation of the spider

specimens were made following Kaston (1972)

and Tikader (1987). The materials were studied

with a Stereozoom Binocular Microscope, model

Zeiss, SV8. All the measurements are taken with

an eyepiece.

The species was identified by following

Tikader (1980, 1987) and was confirmed by the

Zoological Survey of India, Kolkata.

'Accepted October, 2000

2 Department of Zoology, Government P.C. College,

Bagerhat 9301, Bangladesh.

'Entomology Laboratory, Department of Zoology.

University of Calcutta, 35, Ballygunge Circular Road,

Kolkata 700 019, West Bengal, India.

Tibellus shikerpurensis sp. nov.

(Figs 1-6)

General: 9, Brownish-green; cephalo-

thorax brownish-green; legs greenish; abdomen

greenish-white.

Measurements (in mm): Total length

6.40; carapace 2.20 long, 2.00 wide; abdomen

4.20 long, 3.10 wide.

Cephalothorax: Broad, slightly longer

than wide, wider near base, clothed with fine

hairs and pubescence (Fig. 1); cephalic region

raised and produced anteriorly, with straight

anterior margin. Eyes in 2 rows, both the rows

strongly procurved, ocular area wider than long;

postero-laterals distally placed and larger than

others; 2 longitudinal brownish bands running

between the base of posterior eyes and posterior

margin clothed with spines and hairs. Chelicerae

strong and stout, clothed with sharp spines, each

of inner and outer margins with 2 teeth (Fig. 2).

Palps long, filiform, 1.2 mm in length (Table 1)

each covered with sharp spines and setae.

Maxillae longer than wide, medially wide and

anteriorly scopulate (Fig. 3). Labium wider

medially, anteriorly narrowing and scopulate

(Fig. 3). Sternum heart-shaped, pointed,

posteriorly clothed with spines (Fig. 4). Legs long

and slender, with spines and hairs; leg formula

1243 and the measurements (in mm) as in

Table 1.

Abdomen: Longer than wide, posteriorly

narrowing, blunt, clothed with hairs and

pubescence; dorsum decorated with brownish

84

JOURNAL , BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

NEW DESCRIPTIONS

1 mm

0.5 mm

i 1

Figs 1-6: Tibellus shikerpurensis sp. nov.. Female Holotype, 1. Female dorsal view (legs omitted),

2. Chelicera, 3. Maxillae and Labium, 4. Sternum, 5. Epigynum, 6. Internal genitalia

JOURNAL BOMBAY NATURAL HISTORY SOCIETY 100(1), APR. 2003

85

NEW DESCRIPTIONS

Table 1 : Measurements of legs and palps for ( ? ) Tibellus shikerpurensis sp. nov.

markings and patches; epigyne and internal

genitalia as in Figs 5 and 6.

Materia! examined: Type-Data: Holotype:

9 in spirit will be deposited in the Department

of Zoology, University of Dhaka, Bangladesh.

Type locality: Shikerpur, Jhenidah,

18.vii.1993, Coll. V. Biswas.

Paratype: 1 9, same as for the holotype.

Distribution: Bangladesh: District Jhenidah.

Etymology: The species has been named

after the type locality.

Remarks: The species T. shikerpurensis

sp. nov. resembles T. chaturshingi Tikader (Tikader

1980) but stands distinct because of the following

1. Cephalic region raised and produced.

2. Anterior row of eyes and posteromedians

forming a hexagon.

Refer

Begum, A & V. Biswas (1997): A list of the spider fauna

of Barisal division, Bangladesh (Araneae: Arachnida).

Bangladesh J. Zool. 25(2): 207-210.

Biswas, V., H.R. Khan, N.Q. Kamal & A. Begum (1993):

A preliminary study of the rice-field spiders in

Jhenidah, Bangladesh. Bangladesh J. Zool. 21(1):

85-92.

Chowdhury, S.H. & S. Nagari (1981): Rice-field spiders

from Chittagong. Proc. Zool. Soc. Bangladesh , pp.

53-72.

Dyal, S. (1935): Fauna of Lahore 4: Spiders of Lahore.

Bull. Dept, of Zool., Panjab Univ. 1: 1-252.

Kaston, B.J. (1972): Howto know the spiders. 2nd edn..

3. Cephalothoracic bands nearly straight.

4. Abdomen not overhanging cephalothorax

with several black markings.

5. Much different epigynum.

These differences justify the recognition of

the species as new to science.

Acknowledgements

The authors are grateful to Dr. S.C.

Majumder, Scientist-SD, Sunderban Field

Research Station, Zoological Survey of India,

Canning, West Bengal, for confirming the

identity of the species and the Head, Department

of Zoology, University of Calcutta, for providing

laboratory facilities.

N C E S

Wm. C. Brown Co. Pub., Dubuque, Iowa, 272 pp.

Okuma, C., N.Q. Kamal, Y. Hirashima, Z. Alam & T.

Ogata ( 1 993): Illustrated Monograph on the rice-field

spiders of Bangladesh. IPSA-JAICA, Salna, Gazipur,

pp. 1-93.

Tikader, B.K. (1980): The Fauna of India, Spiders:

Araneae, Vol. I, Zoological Survey of India, Calcutta,

247 pp.

Tikader, B.K. (1987): Handbook of Indian Spiders. Director,

Zoological Survey of India, Calcutta. Pp. 25 1 .

Tikader. B.K. & B. Biswas (1981): Spider fauna of Calcutta

and Vicinity. Rec. zool. Surv. India, Occ. Pap. No.

30: 1-149.

86

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

FURTHER CONTRIBUTION TO BIOSYSTEMATICS OF CHEN OP ODIUM,

REPORTING THREE NEW SPECIES FROM NORTH INDIAN PLAINS1

( With three text-figures )

S.C. Pandeya2’3 and Amita Pandeya2

Key words: Chenopodium adpressifolium sp. nov., C. sagittatum sp. nov.,

C. hastatifolium sp. nov., photonastic movements

Three new species of Chenopodium aggregate occurring in the north Indian plains have been

distinguished and described as Chenopodium ci>

C. hastcitifolium sp. nov.

Introduction

Three new species of Chenopodium

aggregate occurring in the north Indian plains

have been distinguished and described as

Chenopodium adpressifolium sp. nov.,

C. sagittatum sp. nov. and C. hastatifolium

sp. nov. Earlier Pandeya et al. (1998) have

communicated two new species of the genus from

the north Indian plains.

Further, in both extensive and intensive

surveys over the last several years, three new

species of the genus Chenopodium have been

distinguished from the north Indian Plains (alt.

100-250 m). All the three species are edible and

occur naturally as weed in winter crop fields, in

gardens and other moist places. They start

growing in November and flower from January

to April. Ecoclimate of the region is Tropical

semi-arid and soils are Pleistocene with fresh

alluvium of great depths. The specimens are

deposited in Raja Balwant Singh College, Agra,

India.

The three suspected species were put

through a Provenance Trial (neutral garden

experiment) at Agra for three consecutive years

'Accepted November, 2001

2Botany Department,

Raja Balwant Singh College,

Agra 282 002, Uttar Pradesh, India.

’Present Address: E-104 Murdhanya Apartment,

Opp. Super Society, Nr. ISRO Colony, Ramdevnagar,

Ahmedabad 380 015, Gujarat, India.

ressifolium sp. nov., C. sagittatum sp. nov. and

for observing any plasticity therein. In neutral

garden experiments, the plants collected from

various habitats are grown together side by side

under similar soil and climatic conditions in

order to eliminate features developed due to

differences of habitat. The quantified description

of the three new species pertains to the neutral

garden experiments.

Clt enopo diuni adpressifolium

Pandeya & Pandeya sp. nov.

Affinis C. album Linn. Sp. PI. 219, 1753.

Differt a C. album Linn, f habitus, positus

foliis, folia crasso, marginemque laminae, folia

photonasticus, nervo rosea.

Herbae annuae, erectae. Herba 1 m alta.

Caule fibro, erecto, angusto costato, ramoso.

Ramosae longior ad 10-25 cm, longis basales

rosettus. Folia ovatus, dentatus, marginemque,

laminae 3-5 cm to 1. 5-3.0 cm, pubescentia in

primordiis, nervo rosea. Petiola 2. 5-3. 5 cm,

rosea. Inflorescentia spikus, pedunculata 18 cm

longum, brevis spikus 1-2 m ad folia axillaris.

Flores perianthus 5, rosea. Stamina 5, longior

quam perianth. Stigma pilosum, bifidis, brevis-

brunnea. Semina brunnea 1.26-1.30 x 0.61-

0.67 mm.

Holotypus 910 et positus Raja Balwant

Singh College Agra, India, lectus Agra ad

February 26, 1999. Isotypus Ibid. S.C. Pandeya

& A. Pandeya. Fig. 1 .

JOURNAL BOMBAY NATURAL HISTORY SOCIETY 100(1), APR. 2003

87

NEW DESCRIPTIONS

5.0

Fig. 1: Chenopodium adpressifolium, a. Habit and maximum size of leaves, leaves turn crimson

during senescence, b. Filaments longer than perianth,

c. Purple perianth upon maturity, d. seeds with pericarp, e. seeds without pericarp

88

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY 100(1), APR 2003

NEW DESCRIPTIONS

An erect annual herbaceous plant. Starts

growing in January, flowers in February /March

and senesces by April/May; Height up to 1 m;

Stem fibrous and hard, somewhat ridged,

greenish-red to purple striped, turning

homogeneously crimson upon maturity;

Branches limited up to 10 cm from the base,

decumbent, longest branch 20-25 cm; giving

mature plant a rosette at the base with intemodes

2-3 cm; Leaves ovate, petiole and lamina in one

line, erect, attached to the node at an angle of

70-75° during day and completely adpressed at

night (photonastic movement), petiole 2.5-

3.5 cm, reddish-green; lamina olive green, veins

reddish-green, dentate margin purple in mature

leaves, dentations sharp and pointing upwards,

lamina 3. 0-5.0 x 1. 5-3.0 cm, 0.56-0.8 mm thick

and brittle, heavy mealiness on primordia, young

stem leaves and perianth, lamina and petiole turn

crimson upon maturity, terminal spike up to

18 cm long, few small spikes (1-2 cm) in axil of

upper leaves; Flowers- perianth 5, central vein

prominent; Stamens 5, longer than perianth,

anthers coming out of flowers, feathery stigma

bifid, small, purple, seeds covered with pericarp

with a circular opening on the top, biconvex, disc-

shaped, dark brown, 1.26-1.3 x 0.61-0.67 mm.

Holotype deposited at Raja Balwant Singh

College, Agra, India. No. 910. Fig. 1.

Etymology: The species has been named

so as its leaves get completely adpressed to the

stem during night owing to photonastic

movement.

The specimen was sent to Prof. Pertii Uotila

of Finland. He (1997, pers. comm.) opined that

C. adpressifolium belongs to the group of

C. album resembling in seed shape, size, and

surface structure, as well as in the general shape

of the leaves. The two species under discussion

differ largely in their habit, position of the leaves,

leaf size, leaf thickness and photonastic

movement of leaves in the former species. He

was of the view that the taxonomy of C. album

in India needs revision.

Chen op odium sagittatum

Pandeya & Pandeya sp. nov.

Affinis C. moquianum Aellen.

C. moquianum Aellen affinis, ab ea differt:

habitus, ramosa longior, folia gigantea, supra,

lobi, lamina sagittus.

Herbae, annuae, erectae, 3.65 m alta. Caule

erecto, roseo, basales circumference c. 12 cm.

Ramosa longior ad 65 cm. Folia petiolata c. 10-

12.5 cm longa, diam 21.5 cm, lamina sagittus

17-20 cm x 10-12 cm, lobus 2-2.5 cm.

Inflorescentia terminalis, spikus, bracts

lanceolatus, rosea, pubescentibus. Stamina 5,

equilongus perianthus. Stigma bifidus, longa

c. 0.64 mm, erectae, ovary diam 0.32 mm.

Semina nigra brunnea, diam 0.91- 0.96 mm.

Holotypus 911 et positus Raja Balwant

Singh College, Agra, India, lectus Agra ad March

12, 1999. Isotypus Ibid. S.C. Pandeya & Amita

Pandeya. Fig. 2.

An erect annual herbaceous plant growing

to 3.65 m height, starts growing in January-

February, flowers from mid-April and spikes fully

mature by May end; Stem ridged, red-striped,

turning completely red up to apex upon maturity,

circumference at the base up to 12 cm branching

throughout, branches straight, stiff and at an

angle of 30-40° from the node, branches parallel

to each other in acropetal order, number of

branches on mature plants 40-50. Several

branchlets on each branch, longest branch up to

65 cm, when young red blotch on nodes; Leaves-

petiole at an angle of 35-40° to stem and lamina

30° to petiole, at night both petiole and lamina

become straight adpressed to the stem at an angle

of 80-85°, lamina boat-shaped at night clasping

the stem, exhibiting the photonastic movement.

Petiole and lamina venation light purple, petiole

10-12.5 cm long, diameter 2.15 cm lamina arrow

shaped (sagitate) with small side lobes pointing

upwards, lamina 17-20 cm long and 10-12 cm

at the broadest part, side lobes 2-2.5 cm,

sometimes bifid, young lamina greyish-green.

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

89

NEW DESCRIPTIONS

Fig. 2: Chenopodium sagittatum , a. Habit with acropetal branching, b. Maximum size of leaves, old leaves

turn crimson, c. Flowering branch, anthers coming out of the flowers, d. Crimson perianth upon maturity,

e. Seeds with pericarp, f. Seeds without pericarp

90

JOURNAL BOMBAY NATURAL HISTORY SOCIETY 100(1), APR. 2003

NEW DESCRIPTIONS

heavy mealiness. Both petiole and lamina turn

crimson during senescence. Lamina 0.34 mm

thick in vertical section, stele with both upper

and lower epidermis red in colour. Inflorescence

terminal along with branchlets up to 15 cm long,

branches and branchlets bear up to 12 cm long

spikes, spikes and lanceolate bracts all turn

crimson upon maturity of seeds. Flowers 1 . 1 mm

across, perianth 5, connate at case, greyish-green

when young, crimson upon maturity, heavy

mealiness. Stamens 5, anthers dehisce soon after

opening of flowers, filaments equal to perianth.

Stigma bifid, purple and erect, 0.64 mm long

above ovary, diameter 0.32 mm. Seeds disc-

shaped, notched, blackish-brown, diameter 0.91-

0.96 mm, width 0.6-0.61 mm. Holotype

deposited at Raja Balwant Singh College, Agra,

India. No. 911. Fig. 2.

Etymology: The species has been named so

as it has a characteristic sagittate lamina margin.

For this species Uotila ( 1 997, pers. comm.)

opined that it might be close to a taxon called

C. moquianum Allen. C. sagitattum can easily

be distinguished from the said group on account

of its very large leaves. The leaves in C.

moquianum are small and resemble Chenopod.

Chenopoilium hastatifolium

Pandeya & Pandeya sp. nov.

Affinis C. ficifolium Sm. FI. Brit. 1 : 276

(1800).

C. ficifolium Sm. Affinis ab ea differt folia

tribus lobis obscuris, oblongo; foliis minuta,

inferio.

Herbae annuae, erectae c. 1 m alta,

rarnosae 20-25, longior rarnosae 60-70 cm.

Caulis rosea green, stripus basales circumference

c. 2.7 cm, folia 3-lobus, lobus oblongo, dentibus

- 2, lamina pubescens, petiolata 3-4.5 cm longa.

Inflorescentia terminalis, spikes 5-10 cm iongis,

Folia basales spikes parvi. Flores parvi,

perianthus 5 basales connatis. Stamina 5

acquilongis perianthus, purplish. Carpella-

stigma longo, bifidus. Semina biconvex, diameter

0.96-1.02 x 0.56-0.61 mm, brunnei.

Hoiotypus 912 et positus Raja Balwant

Singh College, Agra, India, lectus Agra ad

March 15, 1999. Isotypus Ibid. S.C. Pandeya &

Amita Pandeya. Fig. 3.

An annual erect herb, up to 1 m, profusely

branched (20-25 main branches), longest middle

branches 60-70 cm, starts growing in November-

December, flowers from February-March, fruits

in March and senesces by April end. Stem

greenish-red striped, ridged, circumference at

base up to 2.7 cm. Leaves green, completely

crimson upon senescence, 3- lobed, middle lobe

oblong 3.5 x 0.6-1. 2 cm, with two dentations,

basal two lobes pointing upwards, 0.6- 1.8 cm

long, sometimes with 1-2 dentations, lamina 0.2-

0.26 mm thick with heavy mealiness when

young. Petiole 3-4.5 cm long. Inflorescence

greyish-green, turning crimson upon maturity of

seeds due to perianth changing to crimson,

terminal spike 5-10 cm, several small spikes in

axil of leaves; Flowers-perianth 5, connate at

base, central vein prominent. Stamens 5, equal

to perianth. Stigma purple bifid, long and

moustache-like, each arm 0.3-0. 4 mm long.

Seeds biconvex covered with pericarp with a

circular opening on the top. Seeds without

pericarp 0.96-1.02 x 0.56-0.61 mm, dark brown

with a small notch. Holotype deposited at Raja

Balwant Singh College, Agra, India. No. 912.

Fig. 3.

Etymology: The species has been named

on the basis of its hastate leaf margin.

For this specimen Uotila (1997, pers.

comm.) is of the view that it is close to C.

ficifolium although the leaf shape is not very

typical of that taxon. However, the interspecific

taxonomy of the species is not completely known.

Acknowledgements

We thank Prof. Perti Uotila, Finish

Museum of National History, University of

JOURNAL. BOMBAY NATURAL HISTORY SOCIETY. 100(1). APR. 2003

91

NEW DESCRIPTIONS

Fig. 3: Chenopodium hastatifolium, a. Habit with maximum size of leaves, crimson old leaves,

b. Heavy mealiness on flowers, c. Bifid stigma with long arms, d. Perianth turns reddish upon maturity

Helsinki, Finland for critically examining the

three specimens and giving his expert opinion.

We thank Prof. A.B. Bhatt of HNB Garhwal

University, Srinagar (UA) for critically going

through the final manuscript, Dr. V.P. Bhatnagar,

Head and Dr. Anil K. Bhatnagar both of Botany

Department, Dayalbagh Educational Institute,

Agra for their cooperation and facilities,

Dr. R.K.S. Rathore, Head Department of Botany,

RBS College Agra for discussions and Dr. R.S.

Parekh for permission to work on his farm at

Poiya Ghat, Agra.

Reference

Pandeya. S.C. Geeta Singhal & Anil K. Bhatnagar (1998): Biosystematic study of two new species of Chenopodium

from north Indian Plains. J. Bombay nat. Hist. Soc. 95(3): 477-487.

92

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY 100(1), APR. 2003

Professor M.S. Mani

(1908-2003)

OBITUARY

PROF. M.S. MANI

The year was 1 968, as a young postgraduate

student of Zoology at the Department of Zoology,

St. John’s College, Agra, I had the opportunity to

meet Prof. Mani when he used to take classes while

serving as Emeritus Professor. Thus, I had the golden

opportunity to hear his lectures while I was in the

final year M. Sc. class. I still remember the scholarly

presentation and disposition of Prof. Mani.

Professor Mani was an affectionate and highly

respected man, who led a very disciplined life. He

worked hard, and it is not surprising that he wrote

over 35 books and more than 300 papers. Even at

the age of 94, he went to Presidency College and

worked. Personally, he was always extremely kind

towards me. In 1979, when I asked him for some

Chalcididae specimens for my studies on loan, he

sent me several boxes of specimens with an

affectionate letter saying that I need not return them.

He gladly wrote a foreword for my book on Parasitic

Hymenoptera in 2001. I consider myself fortunate

to have been associated with such a great man.

Prof. Mahadeva Subramania was born on

March 2, 1 908 at Thanjavur (Tanjore), Tamil Nadu.

After schooling at K.S. High School, Thanjavur, he

joined the Govt. College, Coimbatore for his

Intermediate in 1928. He later joined the Madras

Medical College, but had to leave due to financial

constraints. In 1933, he proceeded to Calcutta

(Kolkata) in search of a suitable job and later joined

as a tutor-cum-demonstrator in Physics on a part-

time basis at Bangabasi College, Sealdah. At the

same time, he worked at the Indian Museum of the

Zoological Survey of India, Calcutta as an honorary

research student. A few years later, he got a small

job at the ZSI, started working on plant galls and

gall insects, and soon became interested in

Chalcidoidea. In 1 937, he obtained an M.A. degree

from the University of Madras, on the basis of his

research papers in Entomology. He was the only

candidate to be so honoured. In 1937, M.S. Mani

joined the Imperial Agricultural Research Institute

(now the IARI, New Delhi) as a Research Assistant

under Dr. H.S. Pruthi. In 1940, he published his

work "Biological Notes on the Chalcidoidea’ with

Dr. Pruthi. In 1944, Dr. Mani resigned his post at

IARI because of some differences between him and

Dr. Pruthi. For some time, he had no job, but worked

as a German language Translator for the Indian

Army at New Delhi (Dr. Mani knew several

European and Indian languages). In 1945, Dr. Mani

joined St. John’s College, Agra as a lecturer, and

there he prospered. He obtained his D.Sc. degree

from the Agra University for his work on galls and

gall insects of India in 1947. In 1950, he established

the famous School of Entomology at St. John’s

College, Agra, where he initiated research on various

aspects of entomology. It was here that he initiated

pioneering research on High Altitude Entomology

and led several entomological expeditions to the

Himalaya during 1953-56. These records were

published in the JBNHS. Prof. Mani joined the

Zoological Survey of India as Deputy Director in

1956 on an invitation from the Central Ministry,

New Delhi and later served the Institution as

Officiating Director. In 1968, he returned to the

School of Entomology, Agra, as Emeritus Professor

and continued there until 1984, when he went to

Chennai. He worked at the Zoological Survey of

India, Chennai Regional Station as a Principal

Investigator of a DST Project and later shifted to

the Botany Department, Presidency College,

Chennai. Since 1990, he was working there as

Emeritus Professor.

Prof. Mani led an Indian delegation of

Zoologists to the USSR in 1963. He represented

India in the Man and Biosphere (MAB) Committee

on Alpine and Arctic Ecology in Lillehammer,

Norway in 1972. He was a Visiting Professor of

Entomology at Tribhuvan University, Kathmandu,

Nepal in 1 975. Gordon Edwards of Colorado named

him the Dean of High Altitude Entomology in 1971.

He was awarded the Scientist of the Year award by

Presidency College in 2000. Recently, the Ministry

of Environment and Forests, Govt, of India awarded

him the E.K. Janakiammal award for taxonomy,

2001, which carried a cash award of Rs. 1 Lakh.

With the demise of Prof. M.S. Mani, not

only India but the world has lost a renowned

scientist.

■ T.C. NARENDRAN

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, / 00(1), APR. 2003

95

REVIEWS

1 . BEAUTIFUL ORCHIDS OF NEPAL by Keshab R. Rajbhandari and Sushila Bhattarai.

Published by authors. 2001. Pp. i-viii + 1-220 (22 x 14 cm). Price Rs. 1,200/-.

This book covers 101 orchid species

belonging to 43 genera out of the 363 species

belonging to 97 genera found in Nepal. Orchids

attract professionals like botanists,

horticulturists, florists as well as amateur plant

lovers due to their showy flowers, fascinating

structures and variety of colours. This handbook

is useful for identifying some of the common

showy orchids found in Nepal, and perhaps

intended for the lay reader who generally does

not want to read scientific terminology to identify

plants. The book is mainly based on good and

well-identified photographs, some of which are

taken from potted cultivated plants. The book

has limited value for amateurs, as it does not

cover all the orchids from Nepal and it does not

2. A BIBLIOGRAPHY OF THE PLANT

Keshab R. Rajbhandari. Published by The

Museum, University of Tokyo, Hongo 7-3-1

1-160 (25.5 x 18 cm). Price not mentioned,

This is a supplement to the bibliographic

book published in 1994 on the plant science of

Nepal. It gives the titles published during the

last 6 years.

The first 78 pages of the supplement

contain alphabetical entries arranged in the

following order: author names followed by titles

and publication data. The remaining 82 pages

contain the subject index and index to scientific

give identification keys, although the short

descriptions given are sufficient to recognize the

species.

It has references to earlier works on the

subject, index to scientific names and glossary

of botanical terms used in descriptions.

The book is printed on fine art paper, but

this has resulted in unjustified wastage of paper

space after each description. With careful layout,

at least one-third of the printed pages could have

been saved. These days, printing of coloured

plates has become quite a costly affair, however,

the price at Rs. 1,200/- is a little steep for this

book.

■ M.R. ALMEIDA

SCIENCE OF NEPAL (Supplement 1) by

Society of Himalayan Botany [University

, Tokyo 113-0033, Japan], 2001 . Pp. i-xiii +

names of plants (with cross references to authors

in the alphabetical bibliography).

As stated by Samar Bahadur Malla in his

foreword, documentation is a vital pillar in

scientific research and technological

development, and I am sure this Supplement will

be useful in the study of plant wealth of Nepal.

■ M.R. ALMEIDA

3. AROMATIC AND MEDICINAL PLANTS - YIELDING ESSENTIAL OIL FOR

PHARMACEUTICAL, PERFUMERY, COSMETIC INDUSTRIES AND TRADE by

M.P. Shiva, Alok Lehri and Alka Shiva. Published by International Book Distributors,

9/3, Rajpur Road, Dehra Dun, Uttaranchal, 2002. Pp. i-viii + 1-341 (24 x 18 cm), with

12 line-drawing plates and 20 colour photographs. Price Rs. 1,950.00 or $ 65.00.

The book mainly deals with aromatic

plants, so its title “Aromatic and Medicinal

Plants” is somewhat misleading. As ascertained

by J.K. Rawat, Director of Forest Research

Institute, the book describes 60 aromatic

plants.

The following subjects are discussed

under each plant description: 1. Origin and

96

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

REVIEWS

History, 2. Habit, 3. Morphology, 4. Phenology,

5. Distinguishing field characters,

6. Distribution, 7. Access and vulnerability,

8. Climate, 9. Soil types, 10. Silvicultural

requirements, 11. Silvicultural characters,

12. Propagation, 13. Harvesting, 14. Grading

and processing, 15. Storage, 16. Value addition,

17. Substitutes and adulterants, 18. Physico-

chemical properties, 19. Active principles,

20. Production, 21. Uses, 22. Marketing and

trade channels, 23. Export, 24. Recommendation

and future vision, 25. References.

There is a lack of good books on aromatic

plants of India. But in my opinion this book is

far below the required standard for such books.

Many important aromatic plants are missing

which makes the scope of the book very limited.

Some of the important aromatic plants missing

are: Murray a koenigii , Bothriochloa odor at a,

Capillipedium asimilis , Canarium striclum and

Vateria indica.

Many references are incomplete and one

wonders if the authors have referred to the

publications cited or taken the information from

secondary sources, e.g. Bacon (1909): Philipp.

J. Sci. 4A, 131. (p. 100); Gupta (1964): Ind.

Forester 90 - pp. 459 (p. 134)

Both the references do not give the initials

of the author’s name. The second reference

mentions pp. (pages) and gives only one page 459.

The manuscript has not been properly read

before publishing it. Under “Vanilla” the

botanical name Vanilla periflora Andr. with

synonym V fragrans Ames is listed, but the title

of the figure (not so good) is given as Vanilla

planifolia Andr. The author of the Tropical

American orchid Vanilla planifolia is Jackson

(not Andrews). The availability of these two

Vanilla species in the wild as mentioned by the

authors requires validation.

All the species appear randomly in the

book with no specific system being followed. In

some places, the order of appearance has been

based on English names like Mint, Patchouli,

Rosemary, Saffron, Sandalwood; while in the

case of Ajowan, Ajmod, Am(b)a-haldi, etc. they

have been arranged according to their Hindi

names, while in still others like Matricaria

chamomilla they have been arranged according

to their scientific names. This has resulted in

species of the same genera being separated by a

number of pages, (e.g. Curcuma amada on p. 30

and C. aromatica on p. 314; Cymbopogon

citrates on p. 204, C. jwarancusa on p. 1 77 and

C. nardus on p. 110 all arranged according to

English names).

The Index lists some dubious names like

Abelmoschus agallocha (p. 13). This name

perhaps refers to Aquillaria agallocha.

Cymbopogon nardus Rendle has been cited

as a synonym under C. winter ianus Jowitt (see

p. 1 10). However, these two grasses are distinct

species (not conspecific). Cymbopogon

winter ianus Jowitt is a grass having lower glume

with three definite intracarinal nerves, while

typical C. nardus (L.) Rendle is without them. If

due to some reason (e.g. due to variability to a

great extent) these two species happen to be the

same (as they both originally come from Sri

Lanka) Cymbopogon nardus (L.) Rendle has

priority and should be the correct name of this

grass. Incidentally, native Indian grass cultivated

in Tamil Nadu and elsewhere in the country is

Cymbopogon nardus (L.) Rendle var.

confertiflorus (Steud.) Stapf ex Bor.

The authors do not seem to be aware of

the International rules on botanical

nomenclature. For example: Acorus calamus (L.)

Syzygium aromaticum (L.) Citrus Union (L.).

When the author(s) name is cited in parenthesis

it should always accompany the name(s) of the

other author(s) outside the parenthesis. The name

in the bracket indicates that the species was first

published under another generic name, while the

name outside parenthesis indicates the author

who placed the taxon in its proper taxonomic

binomial position. I do not wish to quote other

nomenclatural errors in the book. It would suffice

JOURNAL. BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

97

REVIEWS

here if we agree that no matter how much

information you gather, if it is reported under an

erroneous name it will serve no purpose.

The photograph of Coleus aromaticus

(now Solenosternon amboinicus ) has been

wrongly labeled as Trachyspermum amrni

(Ajowan).

The book lacks a good index. The species

index on p. 339 includes 111 binomials

(including some synonyms); a number of specific

names dealt in the book are missing in the index.

Carum copticum given as p. 81 in the index is

present on p. 23 (not given in the index) under

synonyms of Trachyspermum ammi (L.)

Sprague.

Considering the secondary information

provided in this book, which can be retrieved

from volumes of wealth of india among other

books, the price of the book at Rs. 1,950/- is not

justifiable by any standard.

■ M.R. ALMEIDA

4. RAPTOR WATCH: A GLOBAL DIRECTORY OF- RAPTOR MIGRATION SITES

Compiled and edited by Jorje I. Zalles and Keith L. Bi'.dstein, 2000. BirdLife

Conservation Series No. 9, BirdLife International, Cambridge, U.K. Pp. 419, 22 black

and white photos, 21 tables, 21 figures, 3 appendices (24 x 17 cm). $ 58.00.

A global conservation undertaking was

kick-started in 1988 called ‘Hawks Aloft

Worldwide’. More than 800 raptor biologists and

conservationists worldwide in more than 100

countries on six continents provided information

on known and potential raptor migration sites.

This compendium is based on the information

submitted, and provides the information in a

readable, accessible manner.

The Introductory Chapters mention species

of global conservation concern as listed by

BirdLife International, the regional origin of

breeding populations of migratory species,

countries with taxa that are of conservation

importance, and continental distribution and

migration status (full vs. partial vs. irruptive

migrants) of ail migratory species. In the ‘Global

Analyses’ Chapter, one is told that at least 183

(62%), and possibly 193 (67%) of the world’s

294 species of raptors engage in seasonal

migration. This involves full migrants 19 species

(6% of all raptors) in which at least 90% of ail

individuals leave the breeding area during the

non-breeding season. Around 100-104 species

(approximately 34%) are partial migrants, some

of which range between breeding and non-

breeding areas. Local or altitudinal migrants

comprise 43-60 species (15% plus), their

movements triggered by food availability or local

weather conditions. Asia has the highest number

(66) of migratory raptor species followed by

Africa (6 1 ). Furthermore, 28 (37%) of the world’s

75 Near Threatened, Vulnerable, Endangered

and Critically Endangered raptors are known

migrants. Their protection, which involves

international networking, co-operation,

combined political will and the generation of long

term data bases, will prove to be a difficult,

protracted, but challenging task.

We are informed that 388 raptor migration

watch sites have been identified worldwide, 252

occurring in protected areas. Furthermore, each

observation or watch site that has annually

recorded a minimum of 1 0,000 migratory raptors

is collectively tabulated. The book contains

sufficient information for the reader to decide

where to go to observe migratory raptors, or

where a species is most likely to be seen.

The main part of the book consists of

country and watch site descriptions under

regional headings such as Africa; Asia, the

Middle East, Australia and the Pacific Islands;

Europe; South America, Central America, the

Caribbean and Mexico; North America. Country

98

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

REVIEWS

descriptions contain information on the size

(sq. km), length of coastlines, neighbouring

countries, human population size and growth

rate, per capita GNP, major land-use patterns,

whether party to International Wildlife Law

(UNESCO, Ramsar Convention and CITES),

biogeographical provinces, and total number of

migratory raptors (full and partial migrants). The

watch site descriptions are extensive, providing

facts on location with coordinates, altitude,

biogeographic province, site description, land

tenure, protection status, land use, various threats

if any, migration monitoring activity, main

migratory periods, regular migratory raptor

species, research and conservation activities

undertaken, list of contacts and resource

personnel for the site, and criteria for inclusion

of the site in the global directory.

A large country like North America has

129 watch sites, China 5, a smali country like

Israel 8, and India, sadly none. Smaller countries

like Indonesia have 7, Japan 6, Malaysia 2 and

Nepal 2 watch sites respectively. Raptor

migration in India is poorly known and little

information exists, for example, on the lesser

kestrel and Amur falcon, which migrate through

India in huge numbers. The book highlights the

lack of information on Autumn and Spring

migration over the Indian Himalaya. Raptors

migrate on a broad front of 1,000 km over the

Himalaya, but watch sites need to be identified

and established where climate and geography

create conditions and ‘corridors’ in the form of

N-S oriented river valleys such as the upper Kali

Gandaki in Nepal.

Appendix 1 lists major National (US) and

International raptor organizations and Appendix

2 cites countries that have ratified the convention

on biological diversity (CBD). The last eight

pages are an Index of migratory raptors at Raptor

Watch Global Directory (RWGD) watch sites

with keys to migratory status, two- letter country

codes and names for different watch sites and

number of watch sites at which a particular

species occurs.

In conclusion, raptor watch is well-

produced, both in content and production

that took over 10 years to complete. An ambitious

and pioneering project, it has great conservation

potential through setting in motion conservation

efforts and tourism promotion for watching

migrating birds of prey. For example, more

than 90,000 enthusiasts visit Hawk Mountain

Sanctuary in North America annually. The book

provides quick reference on the distribution

and conservation status, highlights areas/

countries where intensive work is required and

addresses pressing problems. St will help

develop monitoring programmes to track

migrating raptor populations and determine

fluctuations and causes over a long term

period. Besides promoting further research, it

should help to motivate greater international

collaboration in raptor conservation. The book

would also encourage serious raptophiles

to witness the spectacle of migration and

experience wild raptors at close quarters as

volunteers through trapping and banding

programmes. Before migrating raptors can be

protected, we need to know where, when and how

they migrate. This book is the first step in that

direction.

B RISHADNAOROJI

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

99

MISCELLANEOUS NOTES

1. THE HARVARD COLLECTION OF SOUTH ASIAN MAMMALS

Founded in 1859, the Museum of

Comparative Zoology (MCZ) at Harvard

University in Cambridge, Massachusetts,

possesses the oldest systematic mammal

collection in the United States. While not ranked

among the larger mammal collections in terms

of number of specimens housed, the Mammal

Department of the MCZ stands as one of the most

complete collections in the world, in terms of

taxa represented in its holdings.

In a recent review of South Asian

mammals in the collection of the MCZ, we have

identified more than 700 specimens representing

160 different species from the South Asian region

(here taken to include Pakistan, India including

the Andaman and Nicobar Islands, Sri Lanka,

the Maldives, Nepal, Bhutan, Bangladesh and

Upper Myanmar). The species represented are

listed in Table 1. The intention is to inform

researchers, especially those in the South Asian

Table 1 : List of mammal species from South Asian localities in the Museum of Comparative Zoology

at Harvard University

Insectivora

Hemiechirius collaris

Hemiechinus micropus

Crocidura horsfieldi

Suncus etruscus

S uncus mon tan us

Suncus murinus

Suncus stoliczkanus

Nectogale elegans

Soriculus caudatus

Soriculus nigrescens

Euroscaptor micrura

Indian long-eared hedgehog

Indian (Pale) hedgehog

Horsfield's shrew

White-toothed pygmy shrew

Sri Lanka highland shrew

Asian house (Grey musk) shrew

Anderson’s shrew

Elegant water shrew

Hodgson’s brown-toothed shrew

Himalayan shrew

Himalayan (Eastern) mole

Scandentia

Tupaia belangeri Northern (Malay) tree shrew

Chiroptera

Pteropus giganteus

Pteropus melanotus

Rousettus leschenaulii

Rhinopoma hardwickei

Rhinopoma microphyllum

Saccoiaimus saccolaimus

Taphozous longimanus

Taphozous meianopogon

Taphozous nudiventris

Megaderma lyra

Megaderma spasma

Rhinolophus ferrumequinum

Rhinolophus iepidus

Rhinolophus rouxi

Hipposideros atra

Hipposideros fulvus

Hipposideros lankadiva

Hipposideros pomona

Hipposideros speoris

Kerivoula picta

Indian flying fox

Black-eared flying fox

Leschenault’s rousette (Fulvous fruit bat)

Lesser mouse-tailed bat

Greater mouse-tailed bat

Naked-rumped pouched bat

Long-winged tomb (Longarmed sneathtaii) bat

Black-bearded (Bearded) sheathtail bat

Naked-rumped tomb bat

Greater (Indian) false vampire bat

Lesser false vampire bat

Greater horseshoe bat

Blyth’s horseshoe bat

Rufous horseshoe bat

Dusky roundieaf bat

Fulvus roundieaf bat

Indian roundieaf bat

Pomona roundieaf bat

Schneider’s roundieaf bat

Painted bat

100

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

MISCELLANEOUS NOTES

Table 1 : List of mammal species from South Asian localities in the Museum of Comparative Zoology

at Harvard University ( contd .)

JOURNAL , BOMBAY NATURAL HISTORY SOCIETY 100(1), APR. 2003

101

MISCELLANEOUS NOTES

Table 1 : List of mammal species from South Asian localities in the Museum of Comparative Zoology

at Harvard University ( contd .)

102

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

MISCELLANEOUS NOTES

Table 1 : List of mammal species from South Asian localities in the Museum of Comparative Zoology

at Harvard University ( contd .)

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

103

MISCELLANEOUS NOTES

during the 1860s and 1 870s with their donations

of large natural history collections from India.

Another period of significant growth in

accessions from South Asia, especially of bats

and primates, occurred under the enthusiastic

tenure of the great mammalogist G.M. Allen as

Curator of the MCZ, Mammal Department

(during 1 924- 1 942). More recent additions to the

collection have included specimens procured

during expeditions to Nepal and Pakistan in the

1950s.

There is one name-bearing type series of a

South Asian mammal at the MCZ — the holotype

and paratype of Pteropus ariel (= P. giganteus

arid) described by G.M. Allen (1908) from the

Maldives (see Helgen and McFadden, 2001 : 141).

The department holds topotypes of many

mammalian taxa from the region, including

Buclorcas taxicolor , Capra falconeri

cashmiriensis, Soriculus c and at us, S . nigrescens,

Niviventer niviventer monticola , Niviventer eha

and Miniopterus pusillus. Additionally, the

collection also contains specimens of a large

number of South Asian mammals that are rarely

collected.

In the list, taxonomy follows Wilson and

Reeder (1993); common names follow Wilson

and Cole (2000); where these names deviate from

Prater (1948), that author’s common names are

given in parentheses.

December 22, 2002 KRISTOFER M. HELGEN

Mammal Department,

Museum of Comparative Zoology,

Harvard University, 26 Oxford Street,

Cambridge, MA 02138, USA.

JOHN MATHEW

Entomology Department,

Museum of Comparative Zoology,

Harvard University, USA.

Email: [email protected]

CHRISTINE A. MONTA

Department of History of Science,

Harvard University, USA.

References

Allen, G.M. (1908): Notes on Chiroptera. Bull. Mus. Wilson, D.E. & F.R. Cole (2000): Common names of

Comp. Zool. 52: 25-63. mammals of the world. Smithsonian Institution

Helgen, K.M. & T.L. McFadden (2001): Type specimens Press, Washington, D.C.

of recent mammals in the Museum of Comparative Wilson, D.E. & D.R. Reeder ( 1 993): Mammal species of

Zoology. Bull. Mus. Comp. Zool. 157 : 93-181. the world: a taxonomic and geographic reference.

Prater, S.H.( 1948): The Book of Indian Animals. Bombay 2nd edn. Smithsonian Institution Press,

Natural History Society, Mumbai. Pp. 324. Washington, D.C.

2. LITTLE GREBE TACHYBAPTUS RUFICOLLIS : AN ADDITION

TO THE AVIFAUNA OF LADAKH

On the morning of July 1, 1999 between

1000 and 1030 hrs, three adult little grebe

Tachybaptus ruficollis in summer plumage were

observed at Trishul Tso, a small lake created by

the Army, near the headquarters of the Ladakh

Scouts on the Leh-Srinagar road. The birds were

immediately recognised as little grebe

Tachybaptus ruficollis , a species familiar to the

observers. On July 3, 1999 only one bird was

recorded at the lake. HSS checked for the birds

again at the lake on July 9, 1999, but none were

sighted.

There are no previously documented

records of little grebe from Ladakh to the best of

our knowledge. Ali and Ripley (1981), Grimmelt

et al. (1998) and Kazmierczak and van Perlo

(2000) do not mention this area in the species’

distribution.

104

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I

MISCELLANEOUS NOTES

Jayson, E.L. et al. : Fregata arid Gray

Plate 1

Fig. la-b: Lesser frigatebird Fregata ariel Gray in the Kole wetlands

of Thrissur, Kerala

106

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

MISCELLANEOUS NOTES

The upper limit of the species is variously

recorded as 1,800 m (Ali and Ripley 1981),

2,500 m (Ali 1996) and 1,372 m (Fleming et al

1984) for the Indian subcontinent. Interestingly,

the site where the birds were observed is at an

elevation of c. 3,500 m, considerably exceeding

the record from Rara Lake in Nepal on April 1,

1976 at 3,050 m (Inskipp and Inskipp 1991).

Feb. 7, 2001 HARKIRAT SINGH SANG HA

B-27, Gautam Marg,

Hanuman Nagar,

Jaipur 302 021, Rajasthan, India.

Email: [email protected]

References

RISHADNAOROJI

Godrej Bhavan,

4-A, Home Street,

Fort, Mumbai 400 001,

Maharashtra,

India.

Email: [email protected]

MAAN BARUA

107, M.C. Road,

Uzan Bazaar,

Guwahati 781 001,

Assam,

India.

Au, S. (1996): The Book of Indian Birds. 12th revised

and enlarged edn. Bombay Natural History Society,

Mumbai.

Ali, S. & S.D. Ripley (1981): Handbook of the Birds of

India and Pakistan. Vol. 1, 2nd edn. Oxford

University Press, New Delhi.

Fleming, Sr. R.L., R.L. Fleming Jr. & L.S. Bangdel

(1984): Birds of Nepal. 3rd edn., Avalok,

Kathmandu.

Grimmett, R., C. Inskipp & T. Inskipp ( 1 998): Birds of the

Indian subcontinent. Christopher Helm, London.

Inskipp, C. & T. Inskipp (1991): A Guide to the Birds of

Nepal. 2nd edn. Christopher Helm. London.

Kazmierczak, K. & B. van Perlo (2000): A Field Guide

to the Birds of the Indian Subcontinent. Pica Press.

Robertsbridge. U.K.

3. SIGHTING OF LESSER FRIGATEBIRD FREGATA ARIEL GRAY

IN THE KOLE WETLANDS OF THRISSIJR, KERALA

( With one plate )

Kole wetland in Thrissur district is one of

the important wintering grounds for migratory

birds coming to Kerala. The area lies in between

10° 20' - 10° 40' N and 75° 58' - 76° 11' E,

extending to about 1 1 ,000 ha. Kole wetlands are

situated below sea level and paddy is cultivated

during the months of October to April, after

draining water. During monsoon (June to

September) the whole area is inundated. As a part

of ecological studies on wetland birds initiated in

1998, we have been surveying the area regularly.

During the period, a lesser frigatebird Fregata ariel

Gray was recorded from Guruvayur, Thrissur

district on June 16, 2000. The place is situated on

the west coast and is about 4 km away from the

sea. Local people noticed a live bird and brought

it to the Thrissur Zoo. On examination, the bird

was identified as an adult female of the lesser

frigatebird. The bird had a black head and red eye

rings, black throat and white breast extending into

a complete collar around the neck. The bird stayed

alive for a week in the zoo, after which it was

skinned and preserved.

Only a few sighting details of the lesser

frigatebird are reported from Kerala State.

Ferguson and Bourdillon ( 1 904) recorded it from

Trivandrum. Ali (1984) has not reported the

species from Kerala. Faizi (1985) reported the

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

107

MISCELLANEOUS NOTES

species from Quilon, based on a museum

specimen. This is the first report of the species

from central Kerala and also from the Kole

wetlands of Thrissur. The individual seems to

be an accidental straggler that landed in the

coastal zone due to the heavy monsoon winds

prevalent in June.

Refer

Ali, S. (1984): Birds of Kerala. 2Md Edn, Oxford University

Press. Delhi. 444 Pp.

Faizi, S. ( 1 985): An additional record of the Lesser Frigate

Bird Fregcita arid in India. J. Bombay nat. Hist.

4. INDIAN POND-HERONS ARDEOLA

Following the failure of monsoon in our

area in 1 999-2000, Rishi Valley (Chittoor district,

Andhra Pradesh) wore a drab look with the

drying up of ponds and other waterbodies. Even

the percolation tank, which usually harbours

water even in the lean period, was reduced to a

muddy puddle.

I was out on a walk along the road lined

with trees such as Ficus and Spathodea outside

the school campus, on the evening of March 5,

2000, when 1 noticed three Indian pond-herons

(Ardeola grayii ) perched on a banyan tree Ficus

bengalensis , c. 8 m above the ground. It was too

early for them to roost and there was no major

disturbance that might have forced them to fly

to a tree. I paused to look around, and within a

few minutes was able to find the reason.

Just above the Indian pond-herons was a

small group of dragonflies, flying close to the

foliage of the banyan tree. Even as I was watching

them, an Indian pond-heron attempted to snap

at a dragonfly that was just a few inches above

it. This and the attempts that followed were all

unsuccessful for the few minutes that I observed

them. A little later, the dragonflies moved away

from the tree and the Indian pond-herons flew

to forage in the dry, barren fields. A few days

later, I noticed another Indian pond-heron on a

November 25, 2000 *E. A. JAYSON

C. SIVAPERUMAN

Division of Wildlife Biology

Kerala Forest Research Institute

Peechi 680 653,

Kerala, India.

* Email: [email protected]

E N C E S

Soc. 82(1): 191.

Fergusson, FI.S. & T.F. Bourdillon (1904): The Birds of

Travancore, with notes on their Nidification.

J. Bombay nat. Hist. Soc. 16: 1-18.

GRAYII FEEDING ON DRAGONFLIES

tamarind tree outside my house attempting to

catch dragonflies in flight.

With the disappearance of waterbodies, the

Indian pond-herons had to change their foraging

strategies and had taken to vantage points on trees

to catch dragonflies. A few others were seen

stalking insects in dry fields and, following cattle

and goats just like cattle egrets ( Bubulcus ibis )

which were also occasionally seen with the herons.

I had, on an earlier instance, noticed Indian pond-

herons attempting to catch dragonflies from the

ground at Adyar Estuary in Chennai when the

insects were flying low over the islets and water.

Prasad and Hemanth (1992; JBNHS89 : 246) have

reported Indian pond-herons perched on a 7 m

willow tree in Bangalore, feeding on bees that came

to the flowers. Jose noticed (1999; NLBW 39(2):

39) cattle egrets feeding on insects visiting

flowerheads of mango Mangifera indica and

Carissa trees. These observations show how birds

adapt their behaviour to changed circumstances

to survive.

April 16, 2000 V. SANTHARAM

Institute of Bird Studies & Natural History,

Rishi Valley Education Centre,

Rishi Valley 517 352,

Chittoor district, Andhra Pradesh, India.

108

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MISCELLANEOUS NOTES

5. VARIATION RECORDED IN THE IRIS, BILL, LEG AND FOOT COLOURATION

IN CATTLE EGRET BUBULCUS IBIS

A nesting colony of cattle egret Bubulcus

ibis was spotted at Zila Garden, Rajkot. They

started nesting during early June, 2000. During

regular observations, I recorded variations in four

birds of the colony. The iris and bill colour of al!

the breeding birds in the colony was golden

yellow, but in these four birds, the iris was red

and bill reddish-violet. This variation among

breeding populations of cattle egret has not been

recorded by Ali and Ripley (1987), Heinzel et

al. (1973), and Sonobe and Usui (1993).

The breeding birds had pink legs and feet,

while the non-breeding birds had black legs and

feet. The actual changing of colours was not

recorded. (The author has submitted photographic

evidence of the observations — Eds.)

February 6, 2001 TUSHAR M. SANGHANI

“Tushar”

1, Maruti Nagar,

Aerodrome Road,

Rajkot 360 001, Gujarat, India.

References

Ali, S. & S.D. Ripley (1987): Compact Handbook of the

Birds of India and Pakistan together with those of

Bangladesh, Nepal, Bhutan and Sri Lanka. 2nd edn,

Oxford University Press, Delhi, pp.737 + 1 04 plates.

Heinzel, H.. R. Fitter & J. Parslow ( 1 973): The Birds of

Britain and Europe. William Collins Sons and Co.

Ltd., London.

Sonobe, K. & S. Usui (Eds.) ( 1 993): A Field Guide to the

Waterbirds of Asia. Wildbird Society of Japan,

Tokyo.

6. RECORD OF A NESTING COLONY OF PAINTED STORK

MYCTERIA LEUCOCEPHALA AT MAN-MARODI ISLAND IN THE GULF OF KUTCH

{With one text-figure )

Gujarat is well endowed with nesting sites

of painted stork Mycteria leucocephala , both in

the vicinity of freshwater inland wetlands and

on the coast. Dharmakumarsinhj i (1955)

observes that these birds nest “more numerously

on the coastal areas where marine food is easily

available”. However, very little is known about

the coastal nesting grounds of the painted stork

in Gujarat, with the exception of the heronries

at Bhavnagar that have been well documented

by Parasharya and Naik (1990). In this note, I

wish to record the existence of a large nesting

colony of painted stork on the Man-Marodi Island

in the Gulf of Kutch. This site was studied while

I was accompanying teams of students from the

Centre for Environment Education, Ahmedabad,

to the Sundervan campsite at Beyt Dwarka during

1999-2000.

Man-Marodi is a rocky island quite close

to the coast, off Positra, near the mouth of the

Gulf of Kutch (Fig. 1). The island itself appears

as a large block of rocks some 2-3 sq. km in area,

rising at least 20 m above the level of the water.

The chief vegetation on this island is scrub and

small trees such as Acacia and Salvador a,

interspersed by dense stands of Euphorbia. On a

first visit to Man-Marodi on October 20, 1999,

at least 1 00 nests of painted stork were observed.

Most of the nests had 1 or 2 chicks. The nestlings

were in an advanced stage, possibly a month old

as suggested by the colour of their plumage and

bill, indicating that nesting activity at this site

must have started in end August i.e. towards the

end of the monsoon. Since the principal

vegetation of the island is quite short, most of

the nests were observed at c. 1.5 m above ground

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

109

MISCELLANEOUS NOTES

69°

Fig. 1: Map of Man-Marodi Island (Indicated by

arrow) in the Gulf of Kutch

level and quite closely packed. Interestingly,

some nests were made by stacking sticks and

twigs on Euphorbia stands. On November 30,

2000, when this island was visited again, no

painted storks were seen, although information

gathered from local fishermen revealed that the

birds had nested during that year too.

This heronry has, of course, not gone

unnoticed. In his exhaustive review on the birds

of Gujarat, Khacher (1996) refers to Man-Marodi

without actually naming it when he writes,

“There is an unusual nesting colony of the

painted stork on a sandstone island in the Gulf

of Kachchh”. Possibly, what makes Man-Marodi

unusual is that out here, in the absence of true

thermals in the sea, painted stork appear to take

advantage of the wind hitting the rock face,

which creates the effect of thermals and enables

the birds to soar in the sky and gain altitude when

embarking on food finding missions (Lavkumar

Khacher, pers. comm.). The use of Euphorbia

Refe

Dharmakumarsinhji, R.S. (1955): The Birds of Saurashtra.

Times of India Press, Bombay.

Khacher, L. ( 1 996): The Birds of Gujarat - — A Salim Ali

Centenary Year Overview. J. Bombay nat. Hist. Soc.

93(3): 331-373.

Parasharya, B.M. & R.M. Naik(1990): Ciconiiform birds

as a nesting substrate is another interesting

feature of this heronry.

Why is this island preferred by large

numbers of painted stork for nesting? Firstly, the

closely packed vegetation affords a good substrate

for nesting in colonies. Secondly, the Island being

surrounded by the sea on all sides and not being

visited by humans too frequently, the birds

probably find it to be a safe place. But having

said this, I may also mention the problems. For

instance, common crows take their toll on painted

stork eggs. Reportedly, jackals sometimes visit

this island, swimming across from the mainland

or from the neighbouring islands during low tide,

and can pose a danger for the nestlings. Thirdly,

although no humans are known to inhabit this

island as of now, local fishermen sometimes

anchor their boats in the vicinity of Man-Marodi

and reportedly, poach upon the nestlings.

Acknowledgements

I thank Mr. Lavkumar Khacher for

interesting insights about Man-Marodi Island

and Mr. Rajindersinh Jadeja and Mr. Hembha

Vader for providing logistical support, including

a motorboat to visit the islands. I am grateful to

Shri K.V. Sarabhai for encouragement.

March 22, 2001 ABDUL JAMIL URFI

Sundarvan Nature Discovery Centre,

S.M. Road, Jodhpur Tekra,

Ahmedabad 380 015, Gujarat, India.

Permanent Address: c/o Dr. Jalil

A-270, Jamia Nagar,

Okhla, New Delhi 110 025, India.

E N C E S

breeding in Bhavnagar city, Gujarat: a study of their

nesting and plea for conservation. Pp. 429- 445.

In: Conservation in Developing Countries: problems

and prospects. Proceedings of the Centenary Seminar

of Bombay Natural History Society (Eds: Daniel. J.C.

& J.S. Serrao). BNHS, Bombay.

110

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MISCELLANEOUS NOTES

7. FIRST SIGHTING OF LESSER ADJUTANT-STORK LEPTOPTILOS JAVANICUS

FROM SANJAY GANDHI NATIONAL PARK, MUMBAI

A solitary adult lesser adjutant-stork

Leptoptilos javanicus was sighted at Vihar lake,

Sanjay Gandhi National Park (SGNP), Mumbai,

on January 21, 2001 at 1000 hrs. When first

sighted, the bird was searching for food at the

bank of the lake. However, due to human

disturbance, it took off, soared for a while, settled

on a bare tree for 10 min and then resumed its

feeding activities. While in the air, it was mobbed

by three black kites Milvus migrans. However,

the bird looked undeterred by these raptors.

Subsequently, local birdwatchers sighted it thrice

in six days, the last being on January 27, 2001.

According to Ali and Ripley (1987),

L. javanicus is resident, nomadic (during

monsoon) and locally migratory. Apart from

Assam, which is its stronghold, it is scattered

across West Bengal, Bihar, Orissa, Kerala, Tamil

Nadu, Andhra Pradesh, Uttar Pradesh, Delhi,

Kutch, Rajasthan, Nepal, Sri Lanka and

Bangladesh. Lainer (1999) and Choudhury

(1995) have also reported the bird from Goa and

Rf.f

Ali. S & S.D. Ripley (1987): Compact Handbook of the

Birds of India and Pakistan together with those of

Bangladesh. Nepal, Bhutan and Sri Lanka. 2nd edn,

Oxford University Press, Delhi, pp. 737+ 104 plates.

Choudhury, A. (1995): Bird survey of Dibru-Saikhowa

Wildlife Sanctuary. OBC Bull. 22: 15.

del Hoyo. J., A. Elliot & .1. Sargatal (1992): Handbook

Arunachal Pradesh respectively. Though widely

distributed, it occurs in very low densities along

its entire range except Assam, and has been

designated as a 'globally threatened species’ (del

Hoyo et al. 1992; Grimmett et al. 1998).

The breeding season of L. javanicus

stretches from July to January (Grimmett et al.

1998) and thus the appearance of an adult bird

at SGNP in January indicates that it might be

breeding in northern regions of the Western

Ghats. The poor monsoon during 2000 and the

subsequent water scarcity may also have forced

individuals to travel greater distances in search

of food. A detailed study of its probable range is,

however, essential to be able to comment on its

breeding in northwestern Maharashtra.

May 4, 2001 ANISH ANDHERIA

2, Sagar Building,

V.P. Road, Andheri (West),

Mumbai 400 058, Maharashtra, India.

Email: [email protected]

R E N C E S

of Birds of the World, Vol. 1. Lynx Edicions,

Barcelona.

Grimmett, R., C. Inskipp & T. Inskipp (1998): Birds of the

Indian Subcontinent. Oxford University Press,

Delhi.

Lainer, H. (1999): The Birds of Goa (Pt I). J. Bombay

nat. Hist. Soc. 96(2): 203-220.

8. BLACK IBIS PSEUD1BIS PAPULOSA FEEDING ON FROGS FROM CRAB HOLES

On November 5, 6 and 8, 1999, at about

0700 hrs on the K. Sathanur Road, Kalaignar

Karunanidhi Nagar, Trichy, Tamil Nadu, I saw a

pair of black ibis Pseudibis papillosa running into

shallow rainfed pools and beating their wings. Only

on November 9, on closer examination, did I realise

that the splashing of water and beating of wings,

probably by the same pair, had frogs jumping out

of the water into crab holes, situated on the margin

of the dried portion of the pool. The crab holes

were freshly formed. The ibis would thrust their

in-curved bills fully into the crab holes, only the

warts on their head visible, spreading out their

muddy red legs and pull a frog out in c. 1-2 min.

Once the bird had pulled the frog out, it would

throw it on the ground, stab it with its bill and

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

111

MISCELLANEOUS NOTES

then eat it. One bird ate about 10 frogs in 23 min

in this manner. Tadpoles that jumped out and hid

within the hexagonal cracks of the dried patches

of loam were not eaten by the ibis. I have seen

black ibis infrequently in Trichy. They do not breed

around here.

Nov. 14, 2000 J. MANGALRAJ JOHNSON

No. 11, 1 0th Avenue,

Ashok Nagar,

Chennai 600 083,

Tamil Nadu, India.

Email: [email protected]

9. NOTE ON BREEDING OF ANDAMAN TEAL ANAS GIBBERIFRONS

IN SOUTH ANDAMAN ISLANDS, INDIA

The Andaman teal Anas gibber ifrons

occurs in the Andaman Islands and Great Cocos

Islands. The species is not globally threatened,

but is considered ‘threatened’ in the Andaman

Islands because of agricultural development and

disappearance of wetlands (Andrews and

Whitaker 1994; Vijayan et al. 2000). However,

very little information was available on its status

and population until recently (Vijayan 1996;

Vijayan and Sakthivel 1996; Vijayan et al. 2000).

We report here an instance of breeding of the

Andaman teal from the Mahatma Gandhi Marine

National Park (MGMNP), Wandoor in the South

Andaman Island.

MGMNP is spread over c. 281.5 sq. km

and is comprised of 1 5 islands of the Labyrinth

group, along the southwest coast of South

Andaman Island. Only 60 sq. km of the Park is

covered by landmass and the rest comprises open

sea and creeks. The presence of marshy area and

creeks in this Marine National Park provides an

ideal breeding habitat for the Andaman teal

(Harry Andrews pers. comm., Vijayan et al.

2000). On May 28, 1999, while looking for sea

snake nests in and around the Redskin Island,

we approached a small, unnamed islet about 50

to 75 m away from the northeastern coast of

Redskin Island. The islet is about 2-3 m high

from the high tide line, about 7 m long and 3 m

wide. On approaching the islet we flushed two

teals, which we later identified as the Andaman

teal (Anas gibber ifrons). We searched the islet

and found a neatly placed nest, with nine

creamish eggs with small black spots all over, at

the base of a small shrub at the northern tip of

the islet. The nest was c. 20 cm in diameter and

consisted of dried grass, thin twigs and down

feathers of the adult. The nest was easily

accessible to human as well as water monitor

lizard (Varanus satvator andamanensis). Before

landing on this islet, we had sighted a water

monitor lizard basking on a small shrub, but it

disappeared into the water on our arrival. A little

away from the teal nest, four nests of black-naped

tern (Sterna sum air ana) were noticed. Each of

them had one or two eggs that were white with

prominent black spots. When we left the islet

we saw the parent birds return to the nest. The

next day, when we returned to the islet to

photograph the nest of the Andaman teal, eight

of the nine eggs had hatched and the ninth chick

was emerging when we reached. The newly

hatched chicks were grey, with a faint white circle

around the eyes, and a grey beak. The available

definite breeding records to date are from August

to October (Ali and Ripley 1983; Vijayan el al.

2000).

July 13, 2001 SARANG KULKARNI

Wildlife Institute of India,

PB # 18, Chandrabani,

Dehra Dun 248 001, Uttaranchal, India.

Email: [email protected]

MANISH CHANDI

Andaman and Nicobar Environmental Team,

PB # 1, Junglighat, Port Blair 744 103,

India.

112

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MISCELLANEOUS NOTES

References

All S. & S.D. Ripley (1983): Compact Handbook of the

Birds of India and Pakistan together with those of

Bangladesh, Nepal, Bhutan and Sri Lanka Oxford

University Press, Delhi. 737 pp.+ 104 plates.

Andrews. H.V. & R. Whitaker (1994): Preliminary

observation on the Andaman Teal (Anas gibberifrons

albogularis ) in North Andaman Island. Report

submitted to Asian Wetlands Bureau. Malaysia.

Vijayan, L. (1996): Status and conservation of the

Andaman teal Anas gibberifrons albogularis. In:

Proc. Anatidae 2000 Conference, Strasbourg, France,

5-9 December 1994. Gibier Faune Sauvage, Game

Wildlife 13(1): 831-842.

Vijayan, L. & R. Sakthivel (1996): Surveys of Andaman

Teal in winter 1995/96. Threatened Waterfowl

Research Group New/etter 9: 25-27 .

Vijayan, L., R. Sankaran, K. Sivakumar & V. Murugan

(2000): A study on the ecology, status and

conservation prospectives of certain rare endemic

avifauna of the Andaman & Nicobar Islands. Final

Report. Salim Ali Centre for Ornithology and Natural

Flistory, Coimbatore. 184 pp.

10. SOME OBSERVATIONS ON NESTING OF BONELLES EAGLE

HIERAAETUS FASCIATUS (VIEILLOT)

During our regular treks for bird watching,

we have observed a rare phenomenon not

reported elsewhere. This year, for the third year

in succession, the same pair of Bonelli’s eagle

Hieraaelus fasciatus (Vieillot) has nested at the

same site, building their third nest on top of the

previous two nests. The total height of the nest

has now reached a massive 0.9 m; sticks of various

trees gathered from the vicinity have been used to

build this c. 1 m wide nest. The nest is at a height

of c. 12 m on a jambui Syzygium cumini (L.) tree

on the slope of Pavangadh, 962 rn above msi,

18 km northwest of Kolhapur city, and is lined

with fresh green leaves. Nesting at the same site

by the same pair is a phenomenon we have not

observed in other Bonelli’s eagle pairs elsewhere

in the surrounding hilly region.

March 23, 2001 DEEPAL1 KULKARN1

BANDA PEDNEKAR

85, Ruikar Colony, Kolhapur 416 005,

Maharashtra, India.

11. STATUS OF WHITE-BELLIED SEA-EAGLE HALJAEETUS LEUCOGASTER

IN RATNAG1RI DISTRICT, MAHARASHTRA

The white-bellied sea-eagle Haiiaeetus

leucogaster is thinly, but widely distributed and

is listed as vulnerable in the Indian red data book.

No data on the status and population of this

species is available. Its present status, along the

164 km coastline, in Ratnagiri district,

Maharashtra, is given on the basis of a survey

undertaken in the district during 1996-97 and

1997-98. We located 62 nests during our survey.

The identity of the bird was confirmed from the

HANDBOOK.

The species is resident along the seaboard

and offshore islands from about 19° N of Mumbai

down the west coast and up the east to

Bangladesh, Laccadive Is. (now Lakshadweep),

Sri Lanka, and Andaman & Nicobar Islands and

is vagrant in Gujarat (Ali and Ripley 1978).

We undertook a survey to determine the

present status and distribution of the white-

bellied sea-eagie in Ratnagiri district, as the

species is found throughout the coastline of this

district.

Ratnagiri district is situated in the Konkan

region of Maharashtra State (16.30°- 18.04° N

and 73.02°-73.52° E). The coastline of the

district is about 164 km long and is uniformly

rocky and shallow. Various bluffs and

promontories enclose small sandy beaches,

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

113

MISCELLANEOUS NOTES

interspersed by estuaries of the more important

rivers and the mouths of numerous minor

streams. Average width of the district is about

64 km. We surveyed the coastal line, which is

5 to 7 km wide.

The Konkan region experiences a hot and

humid maritime climate with plentiful rain

during the monsoon (June to September ave.

3,000 mm). The mean annual temperature ranges

from 22 °C to 30 °C. The mean relative humidity

is 80% in the region.

The eagle that affects the seacoast and tidal

creeks and estuaries is seen occasionally a few

miles inland along tidal rivers and at freshwater

lakes (Ali and Ripley 1978).

In our survey, we saw most of the nests

along the seacoast, tidal creeks and estuaries. Of

the 62 nests recorded, 4 1 nests were found up to

0.5 km from the seacoast and 20 nests were found

from 0.5 km to 2 km. Only one nest was about

7 km away from the seacoast at Gavde Ambere,

Ratnagiri taluka along the Purnagad creek.

The nests were located while walking

along the coast and collecting information from

the iocals by showing them pictures from the

PICTORIAL GUIDE TO THE BIRDS OF THE INDIAN

subcontinent (Ali and Ripley 1995) and the book

of Indian birds (Ali 1996). All the nests were

visited in the nesting season and photographs

taken wherever possible.

According to Ali and Ripley (1978), the

breeding season is October to January. However,

we observed courtship and nest building from

mid-September to January. Well-grown chicks

were seen in the nest up to the end of March.

Nest size: Most nests located in Ratnagiri

district were 1 to 1 .5 m wide. At Rohile, Taluka

Guhagar the size of a 1 0 year old nest was 1 .7 m

x 1.2 m x 0.7 m.

Height: Most nests located were 20-40 m

high, in lofty trees. Only one nest was found

below 10 m i.e. 7.5 m at Velneshwar, Guhagar

taluka (Approximate measurements were

taken).

Table 1 : Nest height of the white-bellied sea-eagle in

Ratnagiri district

Nest: At Rohile, Guhagar taluka the nest

were made of sticks of Ficus bengalensis, Carissa

carandas , Leea macrophylla, Crota/aria

verrucosa , Loranthus longifolia, Bambusa

arundinacea and other unidentified tree species.

One nest stick was exceptionally long (86 cm)

and thick (2.86 cm).

Pieces of net, plastic paper, bits of woven

bag were also used. Green leaves of Mangifera

indica , Casuarina equisetifolia , Ficus

bengalensis , Grewia asiaticia , Smi/ax

macrophylla , Sapindus laurifolius, Bombax

malabaricum were used to line the nest.

The nesting trees of the eagle as observed

by us are given in Table 2.

Table 2: Tree species used for nesting

by Haliaeetus leucogaster

A = Mandangad; B = Dapoli; C = Guhagar; D = Ratnagiri;

E = Rajapur; T = Total

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MISCELLANEOUS NOTES

Table 3: Alternate nesting sites on same tree

A = Active Nest, I = Inactive Nest

Age: Local enquiry revealed the

approximate age of 44 nests out of a total 62

nests as follows: 21 nests of up to 5 years, 6 nests

of 5 to 10 years and 10 nests of 1 1 to 25 years.

Seven nests were 26 to 50 years old.

Alternate nesting sites: We observed

alternate nesting sites at three places, where both

the nests were in good condition, but were used

in alternate years. Details are given in Table 3.

Territory: Ali and Ripley (1978) define

territory as, “usually a single pair with vast

territory, but not uncommonly several pairs and

nests on the same island”.

In our survey, we have seen two nests

distant from each other. At times we have found

more than one nest in a small area. We sighted

3 nests (nest no. 17 to 19) in Guhagar, and 3

nests (nest no. 51 to 53) in Kashali village,

Rajapur taluka. The nests in both these villages

are hardly 1 km apart from each other.

Human-Bird Interaction: At Narvan,

locals said that the eagle sometimes take poultry.

At Jaitapur, local fishermen said that they get

indications from the eagle’s calls about

availability of fish in the sea and at times also

scare the eagle to drop its prey and eat the fish.

Names: Ali and Ripley (1978) give the

local name as Kankan. It is also referred to as

Sheshari in other regional (Marathi) books.

Besides this we have found some more local

names. In Ratnagiri district, this eagle is called

as ghar (kite) and not garud (eagle). The other

names used by locals are given in Table 4.

Table 4: Local names of the white-bellied sea-eagle

Hunting: Rare hunting incidents were

recorded by the locals. One was on the White

Sandy Beach at Ratnagiri, while two birds were

killed for sport about 4-5 years ago. In Tavsal,

Guhagar taluka one bird was killed for sport.

Mango Trees: Out of 62 nests in the

district, 20 nests are on Mango tree Mangifera

indica. The spraying of insecticides on mango

trees and nesting activity of eagles starts in

October. Due to fear of attack from eagles the

nests are first destroyed and then the spraying is

begun. This causes loss of eggs or chicks. Cutting

of trees also deprives the bird of suitable large

nesting trees.

Nest distribution in Talukas: We surveyed

the white-bellied sea-eagle in 5 talukas of

Ratnagiri district namely Ratnagiri, Guhagar,

Rajapur, Dapoli and Mandangad covering about

164 km of coastal area. Out of 62 nests, there

were 20 nests in Ratnagiri, 1 5 nests in Guhagar,

13 nests in Rajapur, 8 nests in Dapoli and 6 nests

in Mandangad.

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115

MISCELLANEOUS NOTES

A summary of the bird’s nesting activity

is given in Table 5.

Table 5: Summary of activity in 1 996-1 997 & 1 997-1 998

Particulars Activity in the year

1996-1997 1997-1998

Acknowledgements

This survey was supported by grant from

the Salim Ali Nature Conservation Fund, BNHS,

Mumbai. We thank Dr. Asad R. Rahmani,

Director, BNHS and Mr. Rishad Naoroji for their

comments on the report. We also thank Dr. S.

Asad Akhtar for perusing the manuscript.

February 26, 2001 V1SHWAS KATDARE

RAM MONE

Sahyadri Nisarga Mitra,

Near Laxminarayan Temple,

Chiplun 415 605, Ratnagiri district,

Maharashtra, India.

Reference

Ali, S. & S.D. Ripley (1978): Handbook of the Birds of India and Pakistan (Vol. 1). Oxford University Press, Delhi.

Pp 287-289.

12. CANNIBALISM IN INDIAN WHITE-BACKED VULTURE GYPS BENGALENSIS

. IN KEOLADEO NATIONAL PARK, BHARATPUR, RAJASTHAN

On May 22, 1997, at 0600 hrs, five Indian

white-backed vultures Gyps bengalensis and a

few crows Corvus splendens were observed

feeding on a small carcass in the Keoladeo

National Park, Bharatpur, Rajasthan. Some

crows and four vultures were sitting on the

ground close to the carcass waiting for their turn

to feed. The carcass appeared to be small in size

and could not be seen clearly as it was surrounded

by the scavengers. Through the spotting scope,

it appeared to be a carcass of a dark coloured

bird. On closer approach, we found that it was a

dead adult Indian white-backed vulture. It was a

fresh half-eaten carcass. After a few hours, by

0930 hrs, the carcass was almost finished.

The Indian white-backed vultures are

carrion eaters, which feed mostly on large

mammal carcasses (Ali and Ripley 1983). The

reason for cannibalism in Indian white-backed

vulture is not clear, but it was certainly not due

to scarcity of food, as there was an abundant

supply of food for vultures in the Keoladeo

National Park (Prakash 1999).

This observation was recorded when the

vulture mortality was at its peak, at the beginning

of the crash in the population of the Indian white-

backed vulture.

Cannibalism is observed in many predatory

birds such as barn owls Tyto alba , short-eared

owls Asio flammeus , Oriental honey-buzzards

Pernis ptilorhynchus and some sea birds. We

have not come across any reference to

cannibalism in the Indian white-backed vulture,

hence this observation is worth recording.

Acknowledgements

We are thankful to the Ministry of

Environment and Forests for funding and

sponsoring the project. Our sincere thanks are

due to the Rajasthan Forest Department for

permission to work in the Park. We are also

thankful to the Bombay Natural History Society,

especially Dr. A. R. Rahmani, Director, for

encouragement and valuable comments.

116

JOURNAL BOMBAY NATURAL HISTORY SOCIETY. 100(1). APR. 2003

MISCELLANEOUS NOTES

November 7, 2001 GARG1 RAN A

*VIBHU PRAICASH

Bombay Natural History Society,

Hornbill House, S.B. Singh Road,

Mumbai 400 023, Maharashtra, India.

Refer

Ali, S. & S.D. Ripley (1983): Compact Handbook of the

Birds of India and Pakistan together with those of

Bangladesh, Nepal, Bhutan and Sri Lanka. Oxford

University Press, Delhi, pp. 737 + 104 plates.

^Present Address: BNHS Fieid Station,

F-23, HMT Colony, Pinjore 134 101,

District Panchkula,

Haryana, India.

Email: [email protected]

E N C E S

Prakash, V. ( 1 999): Status of vultures in Keoladeo National

Park, Bharatpur, Rajasthan, with special reference

to population crash in Gyps species. J. Bombay not.

Hist. Soc. 96(3): 365-378.

13. WINTERING SITE FIDELITY IN WESTERN MARSH-HARRIER CIRCUS

AERUGINOSUS (LINN.), IN KEOLADEO NATIONAL PARK, BHARATPUR, RAJASTHAN

The Western marsh-harrier Circus

aeruginosus is a common winter visitor to the

Keoladeo National Park, Bharatpur, Rajasthan

(27° 7.6' - 27° 12.2' N and 77° 29.5' - 77° 33.2' E).

The Bombay Natural History Society has been

carrying out intensive studies on the wintering

ecology of the species in the Park since 1996.

Six Western marsh-harriers were studied

during the winter of 1999-2000. Four harriers

were fitted with radio-transmitters, and two were

ringed. One adult female was ringed with a black

band on the right leg with ‘C’ etched on it, and

two rings - one each of plastic (orange coloured)

and aluminium on the left leg. The rings could

be clearly seen with binoculars and telescope

from a distance of 100-200 m. The bird was

caught on January 20, 2000, in the wetland of

Block ‘K’ of the Park by the Stick and Glue

Method, on fish bait. It left for its breeding

ground by the end of March 2000. The bird was

seen every winter since 1996 in this area and

was identified by its peculiar plumage and eye

colour, before ringing. It was recorded again in

winter in the same area on October 14, 2000.

Site fidelity for breeding grounds has

already been recorded in Western marsh-harriers

(Witkowski 1989). However, 1 have not come

across any reference on Western marsh-harriers

returning to the wintering ground, year after year.

Wintering site fidelity has been recorded

in birds. Fischer (1981) found thrashers

Toxostoma sp. in the same wintering territories

for the consecutive year, and Price (1981)

recorded greenish leaf-warblers PhyUoscopus

trochiloides returning to the same wintering

areas in southern India. Among raptors, Steppe

buzzards Buteo buteo vulpinus in South Africa,

rough legged buzzards Buteo lagopus in South

Sweden, and common buzzards Buteo buteo in

Europe have been found returning to the same

wintering areas in successive years (Olsson 1 958,

Newton 1979). One Buteo buteo was seen in the

same place for eight consecutive winters, and

another distinctive bird for twelve winters (de

Bont 1952, Schuster 1940).

Birds have been recorded returning to the

same territories year after year, both at breeding

and wintering grounds, because they are likely

to be more successful as they have to spend less

time in getting familiarised with the habitat,

predators and to some extent food sources. Site

fidelity in birds also reflects on the quality of

habitat in terms of food and habitat availability.

Acknowledgements

I am grateful to the Rajasthan Forest

Department for permission to work in the

JOURNAL , BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

117

MISCELLANEOUS NOTES

Keoladeo National Park, especially Ms. Shruti

Sharma, Director of the Park, for the facilities

extended. I thank the US Fish & Wildlife Service

for funding the project, especially Mr. David

Ferguson for help and encouragement. Thanks

are due to Mr. Peter Bloom for help in trapping

the handers, Mr. J.C. Daniel, Hon. Secretary and

Dr. A.R. Rahmani Director, BNHS for

encouragement and critical assessment of the

manuscript.

Refe

February 6, 200 1 ASHOK VERMA

Bombay Natural History Society

Hornbill House, S.B. Singh Road,

Mumbai 400 023, Maharashtra, India.

Email: [email protected]

Present Address: Sarafa Bazar,

Rekha Naanga Gali,

Bharatpur 321 001,

Rajasthan, India.

E N C E S

De Bont, A. (1952): Breves communications. Le Gerfaut

42: 255.

Fischer, D.H. (1981): Wintering ecology of thrashers in

Southern Texas. Condor 83: 340-346.

Newton, 1. (1979): Population ecology of Raptors. T. and

A. Poyser Ltd. England.

Olsson, O. (1958): Dispersal, migration, longevity and

death causes of Strix aluco , Buteo buteo , Ardea

cinerea and Lams argentatus. Acla Vertebratica

/: 91-189.

Price, T. (1981): The ecology of the greenish warbler,

Phylloscopus trochiloides . in its winter quarters.

Ibis 123: 131-144.

Schuster, L. (1940): Langjahrige Weederkehr eines

Mausebussards ( Buteo buteo) an denselben

Uberwinterungsplatz. Vogelzug //: 86.

W itkowski, J. (1989): Breeding biology, and ecology

of the marsh harrier. Circus aeruginosus , in

the Barcycz valley, Poland. Acta Orn. 25: 223-

320.

14. SOUTHERNMOST RECORD OF EASTERN CALANDRA-LARK

MELANOCORYPHA BIMACULATA AND SIGHTING OF LESSER KESTREL

FALCO NAUMANNI FROM MATHERAN, A HILL STATION NEAR MUMBAI

Matheran is situated at c. 73° 18' E and

18° 28' N in the Western Ghats and is about

105 km from Mumbai. This tiny hill station has

an area of 7.35 sq. km, out of which 3.87 sq. km

is reserved forest while 3.48 sq. km is under

buildings, plots, tanks, roads etc. It has a

maximum elevation of 803 m and an average

rainfall of about 7,500 mm. I was in Matheran

during the second week of April 2000. During

the two day visit, I had the good fortune of sight-

ing the eastern calandra-lark Melanocorypha

bimaculata , of which this was the southernmost

record and the lesser kestrel Falco naumanni , a

Red Data species. The following is a brief account

of the same.

Eastern Calandra-lark

Melanocorypha bimaculata

On April 8, 2000, while returning from

Charlotte Lake, a man-made reservoir in

Matheran, I spotted a solitary, relatively large

member of Alaudidae. It moved hurriedly in the

dry leaf litter, along the bank of the lake, stopping

intermittently to peck at some edible morsel. The

two very distinct features that caught my attention

were its broad white supercilium and a black

patch on the side of the breast. The tail was short,

with a stroke of white on its tip. The bird was

well camouflaged against the leafy background.

It was noticeably stockier and more upright than

the greater short-toed lark Calandrella

brachydactyla. All the field characteristics

pointed at only one candidate - the eastern

calandra-lark. Later, on April 9, two more birds

were sighted near One-Tree-Hill, the

southernmost tip of Matheran. They were

foraging in a rocky slope interspersed with dry

grass beds. They remained in sight for over

118

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY. 100(1). APR. 2003

MISCELLANEOUS NOTES

15 min before disappearing behind a ledge.

The handbook states that M bimaculata is

a winter visitor to Kashmir, Punjab, Haryana,

Rajasthan, Uttar Pradesh, and an occasional visitor

to Kutch. Himmatsinhji (1960) and Tiwari (1993)

have also reported the bird from Kutch. According

to the handbook, there is no record of the bird from

Maharashtra. However, Grimmett et al. (1999)

mention its presence up to NW Maharashtra. There

is also a confirmed unpublished record of two birds

near a waterbody in Nasik dist. (Raha, B. pers.

comm, in 1999). The present record is therefore

the southernmost one for M. bimaculata.

Lesser Kestrel Falco naumanni

On the afternoon of April 9, 2000, I saw a

bird of prey that had just landed on a barren tree

on the rocky slope; 50 m separated the two of us.

It was difficult to decipher its identity from that

distance, but from the size, it looked like a falcon.

On a keener view, it resembled the female of the

common kestrel Falco tinnunculus.

After preening for a few minutes, the bird

began hovering in a typical kestrel-like fashion

with short bouts of rapid wing beats interrupted

by equally short flights. 1 needed a closer look to

see other characteristics to confirm my initial

identification. The bird glided in my direction and

was below me, providing a dorsal view. The flight

was definitely less laboured than that of the

common kestrel. Just when 1 was weighing up its

true identity, the bird got hold of a strong wind

current that carried it at least 20 m above me in a

fraction of a second. The underbelly was streaked

while the wings had darker wing margins.

Meanwhile, the bird spotted something in

the grass and swooped down on it. At this

juncture it went out of sight while I waited to get

another glimpse of it. Within moments it returned

with something in its bill and settled on a

Ficus sp. jutting from the vertical rock face to

my right, but at a lower level than me. While the

bird dismembered its booty, I crept up to almost

15 m from it. The head was pale and the

conspicuous moustachial stripe of the common

kestrel was not visible. The bird was holding on

to a grasshopper with its claws. The claws lacked

the dark tinge of the common kestrel and by then

1 was convinced that it was a female of the lesser

kestrel Falco naumanni.

According to Ali and Ripley (1987), Falco

naumanni is more patchily distributed than its

larger cousin, F. tinnunculus and is an irregular

passage migrant to East Africa. A few stragglers

are known to remain behind and have been

sighted between November and April in Ambala,

Delhi, Lucknow (U.P.), Dinapur (Bihar),

Balasore (Orissa), Kolkata (Bengal), Dibrugarh,

Naga hills and N.E. Cachar (Assam), Manipur,

to as far south as Chennai (Coonoor-Nilgiris).

Within Maharashtra, a flock of several hundred

birds has been sighted in Sholapur, apart from a

flock of a dozen birds in Ahmednagar in January.

Apart from this, there have been unpublished

records of the bird from Nasik (Raha, B. pers.

comm., 1999). F. naumanni is also recorded from

Sri Lanka (Hoffmann, 1996).

Though rare, it is widely distributed in India.

The inadequate data on the bird could probably be

attributed to its resemblance to F. tinnunculus. The

females of the two species are difficult to

differentiate in the field. Only a trained eye can

distinguish between the two birds from their

hovering styles — instead of the up and down wing

strokes of the common, the lesser kestrel beats its

wings forward and backward. This is apparently

because the body of the lesser kestrel is raised 45°

while that of the common is held parallel to the

ground. The most important identification is the

paler, almost colourless claws of F. naumanni

compared to the black ones of F. tinnunculus. This

is the first record of this red data book species from

Matheran and therefore worth mentioning.

November 25, 2000 ANISH ANDHER1A

2, Sagar Building, V.R Road, Andheri (W),

Mumbai 400 058, Maharashtra, India.

Email: [email protected]

JOURNAL , BOMBAY NATURAL HISTORY SOCIETY. 100(1), APR. 2003

1 19

MISCELLANEOUS NOTES

References

Ali, S. & S.D. Ripley (1987): Compact Handbook of the

Birds of India and Pakistan together with those of

Bangladesh, Nepal, Bhutan and Sri Lanka. 2nd edn,

Oxford University Press, Delhi, pp.737 + 104 plates.

Grimmett, R., C. Inskipp & T. Inskipp ( 1 999): Birds of the

Indian Subcontinent. Oxford University Press, New

Delhi.

Himmatsinhji, M.K. (1960): The Eastern Calandra Lark

( Melanocorypha bimaculata ) in Kutch. J. Bombay

nat. Hist. Soc. 57: 408.

Hoffmann, T.W. (1996): New bird records in Sri Lanka

and some connected matters. J. Bombay nat. Hist.

Soc. 93(3): 382-388.

Tiwari, J.K. (1993): Sighting of Eastern Calandra Lark,

Melanocorypha bimaculata (Blyth) in Kutch.

In: Proc: Changing Scenario of Bird Ecology and

Conservation, Ornithological Society of India,

Bangalore.

15. UNUSUAL FEEDING ASSOCIATION BETWEEN SIBERIAN CRANE

GRUS LEUCOGERANUS AND WILD BOAR SUS SCROFA

IN KEOLADEO NATIONAL PARK, BHARATPUR, RAJASTHAN

Siberian cranes Grus leucogeranus are

wetland-dependant birds and unlike other

cranes that often forage in dry upland areas

near wetlands, they usually forage in ankle

deep shallows (Sauey 1985). Three Siberian

cranes were observed feeding in dried up areas

of wetland in the Keoladeo National Park,

Bharatpur, India. The cranes were in areas

where the soil appeared ploughed and were

observed picking up tufts of grass and putting

them aside, and then picking up small items

of food. After the cranes had moved away from

the foraging sites, we noticed that the area

had wet soil and was dug up by boars Sus

scrofa , as was evident from the hoof marks

and droppings of wild boars. All the grass was

uprooted; tufts of roots and partly eaten insect

larvae were lying all over. A bunch of larvae

were wriggling under the tufts of grasses,

which were lying all over the dug up areas.

Wild boars are known to feed on roots

of grasses and sedges, and also on insects

(Prater 1971). They had probably uprooted the

grasses to get to these larvae. The cranes took

advantage of the ploughing by the wild boars,

as they themselves would have found it

difficult to uproot the grasses in the dried up

area where the soil had hardened after drying.

The insect larvae were very small and

individually not of much food value to cranes,

but since they were available in abundance,

the cranes may have found it profitable to feed

on the larvae, which are not on their regular

diet. Only the captive bred released birds,

which were a part of an experiment to augment

the wild population of Siberian cranes, were

observed feeding on larvae. It was probably

behaviour learnt from sarus cranes, as the

Siberian cranes were associated with them

more than with the wild Siberian cranes. The

captive released birds would often venture out

with the sarus cranes to forage in the dry areas.

The Siberian cranes are known to feed largely

on vegetable matter and occasionally on

aquatic insects in their wintering grounds (Ali

and Ripley 1983, Vijayan 1991).

Birds are known to benefit from feeding

associations with other animals. Egrets

Egretta spp., drongos Dicrurus spp., starlings

Sturnus spp., mynas Acridotheres spp., and

many other species follow grazing animals and

pick up insects disturbed by the movement of

these animals in the grass (Ali and Ripley

1983). However, we have not come across any

reports of Siberian cranes, which are largely

wetlands birds, getting direct benefit from the

action of a terrestrial animal like the wild

boar.

120

JOURNAL BOMBAY NATURAL HISTORY SOCIETY 100(1). APR. 2003

MISCELLANEOUS NOTES

Acknowledgements

We thank the Rajasthan Forest Department

for permission to work in the Park, U.S. Fish

and Wildlife Service for funding the project and

the Bombay Natural History Society, especially

Dr. A. R. Rahmani, Director, for encouragement

and valuable comments.

Refer

Ali, S. & S.D. Ripley (1983): Compact Handbook of the

Birds of India and Pakistan together with those of

Bangladesh, Nepal, Bhutan and Sri Lanka. Oxford

University Press, Delhi. Pp. 737 + 104 plates.

Prater, S.H. (1971 ): The Book of Indian Animals. Bombay

Natural History Society, Bombay. Pp. 324.

November 7, 2001 GARGI RANA

♦VIBHU PRAKASH

Bombay Natural History Society

*Present Address: BNHS Field Station,

F-23, HMT Colony, Pinjore 134 101,

District Panchkula, Haryana, India.

Email: [email protected]

N C E S

Sauey, R.T. (1985): The Range. Status and Wintering

Ecology of Siberian Cranes Gms leucogeramis . Ph.D.

Thesis, Cornell University, Ithaca, NY. Pp. 411.

Vu ayan. V.S. (1991); Keoladeo National Park Ecology

Study. Final Report 1980-90. Bombay Natural

History Society, Bombay. Pp 337.

16. COMMON COOT FULICA ATRA FROM KYONGNOSLA IN EAST SIKKIM

In early March 2000, there were reports of

migrating water birds in groups of 50+ from the

Kyongnosla area of east Sikkim (c. 3000 m). On

March 15, 2000 one bird that had fallen from

the sky from its group of c. 1 00 birds was caught

and taken to a nearby settlement. Mr. Bishnu

Sharma, Supervisor, Kyongnosla Alpine

Sanctuary, immediately retrieved it and brought

it to my office at Deoral i, Gangtok the next day

in a cardboard carton. The bird was an adult

common coot Fulica atra. It was alert and active,

stabbing at my hand with its bill, and it drank

water copiously. It seemed unhurt and had no

external injuries. As I had no apparatus to ring

or measure it, or take photographs, I took it to

my residence at the Forest Colony, Baluakhani,

Gangtok (1,800 m) that evening, where it

escaped. For over an hour it wandered in the

garden before fluttering down to the road and

going into the forest scrub further down. I

R E F E

Ali, S. (1962): The Birds of Sikkim. Oxford University

Press. Delhi.

An, S. & S.D. Ripley (1983): Handbook of the Birds of

watched for it till dark and could not find it the

next morning.

The common coot, which has a wide

distribution range over the Indian subcontinent,

has not been recorded from Sikkim so far, though

the area is a well known traditional flyway for

water birds on return migration northwards in

spring (Ali and Ripley 1 983; Ali 1 962) probably

due to paucity of field surveys. This could

therefore be considered a new record for Sikkim.

March 22, 2001 USHA GANGULI-LACHUNGPA

Department of Forests,

Environment & Wildlife,

Government of Sikkim, Deorali,

Gangtok 737 102, Sikkim, India.

Email: [email protected]

BISHNU LAL SHARMA

Kyongnosla Alpine Sanctuary

E N C E S

India and Pakistan together with those of

Bangladesh, Nepal. Bhutan and Sri Lanka. Oxford

University Press, Delhi.

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17. SIGHTING OF THE LESSER FLORICAN SYPHEOTIDES INDICA

IN GIR INTERPRETATION ZONE, DEVALIYA, GIR SANCTUARY

On June 27, 2000, I had gone to the Gir

Interpretation Zone (GIZ), Devaliya, specially

developed for tourism in the Gir Sanctuary. I had

gone to check the health of the lions. The GIZ

has a fenced area of 412 ha, with c. 100 ha of

Savannah grassland. During the visit,

Sardulbhai, one of my staff, informed me that

an unknown bird similar to a small peahen had

been spotted near the lions’ feeding cage. On

reaching the spot, I saw a domestic hen-sized

bird with a crest on its head, at some distance.

When we approached, the bird ran into a patch

of tall grass and hid there in silence. To my great

jubilation, I confirmed it as a male lesser florican

(. Sypheotides indica) through my binoculars and

immediately took some photographs. It is for the

first time that a lesser florican has been

photographed in the Gir Protected Area. The

same species was reported from the Gir Protected

Area (and the sighting confirmed) during May

and June, 1982. According to Sankaran et al.

(1992) the lesser florican sometimes visits Babara

Vidi grassland, Maliya taluka, Junagadh district,

an ideal habitat for lesser florican. Our field staff

and researchers have observed florican in this

area several times.

From June 27 to July 10, I visited GIZ,

Devaliya regularly, twice a day, morning and

evening. The florican was mostly seen in the

grasslands. I saw it display twice. To ascertain

the presence of other lesser florican (especially

female) in GIZ, I used a group of trackers, but

sighted only the same male. After July 10, 2000,

the bird left the GIZ, Devaliya.

November 22, 2002 B.P. PATI

Wildlife Division, Sasan Gir 362 135

Junagadh district, Gujarat, India.

Reference

Sankaran, R., A.R. Rahmani & U. Ganguli-Lachungpa (1992): The distribution and status of the lesser florican

Sypheotides indica (J.F. Miller) in the Indian subcontinent. / Bombay nat. Hist. Soc. 89: 156-179.

18. GREY-HEADED LAPWINGS VANELLUS CINEREUS SEEN AROUND

MACHILIPATNAM, KRISHNA DISTRICT, ANDHRA PRADESH, INDIA

On February 17, 2001, at around 0645 hrs,

I was watching waders of several species, along

with other birds, on a largely flooded field,

1-2 km northwards of Chilakalapudi,

Machilipatnam. Amongst the waders, which

included several wood sandpipers, little ringed

plovers, Pacific golden-plovers and a few little

stints, were twelve larger birds. An initial look

was enough to place them with lapwings

( Vanellus ) and the birds looked superficially

similar to pictures of white-tailed lapwing, which

I had seen.

Their posture was upright, with a small to

medium sized, smooth round head with no crest.

Size was similar to red-wattled lapwing ( Vanellus

indicus) of which there were three in a nearby

field. The head and neck were a uniform pale

grey colour with no supercilium or eye-stripe.

The bill was bright yellow and no wattles could

be seen on the face. A wide ( 1 -2 cm) black breast

band separated the grey of the neck from the belly,

which was white. Legs were bright yellow, the

same colour as the bill. The wings, which were

seen when one of the birds flew a short distance,

showed a very distinctive ‘tri-coloured’ pattern,

with black primaries, white secondaries and

brownish coverts, in three, straight edged blocks

of colour.

The birds were watched for 30-45 minutes

through a Bushnell Spacemaster Telescope with

122

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MISCELLANEOUS NOTES

a 22x wide-angle lens and 8x32 Leica binoculars

at a distance of perhaps 50-100 m (some of the

birds were further away). Light conditions were

good, with the early morning sun coming from

just behind my right shoulder.

After consulting birds of the Indian

subcontinent by Grimmett et al. (1998), I was

confident enough to identify the birds as grey-

headed lapwings ( Vanellus cinereus ), which

according to that book had not been recorded in

Andhra Pradesh before.

On Friday February 23, at a different site,

just east of Machilipatnam, 5 more grey-headed

lapwings were seen, in groups of 2 and 3 in paddy

fields on either side of a small road going down

to the sea.

With the exception of the wing pattern (the

birds did not fly) all the same features were

observed, and in addition the black tip to the

yellow bill was noted.

Six grey-headed lapwings were seen again

just north of Chilakalapudi, Machilipatnam on

March 13, 2001 and one on April 3, 2001.

March 24, 2001 COLIN CONROY

13 Avenue Road, Southall,

Middlesex, UB1 3BL, U.IC.

Email: [email protected]

19. THE GREY-HEADED LAPWING IN TAMIL NADU - A REJOINDER

In his note on the grey-headed lapwing

Vanellus cinereus (Blyth), [JBNHS 97(2): 277],

Gopi Sundar writes that in peninsular India, this

species has only been recorded in Bangalore in

1987 prior to his sighting in Kaliveli Tank near

Pondicherry in 1997. He further writes,

“Perennou and Santharam have conducted

detailed ornithological surveys in this region and

have not come across this species.”

I would like to point out that this is not

true. Gopi Sundar has only referred to two of

our publications and hence overlooked my note

published in Blackbuck 3: 25-27 (1987). In this

note, I had recorded the sighting of the grey-

headed lapwing in Madras city (now Chennai)

on January 1 1, 1987. The bird was spotted on a

mud flat of the Adyar Estuary from the

Theosophical Society estates. T.R. Shankar

Raman had spotted it and all the nine bird

watchers present on that occasion had a good

look at the bird, which was a juvenile. The

lapwing was not seen on subsequent visits. This

record also finds a mention in the birds of the

Indian subcontinent (Grimmett et. al ., 1998

Oxford).

May 4, 200 1 V. SANTHARAM

Institute of Bird Studies and

Natural History,

Rishi Valley School, Rishi Valley P.O.

Pin 517 352, Andhra Pradesh,

India.

20. A SIGHT RECORD OF BUFF-BREASTED SANDPIPER

TRYNGITES SUBRUFICOLLIS IN GOA

At about 1 225 hrs on November 1 8, 2000,

while birdwatching with a Sunbird tour group

in recently irrigated paddy fields at Santa Cruz,

Tiswadi taluka, Goa, Rick Heil found a buff-

breasted sandpiper Tryngites subruficollis. The

bird performed superbly and was watched, by RH

and five other observers including Paul 1. Holt,

David Hemmingway and Mark Newsome,

uninterrupted for about 50 min and at ranges

down to c. 75 m. We used a variety of binoculars

and telescopes, some of the latter with eyepieces

of up to 60x magnification. RH and PIH both

had extensive experience of the species and all

observers were completely confident of RH’s

initial identification. RH, PIH and MN took field

notes and PIH did a hurried sketch (copies of

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

123

MISCELLANEOUS NOTES

these are held by PIH). Unfortunately, the bird,

a juvenile showing no signs of moult to first-

winter plumage, could not be located when other

observers searched for it later the same afternoon.

Throughout the observation we were aware

that, although the buff-breasted sandpiper is a

very distinctive shorebird, it has, on occasion,

been confused with the ruff Philomachas pugnax

(several of which were present nearby) and with

the extralimital Baird’s sandpiper Calidris

bairdii.

Description: The following description is

a synthesis of notes taken by PIH, RH and MTN.

Structure: A medium sized wader,

fractionally smaller than the accompanying wood

sandpipers Tringa glareola and proportionately

slightly larger-headed and with a shorter and

thicker neck than that species. The bird’s head

nevertheless appeared relatively small and its

size, in combination with its plumage pattern

and shape (in particular its steep forehead, flat

crown and angular nape), created a soft, open-

faced and almost dove-like impression. The

bird’s body also appeared slimmer, sleeker and

more attenuated at the rear than those of the wood

sandpipers.

The folded wings extended beyond the tip

of the tail by a distance approximately equal to

half the length of the bill, while two, and just

possibly three, primary tips were visible beyond

the longest, lanceolate and remarkably wispy

tertial.

The bill, proportionately shorter than that

of the wood sandpiper, was approximately equai

to the length of the head. It was very straight,

the lower mandible having an almost

imperceptible arch along its lower edge, and

tapered to a rather fine tip. The legs were

similarly proportioned to those of the wood

sandpiper.

Plumage: The whole face, front and sides

of the neck and virtually the entire underparts

were a remarkably uniform fawn-, or cinnamon-

buff colour while the feathers of the upperparts

were dark centred and neatly scalloped.

Head: A rather large, dark eye was very

conspicuous on an otherwise plain, ‘open’ face.

The ear-coverts were slightly duskier, setting off

paler buff supercilia and there was a pale buff,

broken eye-ring, more noticeable above the eye.

The forehead was unmarked, though the crown,

nape and hind-neck to mantle were marked with

fine blackish streaks.

The upperparts were attractively patterned.

Dark, almost blackish centres to the mantle,

scapulars, wing coverts and tertials contrasted

crisply with their broad buff or whitish-buff

fringes. Compared to the rest of the upperparts,

the centres to the wing coverts were paler, the

fringes broader and more diffusely demarcated.

The resulting pale wing panel contrasted quite

well with the lower scapulars. The wing coverts

also exhibited the anchor-shaped internal

markings that are typical of juvenile buff-breasted

sandpipers (A narrow blackish shaft being

connected to a broader blackish sub-terminal

crescent that traced the feather’s shape). A

similar pattern was also discernible in some of

the lower scapulars. There were no signs of any

moult, all of the upper-part feather tracts

appeared rather fresh and were arranged in neat,

linear rows. Individual feathers, particularly the

larger wing coverts and rear lower scapulars,

were easily identified and counted.

Wings: Although the bird was seen only

very briefly in flight, it opened its wings, to

stretch or to preen on a couple of occasions, and

we were then able to discern some details of the

wing pattern. The flight feathers were dark and

contrasted well with the paler inner wing-coverts.

A narrow, very indistinct, or even obscure (RH),

pale-buff wing-bar was just apparent across the

tips of the greater coverts. The under-wing

coverts appeared clean white, contrasting with

the underside of the flight feathers and especially

with a relatively conspicuous blackish comma,

or crescent, formed by the primary under-wing

coverts.

124

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MISCELLANEOUS NOTES

Underparts: The chin was fractionally, but

perceptibly, paler than both the face and the fore

neck, and while the fawn-buff neck, breast and

belly were concolorous the remainder of the

underparts gradually faded paler and whiter from

the rear belly through the vent to the undertail

coverts. Isolated patches of bold, blackish flecking

or spotting extended to either side of the breast.

Bare parts: The bill appeared dark and

there was possibly a very small, slightly paler

area at the extreme base of the lower mandible.

The legs were a uniform olive-yellow, the colour

of English mustard, and had a slightly more

orange hue than those of all the accompanying

wood sandpipers.

Behaviour: During our observation the

bird fed busily. It was almost constantly on the

move, walking around the paddies with a

distinctive head-bobbing, almost dove-like

motion and high stepping, tripping gait. No

interaction was noted between it and any of its

equally busily feeding companions. As is typical

of the species, the bird remained silent.

Status: Buff-breasted sandpiper is almost

exclusively Nearctic in its distribution. It breeds

across the Arctic belt of North America from

Alaska to western Canada, as well as on Wrangel

Island and perhaps the Chukotskiy peninsula,

Siberia (Cramp and Simmons 1983). It winters

in northeast South America. Brazil (1991) listed

12 records from Japan and, although the species

has not been recorded anywhere in southeast Asia

(Robson 2000), there is at least one report from

the east coast of Taiwan (MacKinnon and

Phillipps 2000), and at least eight sightings in

Australia (Pringle 1 987). There are four previous

records from the Indian subcontinent. Three of

Refer

Brazil, M. (1991 ): The Birds of Japan. Christopher Helm,

London.

Cramp, S. & K.E.L. Simmons (1983): Birds of the Western

Palaearctic. Vol. 3. Oxford University Press. London.

Hoffmann, T. (1991): Notes on accepted sight records of

birds in Sri Lanka. J. Bombay nat. Hist. Soc. 88(2):

these reports are from the east coast of Sri Lanka

- a specimen collected by T.S.U. De Zylva at

Kalametiya near Hambantota on the March 5,

1 960, and two subsequent sight records, one near

Trincomalee in November 1974 and the other at

the Bundala Sanctuary on the January 19-23,

1985 ( CBCN 1960: 14, CBCN 1974: 41, CBCN

1985: 2, Hoffman 1991 and Lam fuss 1998). The

only previous record from India was of a bird seen

by Per Undeland at Harike Bird Sanctuary, Punjab

on the May 18,1 995 (Robson 1 996). Perhaps even

more surprisingly, there are three recently accepted

records ( 1 2. ii i. 198 1 , 31.x. - 14. xi. 1997 and

1 3- 1 9.xi. 1 999) from the Seychelles (David Fisher

pers. comm, and Adrian Skerrett in lift, to P1LI).

Acknowledgements

Krys Kazmierczak kindly provided

information on the previous subcontinent

records, supplied several references, contact

addresses and reviewed a draft of this note.

Guenter Lamfuss responded speedily to our

requests for further information on the Sri

Lankan records and both David Fisher and

Adrian Skerrett provided information concerning

the species’ status in the Seychelles.

March 24, 2001 PAUL I. HOLT

New Laund Farm, Greenhead Lane,

Fence, Burnley, Lancashire,

BB12 9DU, UK.

Email: [email protected]

RICK HEIL

20 MacArthur CIR,

Peabody, MA 01960, USA.

E N C E S

381-383

Lamfuss, G. (1998): Die Vogel Sri Lankas: ein Vogel- und

NaturfLihrer. [The Birds of Sri Lanka: a Bird and

Nature Guide (In German).] Heidelberg: Kasparek

Verlag.

MacKinnon, J. & K. Phillipps (2000): A field guide to the

JOURNAL. BOMBAY NATURAL HISTORY SOCIETY. 100(1). APR. 2003

125

MISCELLANEOUS NOTES

birds of China. Oxford University Press, London.

Pringle, J.D. (1987): The Shorebirds of Australia: the

National Photographic Index of Australian Wildlife.

Angus & Robertson, North Hyde.

Robson, C. (1996): From the field: India. Oriental Bird

Club Bull. 23: 50.

Robson, C. (2000): A Field Guide to the Birds of South-

east Asia. New Holland, London.

21. GROUP FISHING OF HOUSE CROWS (CORVUS SPLENDENS)

WITH RIVER TERNS ( STERNA AURANTIA)

( With one plate)

On July 6, 2000, during a visit to Bigwan,

Indapur taluka, Pune district, Maharashtra State

for bird watching, I noticed a flock of about

40 river terns ( Sterna aurantia) and 10 house

crows ( Corvus splendens ), flying together at one

spot, in the backwaters of Ujani Dam. As I

approached closer, I saw fishermen emptying

their catch from the nets.

The house crows and river terns were

diving for the dead fish, which had fallen from

the fishing nets, and were floating on the water.

The crows had mastered the technique of

hovering and accurately picking up the fish, just

like the river tern.

The house crow is not a water bird; whether

this was a natural instinct or it had mastered the

technique while observing the river terns is a

big question. Crows are known to be territorial

and aggressive, but in this case they neither

harmed nor quarreled with the terns.

Another observation worth noting was that

while the river tern picked up and swallowed

the fish in flight the crow would pick up a fish,

fly to the shore to eat it, and then fly back to

catch another one.

April 12, 2001 SATTYASHEEL N. NAIK

781/782, Shukrawar Peth,

‘Laxmi Chhaya’, Opp. Jain Mandir,

Pune 411 002, Maharashtra, India.

22. TERNS OF THE VENGURLA ROCKS, A REVIEW AND UPDATE

The breeding colony of maritime terns on

the Vengurla Rocks is arguably the Indian

subcontinent’s most important one, if not for the

sheer quantity of nesting birds then for the

number of constituent species. Though it has been

known for at least 125 years that a ternery exists

on Burnt Island, one of the islets forming the

Vengurla Rocks, all knowledge about it rested

on local lore, second-hand accounts and

circumstantial proof. No ornithologist had ever

seen terns there before 1981.

The Vengurla Rocks are a tiny archipelago

of rock outcrops situated 16.5 km off the South

Konkan coast from a point about halfway between

Vengurla and Malwan (Sindhudurg district,

Maharashtra State, India), situated at

15° 43' 24" N and 73° 27 42" E. The four largest

ones form a group of precipitous rocky islets at a

distance of several hundred metres from each other.

Among them, the smallest and westernmost is

crowned by the ruins of the old lighthouse. This

was replaced around 1935 by a more modern one

on the largest of the rocks, manned by a crew of

eight that resides on it throughout the year. Burnt

Island, the second largest of the islets, lies closest

to the mainland. It is about 300 m long, 100 m

wide and 30-50 m high, and sparsely covered by

coarse grass and a few stunted bushes. The islet

remains free from human interference and A.O.

Hume’s (1876) detailed description is as accurate

and valid today as it was 125 years ago.

The group of islets as a whole presents a

forbidding aspect, access is limited to the fair

season, October to May, and landing is risky even

at the best of times. In January 1998, permission

given to me to stay at the Inspection Quarters of

126

JOURNAL BOMBAY NATURAL HISTORY SOCIETY. 100(1). APR. 2003

MISCELLANEOUS NOTES

Naik, Sattyasheel N.: Corvus splendens & Sterna aurantia Plate 1

Fig. la-b: House crows fishing with river terns

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127

f

MISCELLANEOUS NOTES

the lighthouse island was revoked by the

Directorate of Lighthouses and Lightships, after

a crewman was killed when trying to alight from

the departmental boat on to the rock. During the

southwest monsoon, the time when the terns breed,

the Vengurla Rocks are totally inaccessible, and

the lighthouse crew is cut off from the world

between May 31, when a country-craft under

contract brings the last supplies, and early

September, when communications are resumed.

Antecedents

it was the breeding colony of edible-nest

swiftlets on Burnt Island that caught the interest

of ornithologists in the second half of the 19th

century. The taxonomic status of these enigmatic

swiftlets was heatedly discussed then, and the

ornithologists’ demand for bird specimens must

have been as great and urgent as the Chinese

gourmet’s for their soup-providing nests. All we

came to know about the equally enigmatic temery

was an incidental spin-off of these studies.

A.O. Hume (1 876), in February 1 875, visited

three of the islets and spent several hours on Burnt

Island on his way to the Laccadives (now

Lakshadweep). He did not see any terns, but

collected desiccated remains of terns and eggs.

G.W. Vidal (1880), solely interested in swiftlet

specimens, sent his ‘shikaree’ on collecting trips

to the Vengurla Rocks in April 1878, December

1879 and February and April 1880. Some years

later, J. Davidson apparently got a number of tern

eggs collected by the crew of the old lighthouse

{vide Whistler in Abdulali 1939).

The issue of the specific status of the Indian

edible-nest swiftlet Collocalia unicolor (Jerdon)

having been settled by then, nobody seems to

have taken interest in the Vengurla Rocks until

Humayun Abdulali took up the challenge over

40 years later. He turned out to be the most

enterprising of all the visitors, managing to land,

and even stay overnight, on Burnt Island in

February 1938 and 1941 (Abdulali 1939, 1941).

Forty years later, in May 1981, he undertook

another collecting trip to Burnt Island, this time

of only a couple of hours duration. It yielded,

besides various desiccated remains of adult birds,

chicks and eggs, the first sight-record of live

terns: a group of some 50 birds, identified as

Sterna anaethetus and a few S. fuscata, circling

half a mile from the Rocks (Abdulali 1983).

A critical appraisal of the old breeding records

By 1982, the following six tern species

were established and listed in the handbook and

synopsis - as breeding on the Vengurla Rocks.

One of these, the sooty tern Sterna fuscata ,

should be omitted.

Roseate tern Sterna c/ougallii Montagu

(Syn. No. 466): A number of eggs collected on

Burnt Island by the lighthouse staff sometime

during the last decade of the 19th century were

ascribed by J. Davidson to this species ‘without

keeping notes on how he arrived at this

identification’ (Whistler in Abdulali 1939). This

probably had prompted E.H. Aitken to state in

his common birds of Bombay (c. 1900): ‘The

Roseate Terns breed in the Vingurla Rocks (sic)

during the monsoon.’

Addled eggs and desiccated bird-remains,

collected by Abdulali in 1938 and 1941, were

examined by Hugh Whistler at the British

Museum, London, and Salim Ali at the Bombay

Natural History Society and identified as

belonging to this species (Abdulali 1939, 1941).

White-cheeked tern S. repressa Hartert

(Syn. No. 467): Out of Abdulali’s 1938 and 1941

material, Ticehurst examined the mummy of a

juvenile tern, and Whistler and Salim Ali several

addled eggs that they ascribed to this species

(Abdulali 1939, 1941). Apparently, there were

no signs of this tern having bred at the time of

Abdulali’s 1981 visit to Burnt Island.

Bridled tern S. anaethetus Scopoli (Syn.

No. 471): Hume (1876) found, in February 1875,

on Burnt Island “innumerable addled, broken and

more or less decayed tern’s eggs, while all about

in similar situations, mummies, desiccated

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129

M ISC ELLA NEO US NO TES

corpses of young birds of all ages and a few old

S. anaethetus lay scattered leaving no possibility

of doubt as to the species which chiefly breeds

there.” The bird and egg remains he collected

were identified at the British Museum, London.

Sooty tern S. fuscata Linn. (Syn. No.

474) : Salim Ali, when examining the material

collected by Abdulali in February 1938, had

noted: “... Two brown wings, however, measure

270 and 294 mm, and apparently belong to the

sooty tern (S. fuscata). This is a new record for

Vengurla; the nearest hitherto known breeding

place is in the Laccadives. A beak measuring

40 mm may support this identification.” (vide

Salim Ali in Abdulali 1939).

This record of a mere ‘occurrence’ of this

tern has found its way as a ‘breeding record’ into

works like Ali and Ripley (1983), Harrison

( 1 985), Grimmett et al. ( 1 998) and Kazmierczak

(2000). 1 feel that Salim Ali had jumped here to

an inadmissible conclusion: The presence of a

pair of wings and a beak on Burnt Island hardly

constitutes proof of the bird having bred there.

Ripley (1982) did not follow Salim Ali’s

reasoning and omitted this ‘breeding record’

from his synopsis.

Little tern S. albifrons Pallas (Syn. No.

475) : Desiccated remains of mostly downy chicks

with emerging feathers, collected by Abdulali in

May 1981, were identified by Dr. Jon Fjeldsa at

the Zoological Museum of the University of

Copenhagen, Denmark, as being of this species

(Abdulali 1983).

Though a modicum of doubt must adhere

to a specific identification of small chicks without

any supporting evidence, especially if the

specimens are of the smallest species out of a

mixed breeding colony that abounds in chicks of

species only slightly larger than the bird in

question, Dr. Jon Fjeldsa remains confident of

his identification (in litt. 314.2001).

Large crested tern S. bergii Lichtenstein

(Syn. No. 478): Addled eggs and mummified

remains of a juvenile bird, collected by Abdulali

in February 1938 and 1941, were examined by

Whistier at the BMNH, London, and Ticehurst

and Salim Ali at the BNHS, Bombay, and

ascribed to this species.

Recent visits

In May 1988, Stig T. Madsen, a Danish

bird watcher interested mainly in seabirds,

undertook an exploratory cruise through the

group of islets on a fishing trawler chartered at

Vengurla (Madsen 1988). His report of having

seen c. 20 each of S’, anaethetus , S. repressa and

S. bergii and 300-500 unidentified terns

prompted me to accompany him on a three day

stay at the inspection quarters of the Lighthouse

Is., at the end of March 1989. To reach the Rocks

we hired a motorized and outrigger-fitted dugout-

canoe at Kochra, a fishing village at the mouth

of a narrow creek between Vengurla and Malwan,

at the closest distance to the Vengurla Rocks.

By the same mode of transport, 1 visited

the Lighthouse Is., again for three days each time,

in mid-May and mid-October 1989. A

Zeiss 30x60 refractor was used to observe tern

activity on close-by Burnt Is. and in the entire

‘archipelago’. A further trip, rounding Burnt

Island for a couple of hours and spending the

night on the canoe in its vicinity, was made in

mid-September. In mid-December 1989, 1 landed

on Burnt Island, but suffering from a bout of

vertigo, was not able to collect more than a couple

of tern mummies.

These visits spread more or less randomly

throughout the fair season, served to determine or,

at least, to narrow down the time when the terns

appeared on and left from their breeding ground.

After a hiatus of nearly 8 years, a new series

of visits was undertaken in an attempt to get a

glimpse of the breeding terns themselves.

Using a hired, diesel-powered, open

fibreglass boat out of Kochra, 1 circled Burnt Is.

on June 5 and 9, and September 30, 1997,

accompanied by Gordon Frost and Nick Manville

on two of these outings. After an uneventful visit

130

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MISCELLANEOUS NOTES

on April 30, 1998, an unexpected lull in the

southwest monsoon allowed me to see the

breeding colony at its apparent peak on August

16, 1998. During this crucial visit, an immense

‘cloud’ of well over 15,000 terns could be

observed on the precipitous leeward (towards the

mainland) side of Burnt Island. In addition to

that, a conservatively estimated 1 0,000 terns were

spread over the less steep seaward side, but the

almost impenetrable spray created by thunderous

breakers and the heaving and tossing of the small

boat made closer observations and a species-wise

break-up impossible.

These 12 visits, spread over the years 1988-

89 and 1 997-98, covered all seasons and all months

except January, February, July and November.

Breeding terns

Roseate tern Sterna dougallii Montagu (Syn.

No. 466):

March 25, 1989 : Flocks of up to c. 700

smallish white terns, too distant for identification,

visited the vicinity of the Vengurla Rocks.

June 5, 1997 : The first positive sighting

of roseate terns on Burnt Island, just two days

before the regular and 1 0 days before the actual

onset of the southwest monsoon. Well over 800

birds in full breeding plumage were sitting openly

on rock-ledges, not more than 10 m above the

waterline or flying close to the islet. Though

almost all these terns seemed to be paired off,

there was no visible breeding activity as such.

August 16, 1998 : A conservatively

estimated number of 10,000 adult birds milled

around the islet in a veritable feeding frenzy.

They kept coming in, mainly from the seaward

side, with small fish in their bills, diving into

rock fissures and crannies and wider tussocks of

the lush coarse grass, all located on the lower

half of the islet obviously feeding the downy

chicks and youngsters hidden there, and taking

off immediately to haul in more. Very few fully-

fledged juveniles were perched openly on rock-

ledges, while many adults were apparently still

incubating clutches of eggs or brooding freshly

hatched chicks.

September 9, 1997 : Hectic feeding of

concealed youngsters by over 4,000 adult birds

continued. Only a dozen or so fully-fledged

juveniles were visible. The rosy tinge of the

breeding plumage had already faded in about half

of the adults.

September 30, 1997 : Only just over 80

birds, including 5 juveniles were present. Feeding

activity had ceased. The adults breeding plumage

had almost completely faded.

October 14, 1989 : All roseate terns had

left Burnt Island.

Bridled tern S. anaethetus Scopoli (Syn. No.

471):

May 10, 1981: Up to 50 birds were present

in the group of islets.

May 13, 1989: The activity of the over 400

birds present was loosely centered on Burnt

Island. Roughly one half of the birds were already

paired off, standing for long stretches of time on

rock-ledges and under overhanging rocks of the

seaward side. The other half were wheeling in

circles over the islet and chasing each other in a

sort of aerial nuptial display. At dusk, the terns

dispersed; no roosting on any of the islets was

observed. During the night, no calls were heard

from the vicinity.

June 5, 1997: More than 400 bridled terns

had occupied crags and fissures on the lower half

of the seaward side, apparently incubating.

August 16, 1998: Well over 3,000 adult

birds were present, frenziedly feeding chicks and

youngsters that were hidden from view. Very few

fully-fledged juveniles were visible.

September 9, 1997: Over 300 adult birds

were perched on the leeward side of Burnt Island;

no ju veniles were visible; all feeding activity had

stopped.

September 17, 1989: Over 200 terns were

either wheeling around the islet or surface

feeding in the vicinity. Going by their calls, they

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

131

MISCELLANEOUS NOTES

appeared to keep up their activity throughout the

night. Between the Vengurla Rocks and the

mainland, c. 60 birds per hour were continually

passing on their autumnal southward mass

migration, presumably from their breeding

grounds on the southern shores of the Red Sea

and the Persian Gulf.

September 30, 1997: All bridled terns had

left.

Large crested tern S. bergii Lichtenstein (Syn.

No. 478):

This tern occurs along the South Konkan

coast throughout the year in small groups and

loose flocks of up to 800 (Lainer 1999a). Around

the Vengurla Rocks, small numbers (up to 25

birds) were commonly seen during the dry

season.

June 5, 1997: Over 1,000 birds were

perched on rock-ledges, crags and pinnacles on

the highest parts of Burnt Island. No breeding

activity was apparent.

August 16, 1998: More than 2,000 adult

birds were feeding chicks and youngsters that

were hidden from view. Very few ready-to-fly

immatures were visible.

September 9, 1997: Only c. 200 adult terns

occupied the top of the islet. Feeding activity had

ceased; no juvenile birds were to be seen.

September 30, 1997: Over 300 birds were

spread all over the group of islets and tide-washed

rock-pinnacles in the vicinity.

October 14, 1989: Not a single tern

present on Burnt Island.

Non-breeding terns

Caspian tern Hydroprogne caspia (Pallas)

(Syn. No. 462):

Status: On passage in the vicinity of the

Vengurla Rocks.

On March 25, 1989 over 10 birds were

seen moving in the ‘archipelago’ and two

between the Vengurla Rocks and the mainland

on September 30, 1997.

Common tern Sterna hirundo Linn. (Syn. No.

465):

Status: Uncertain.

A desiccated head of a tern, collected by me

on Burnt Island on December 15, 1989, was

identified by S.A. Hussain at the BNHS, Mumbai,

as belonging to this species. Unfortunately, the

specimen has not been preserved in the collection.

White-cheeked Tern S. repressa Hartert (Syn.

No. 467):

Status: Formerly a breeding summer

visitor, now probably only on passage during the

annual southward mass-migration from July to

September and the less spectacular return flight

from March to May.

Madsen (1988) reported the sighting of

c. 20 birds on the seaward side of the group of

islets, on March 21,1 989. This species may have

been present in the roving flocks of unidentifiable

small white terns observed in May 1989.

Sooty tern S. fuscata Linn. (Syn. No. 474):

Status: Rare or occasional visitor.

Abdulali (1983) reported the sighting of a

few of these terns among a flock of c. 50 bridled

terns circling half a mile from the Rocks. On

April 30, 1998 I saw 8 birds in the vicinity and

identified them tentatively as of this species.

Sooty terns are uncommon, but far from rare,

visitors olf the South Konkan coast especially in

August-September during the annual mass-

migration of white-cheeked and brown-winged

(now bridled tern) terns (Lainer 1999a).

Lesser crested tern S. bengalensis Lesson (Syn.

No. 479):

Status: A roving visitor, regularly on passage.

Over 50 birds, in small groups, were

observed on northward migration on March 26,

1989. Southward passage in twos and threes was

recorded in September and October 1989 and

1997. Their flight path invariably lies between

the Vengurla Rocks and the mainland.

132

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MISCELLANEOUS NOTES

Sandwich Tern & sandvicensis Latham (Syn.

No. 480):

Status'. Unusual, on passage.

At least 3 birds were observed in the

vicinity on March 25, 1989. Groups of 20-25

birds kept passing in straight and purposeful

northward flight, about a kilometre from the

Rocks on May 13, 1989. Sandwich terns are

common visitors all along the South Konkan

coast throughout the year (Lainer 1 990).

Brown noddy Anous stolidus (Linn ,) (Syn. No.

481):

Status'. Stray or rare visitor.

On September 9, 1997, 18 birds were

observed perched on rock-ledges and stunted

bushes on the leeward side of Burnt Island. Three

weeks later, only a single brown noddy was found

there (Lainer 1999b).

Comments

Of the six tern species that were thought

to breed on Burnt island (S. dougallii,

S. repressa, S. anaethetus, S. fuscataS. albifrons

and S. bergii) the sooty tern should be omitted.

A minimum of 10,000 S. dougallii , 3.000

S. anaethetus and 2,000 S. bergii bred during the

southwest monsoon of 1998 on the leeward side

of Burnt Island. These numbers would increase

by 30% if one assumes that the over 10,000 terns

on the seaward side were of an identical or at

least similar species break-up.

Egg-laying and incubation started after

the traditional, regular onset of the southwest

monsoon in the area (June 5-7), probably after

the actual onset, if that should be at a later

date.

The simultaneous presence of fully-fledged

juveniles and adult birds still incubating in mid-

August suggests that second broods are taken up,

possibly after the loss of the first.

The surprisingly small number of fully-

fledged juveniles present towards the end of the

breeding season suggests that youngsters leave

the colony as soon as they are able to fly or soon

after. This ties in well with the annual southward

mass migration of S. anaethetus and S. repressa

off the coast of Goa, where 1 noticed that the

earlier part of the migratory stream (starting

usually at the end of August) is a trickle of mainly

juvenile birds, while the adults follow in a solid

rush from mid-September onwards.

The mixed ternery shows a strictly

stratified structure: S. dougallii , the smallest of

the constituent species, occupies the lowest rock-

strata, up to 10-15 m above the high-water line.

The medium-sized S. anaethetus , though

mingling with S. dougallii in the upper reaches

of their zone, breed at an intermediate height

while the large S. bergii adhere to the uppermost

parts of the islet.

Records of other bird species

Apart from terns, the following 26 bird

species were recorded on the Vengurla Rocks or

in the close vicinity:

1. Indian pond-heron Ardeola grayii

(Sykes) (Syn. No. 42): A single bird on Burnt

Island in December 1989.

2. Western reef-egret Egretta gu laris

(Bose) (Syn. No. 50): Single birds were seen in

March and November on Burnt Island.

3. Bra inn my kite Haliastur indus

(Boddaert) (Syn. No. 135): Singles among the

islets in March and May.

4. White-bellied sea-eagle Haliaeetus

leucogaster (Gmelin) (Syn. No. 173): Pairs and

singles were observed in February, March, May,

October and December.

5. Western marsh-harrier Circus

aeruginosus (Linn.) (Syn. No. 193): Two

immature birds over Burnt Island on September

30, 1997.

6. Osprey Pandion haliaetus (Linn.)

(Syn. No. 203): One to three birds were seen in

the archipelago in February, March, May (!),

September, October and December.

7. Peregrine falcon Falco peregrinus

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133

MISCELLANEOUS NOTES

Tunstall (Syn. No. 209): Singles were hunting

among the islets in March, October and

December.

8. Common kestrel Falco tinnunculus

Linn. (Syn. No. 222): A pair was recorded in

February 1875 by Hume (1876).

9. Common Sandpiper Tringa

hypoleucos Linn. (Syn. No. 401): Several birds

recorded in October and December.

10. Ruddy turnstone Arenaria interpres

(Linn.) (Syn. No. 402): Up to 4 birds on rocks

surrounding Burnt Island in May and August (!).

1 1 . Jaeger sp. Stercorarius sp.: Two adult

birds in non-breeding plumage among the islets

on March 25, 1989.

12. Heuglin’s gull Larus heuglini Bree

(Syn. No. 450): Nine birds passing through the

archipelago on northward migration on March

26, 1989.

13. Brown-headed gull Larus

brunnicephalus Jerdon (Syn. No. 454):

Hundreds among the islets in March and

December 1989.

14. Black-headed gull Larus ridibundus

Linn. (Syn. No. 455): Less numerous than the

preceding species in December 1989.

15. Blue rock pigeon Columba livia

Gmelin (Syn. No. 517): Breeding resident on

Burnt Island. Present throughout the year except

in July. Hume (1876) saw a few of these birds on

Burnt Island in 1875, Abdulali (1983) estimated

about 200 of what he deemed feral pigeons in 1981;

I noted well over 500 of them on June 5, 1997.

16. Oriental turtle-dove Streptopelia

orientalis (Latham) (Syn. No. 531): One coming

in from the sea and alighting on the base of the

lighthouse in October 1989; two birds on

Lighthouse Is. in December 1989.

17. Indian edible-nest swiftlet Coliocaiia

unicolor (Jerdon) (Syn. No. 685): Breeding

resident on Burnt Island, absent during the

southwest monsoon. The largest number 1 noted

were “tens of thousands, a virtual cloud over

Burnt Is.” on March 25, 1989.

18. House swift Apus affinis (J.E. Gray)

(Syn. No. 703): Up to 200 birds over the

archipelago in March and December.

19. Common hoopoe Upupa epops Linn.

(Syn. No. 765): A single on Lighthouse Is. on

October 14, 1989.

20. Greater short-toed Sark Calandrella

brachydactyla (Leisler) (Syn. No. 886): A couple

of flocks of c. 25 birds each passing over

Lighthouse Is. on October 15, 1989.

2 1 . Black Drongo Dicrurus macrocercus

Vieillot (Syn. No. 963): Two birds commuting

between Lighthouse Is. and Burnt Is. on October

14 and 15, 1989.

22. House Crow Corvus splendens

Vieillot (Syn. No. 1049): Singles on the

Lighthouse Is. in March and December.

23. Black-headed cuckoo-shrike

Coracina melanopiera (Ruppell) (Syn. No.

1079): A juvenile bird perched all morning in

the gantry of a crane on Lighthouse Is. on October

16, 1989.

24. Asian brown flycatcher Muscicapa

dauurica Pallas (Syn. No. 1407): A single bird

foraging on the rocky ground of the totally barren

Lighthouse Is. on October 15, 1989.

25. Blue rock-thrush Monticola solitarius

(Linn.) (Syn. No. 1726): Noted on all the islets

of the group in February, March and October.

26. Grey wagtail Motacilia cinerea

Tunstall (Syn. No. 1884): Two birds on

Lighthouse Is. on October 15, 1989.

Acknowledgement

I am grateful to the Director, Directorate of

Lighthouses and Lightships, Mumbai, for

permission to use the Inspection Quarters on

the Lighthouse Island on four occasions in 1989.

July 12, 2001 HEINZ LA1NER

Praias de St. Antonio,

Anjuna 403 509, Goa.

Email: [email protected]

134

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MISCELLANEOUS NOTES

References

Abdulali, H. ( 1 939): Swifts and Terns at Vengurla Rocks.

J. Bombay nat. Hist. Soc. 41(3): 661-664.

Abdulali, H. (1941): The Terns and Edible-nest Swifts at

Vengurla, West Coast, India. J. Bombay nat. Hist. Soc.

43(3): 446-45 L

Abdulali, H. (1983): Pigeons ( Columba livia) nesting on

the ground - some more bird notes from the Vengurla

Rocks. J. Bombay nat. Hist. Soc. 80(1): 215-217.

Ali S. & S.D. Ripley (1983): Compact Handbook of the

Birds of India and Pakistan together with those of

Bangladesh, Nepal, Bhutan and Sri Lanka. Oxford

University Press, Delhi, pp. 737 + 104 plates.

Grimmett, R., C. Inskipp & T. Inskipp (1998): Birds of the

Indian Subcontinent. Christopher Helm, A & C Black,

London.

Harrison, P. (1985): Seabirds. Revised edition. Christopher

Helm, A & C Black, London.

Hume, A.O. (1876): The Laccadives and the West Coast.

Stray Feathers 4: 413-483.

Kazmierczak, K. (2000): A Field Guide to the Birds of

India. Om Book Service, New Delhi.

Lainer, H. (1990): On the Status of the Sandwich Tern

Sterna sandvicensis on India’s West Coast. J. Bombay

nat. Hist. Soc. 87(2): 298-299.

Lainer, H. (1999a): The Birds of Goa. J. Bombay nat.

Hist. Soc. 96(2): 203-220, (3): 405-423.

Lainer, H. (1999b): The Noddy Tern (Brown Noddy)

Anous stolidus off the South Konkan Coast. J. Bombay

nat. Hist. Soc. 96(3): 469.

Madsen, S.T. (1988): Terns of the Vengurla Rocks.

Hornbill 1988(1): 3-4.

Ripley, S.D. (1982): A Synopsis of the Birds of India

and Pakistan. Bombay Natural History Society,

Bombay.

Vidal, G.W. (1880): First list of the birds of the south

Konkan. Stray Feathers 9: 1-96.

23. RANGE EXTENSION OF ALEXANDRINE PARAKEET

PSITTACULA EUPATRIA IN GUJARAT

In Gujarat State, the Alexandrine parakeet

Psittacula eupatria is known to occur in Mahal

(Dang), Waghai (Surat), Rajpipia (Narmada), and

Gangasagar (Banaskantha) districts (Ali 1954). In

recent publications, the species is reported only

from the Rajpipia forest, but is uncommon (Monga

and Naoroji 1983, Desai et al. 1993, Narve et al.

1 997). Khacher ( 1 996) and Grimmett et al. (1998)

do not add new information about its distribution

in Gujarat.

In Oct. 1992 and 1993, we had seen the

species feeding on maize Zea mays cobs at Vatrak,

Modasa and Himmatnagar of Sabarkantha district.

After that, almost every year, a few pairs can be

seen on a river bridge at Vatrak. We saw the bird

first at Anand (Anand district), on Apr. 17, 1999

in the Institute of Rural Management campus.

After that there were several records every month.

At Nadiad (Kheda district), Shri Prathmesh Patel

(pers. comm, in 1999) saw 3 wild birds hovering

around his captive P. eupatria during Nov. 1999.

At the Indian Petrochemicals Corporation Ltd.

Township, Vadodara (Vadodara district), it was

heard frequently over the last 3 years. We saw one

bird on a banyan tree ( Ficus religiosa) on Jul. 14,

1999 at Jambughoda Wildlife Sanctuary and

several birds on Feb. 20 & 2 1 , 2000 at Ratanmahal

Wildlife Sanctuary (both in Panchmahal district).

There is a recent record of its occurrence from

Ahmedabad city (Ketan Tatu pers. comm, in 2000)

and BMP saw one bird in flight on Nov. 6, 2000

at Kalupur Railway station, Ahmedabad. At

Bhavnagar, four adult birds are regularly seen since

1998 (Indra Gadhavi pers. comm, in 2000).

The above records prove that the species has

extended its distributional range in Gujarat State

and also breeds in some parts of Kheda and

Sabarkantha districts. Like elsewhere in India

(Mehrotra and Bhatnagar 1979), it raids maize

crops in Gujarat too. Our observation of a few pairs

around holes on the river bridge indicate a search

for unusual nesting structures by these birds,

March 23, 2001 AESHITA MUKHERJEE

C.K. BORAD

*B.M. PARASHARYA

AINP on Agricultural Ornithology,

Gujarat Agricultural University,

Anand 388 110, Gujarat, India.

* Email: [email protected]. in

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

135

MISCELLANEOUS NOTES

Refer

Ali, S. (1954): The Birds of Gujarat, Part I. J. Bombay

nat. Hist. Soc. 52(2 & 3): 374-458.

Desai, I.V., B. Suresh & B. Pilo (1993): Birds of

Shoolpaneshwar Wildlife Sanctuary. Pavo31(l cfc 2):

5-72.

Grimmett, R., C. Inskipp & T. Inskipp ( 1 998): Birds of the

Indian Subcontinent. Oxford University Press, Delhi.

Khacher, L. (1996): The Birds of Gujarat — A Salim Ali

Centenary year review. J. Bombay nat. Hist. Soc.

93(3): 331-373.

24. MIMICRY OF A CROW CHICK BY AN

AS A DEFENCE AGAINST ATTACK BY

The Asian koel Eudynamys scolopacea is

a common brood parasite of corvids, mynas and

other species throughout India and much of

southeast Asia. In India, this cuckoo’s

predominant host is the house crow Corvus

splendens and to a lesser extent the jungle crow

C. macrorhynchos (Lamba 1976). This author

has observed that the high populations of house

crows in towns and villages accounts for the large

concentrations of Asian koels that may

sometimes occur there. Despite the commonness

of this brood parasitic relationship, the exact way

in which Asian koel gains access to the hosts’

nest for egg laying remains a point of

controversy.

An interaction observed between an Asian

koel and a group of house crows in the village of

Anjuna, Goa on December 17, 1999, at a time

when house crows were breeding in the area, may

throw light on this question.

Observation

At dusk, about 1800 hrs, a group of five

house crows were seen noisily chasing a female,

or perhaps a juvenile, Asian koel. The latter

alighted in the top of a coconut palm and was

immediately surrounded by the house crows that

were cawing loudly and aggressively. One crow

moved within c. 30 cm of the Asian koel, and

ENCES

Mehrotra, K.N. & R.K. Bhatnagar (1979): Status of

Economic Ornithology in India - Bird Depredents

(sic). Depredations and their Management. Indian

Council of Agricultural Research, New Delhi.

Monga, S.G. & R. Naoroji (1983): Birds of the Rajpipla

forests — south Gujarat. J. Bombay nat. Hist. Soc.

80(3): 5 75-612.

Narve, D.S., C.D. Patel & N.P. Pandya (1 997): Avifaunal

diversity in Shoolpaneshwar Sanctuary. Tigerpaper

24(1): 17-22.

ASIAN KOEL EUDYNAMYS SCOLOPACEA

HOUSE CROWS CORVUS SPLENDENS

seemed about to launch a pecking attack. At that

moment, the Asian koel responded to this threat

by opening its mouth wide to reveal the bright

red gape, which was held upwards, and emitting

a call closely resembling that of a begging house

crow chick.

This display by the Asian koel resulted in

an instantaneous halt to the attack by the

approaching crow, while the other four crows also

became silent and passive. After about two

minutes, during which time no further aggressive

behaviour was displayed by the crows, the Asian

koel slipped away into the semi-darkness of the

coconut grove. The light was now failing rapidly

and after a few minutes more of perching around

abstractedly, the crows also flew off singly.

Discussion

House crows are well known to behave

aggressively towards both sexes of the dimorphic

Asian koel, particularly during the breeding

season, chasing them on sight even far from nest,

sometimes physically attacking them (Hume

1889; Lamba 1963) and even in rare cases killing

them (Lamba 1976). The peak breeding season

for house crows in southern India is April to May

but some breeding, as in the present case, takes

place in November and December (Lamba 1963).

Given this aggressive behaviour on the part

136

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

MISCELLANEOUS NOTES

of the hosts, there has been discussion and

speculation over the years as to how the female

Asian koel finds the opportunity to deposit her

eggs in the vigilant house crow’s nest. Dewar

( 1 907) and Dharmakumarsinhj i ( 1 954) described

several instances in which Asian koels appeared

to take advantage of this aggressive response to

facilitate access to the house crow’s nest for the

egg loaded female Asian koel. Dewar proposed

a scenario in which the black male Asian koel

lures the sitting house crow away from the nest,

so that the cryptically marked brown-barred

female Asian koel can slip in to deposit its egg.

Although Lamba ( 1 963) at one time accepted this

hypothesis, by 1976, his own extensive studies

had led him to believe that such observations

were coincidental rather than a well-orchestrated

strategy and that, in fact, it was simply a case of

the female Asian koel taking any opportunity to

get to the unoccupied nest. Furthermore, Eates

(undated) described three instances where female

Asian koels were seen in nests alongside

incubating house crows, flapping and jockeying

for position, and that calls resembling those of

young house crows were heard. In each case, the

Asian koel laid an egg and was not attacked by

the resident crow. This suggests that the female

Asian koel produced a call like a young house

crow to appease the rightful occupant of the nest.

The instance described here, not only

involved mimicry of a crow chick, but also of

the chick’s gape and begging behaviour. The

Asian koel had a bright red gape, resembling

that of a house crow chick. Goodwin (1986) also

described the inside of the house crow chick’s

mouth as fleshy red and Lamba (1976) as blood

red. Interpretation of the observation described

Refe

Dharmakumarsinhji, R.S. (1954): Birds of Saurashtra.

Published by the Author, Bhavnagar. 562 pp. Cited :

Lamba, B.S.(1976).

Dewar, D. (1907): An enquiry into the parasitic habits of

the Indian koel. J. Bombay ncit. Hist. Soc. 1 7: 765-

here depends on whether the Asian koel in

question was an adult or a juvenile. Stuart

Butchart (pers. comm.) pointed out that fledgling

Asian koels have a bright red gape, but that this

probably becomes duller in adults, as is the case

with other cuckoos. As no gape flange was

visible, the bird would not have been a recent

fledgling. However, older fledglings do indeed

resemble females. If the Asian koel were a

juvenile, it may have provoked a mixed response

from the crows: mobbing whilst in flight,

followed by tolerance once begging was initiated.

In any case, it seems likely from the various

published descriptions mentioned above, that

female Asian koels retain this fledgling-like

behaviour and may resort to mimicry of house

crow chicks to avoid physical attack when they

have been cornered and are unable to escape. In

the case of Eates’ observations, perhaps the

female koels were in the process of egg laying in

an unoccupied nest when the house crow returned

and, through mimicking a chick, the Asian koel

was able to finish depositing an egg and depart

without attack. The key to understanding this

interaction is whether the gape colour of the adult

female Asian koel is able to elicit the appropriate

parental response in house crows. Clearly, such

an ability would have enormous survival value,

not only for the individual Asian koel, but also

for the species as a whole.

April 24, 2001 COLIN RYALL

Centre for Environmental Management,

Farnborough College,

Boundary Road, Farnborough,

Hampshire, GUM 6NU, UK.

Email: [email protected]

E N C E S

782.

Eates, K.R. (unpublished): Memories grave and gay of a

field naturalist. Section 4, Crows, British Museum

(Nat. Hist.) tiles. Cited: Goodwin, D. (1986).

Goodwin, D. (1986): Crows of the World. Cornell

JOURNAL. BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

137

MISCELLANEOUS NOTES

University Press. Pp. 100.

Hume, A.O. (1889): Nests and eggs of the Indian Birds:

Vol. 1, 2ndedn. R.H. Porter, London.

Lamba, B.S. (1963): The nidification of some common

Indian birds Part \ .J. Bombay nat. Hist. Soc. 60(1):

121-133.

Lamba, B.S. (1976): The Indian crows: a contribution to

their breeding biology, with notes on brood

parasitism on them by the Indian Koel. Rec. zoo/.

Surv. India 71: 183-300.

25. BUFFY FISH-OWL {KETUPA KETUPU) IN SUNDARBANS, BANGLADESH

{With one plate )

Three species of fish-owl are known from

the Indian subcontinent: the brown fish-owl

{Ketupa zeylonensis), the tawny fish-owl ( Ketnpa

flavipes) and the buffy fish-owl ( Ketnpa ketnpn).

The last sightings of the buffy fish-owl from

Assam were recorded by Stevens (1915) and

Baker (1927). Stevens writes “Common in the

forest streams which emerge into the Dibru” and

mentions localities “Rungagora, 1902” and

“Dejoo, North Lakhimpur, 1911”. Baker says “I

found it not very rare in the hills of South Assam

and Coltart obtained one specimen in

Dibrugarh”. Further, “Coltart and I obtained eggs

in the Khasia Hills and North Cachar”.

The buffy fish-owl, also called the Malay

fish-owl, is common in southeast Asia. According

to Koenig et al. ( 1 999) and del Hoyo et al. ( 1 999),

its present range of distribution covers Myanmar,

Thailand, Vietnam, the Malaysian Peninsula and

Indonesia (including Sumatra, Java, Borneo).

Fish-owls are not rare in the Sundarbans,

and the huge mangrove forest in the lower delta

of the Ganga, but are usually hard to identify

when only seen in flight in poor light conditions.

Some more recently published bird lists (Khan

1986; Hussain and Acharya 1994) mention

brown fish-owl and tawny fish-owl, or only

brown fish-owl, to occurring here.

It was only in November 2000, that we

were able to photograph fish-owls in the

southeastern Sundarbans of Bangladesh (Kotka

Sanctuary - Plate 1) in early mornings and late

afternoons. Analysing the photographs of three

different individuals, all of them could be

identified as buffy fish-owls.

Main characteristics of the buffy fish-owl:

— at 38-48 cm it is smaller than the brown fish-

owl (56 cm) and tawny fish-owl (58-61 cm);

— lacking fine horizontal cross-barrings on

underparts (typical for the brown fish-owl);

— colour above rich brown (against rich orange-

rufous to tawny of the tawny fish-owl, and

duller brown of the brown fish-owl);

— bare tarsi (against partly feathered tarsi of

the tawny fish-owl);

— relatively short tail with few whitish bands

(against longer, more narrowly barred tail

of the tawny fish-owl);

— wings more broadly and buff sh-white barred

(against less broad orange-buff barrings of

the tawny f sh-owl).

This is, as far as known, the first record of

the buffy fish-owl from Sundarbans and a re-

discovery of the species in the Indian

subcontinent (last record by Baker) after almost

80 years! It is perhaps not impossible that the

buffy fish-owl was overlooked or mistaken for a

tawny fish-owl, as Ali and Ripley (1969) along

with their pictorial guide (1983), widely used. in

the past decades for identification of owls on the

Indian subcontinent, does not mention the

species. We became aware of the buffy fish-owl

when consulting Grimmett et al. (1998).

The fauna of Sundarbans has, besides the

buffy fish-owl, several other affinities with the

Malaysian Region. Paynter (1970) mentioned

laced woodpecker, blue-winged pitta, mangrove

whistler and orange-bellied flowerpecker in this

context (specimens collected in 1958 in the

Sundarbans of Bangladesh). Other species from

138

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY 100(1), APR. 2003

MISCELLANEOUS NOTES

Neumann-Denzau, Gertrud and Helmut Denzau: Ketupa ketupu

Plate 1

Fig. 1: Buffy fish-owl Ketupa ketupu in Sundarbans, Bangladesh

a. Individual 1 (front view), b. Individual 1 (side view), c. Individual 2, d. Individual 3

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

139

MISCELLANEOUS NOTES

Sundarbans with the same affinity are the masked

finfoot, and not to forget the Javanese rhino

among the exterminated Sundarbans mammals.

More careful observations in future will surely

bring forth other interesting discoveries from the

Indian and Bangladesh Sundarbans.

Mar. 23, 2001 GERTRUD NEUMANN-DENZAU

HELMUT DENZAU

fm Brook 8,

2 4321 Ranker,

Germany.

Email: [email protected]

References

Ali, S. & S.D. Ripley (1969): Handbook of the Birds of

India and Pakistan together with those of

Bangladesh, Nepal, Bhutan and Sri Lanka. Oxford

University Press, Bombay, Vol. 3.

Baker, E.C.S. (1927): The Fauna of British India. Taylor

and Francis, London, Vol. 4: 410-411.

del Hoyo, J., A. Elliott & J. Sargatal (Eds) (1999):

Handbook of the Birds of the World. Lynx Edicions,

Barcelona, Vol. 5 (Buffy fish-owl : 1 94).

Grimmett, R., C. Inskipp & T. Inskipp (1998): Birds of

the Indian Subcontinent. Christopher Helm,

London.

Hussain, Z. & G. Acharya (1994): Mangroves of the

Sundarbans. Vol. 2. Bangladesh, IUCN, Bangkok.

Pp. 248.

Khan, M.A.R. (1986): Wildlife in Bangladesh Mangrove

Ecosystem. ./. Bombay nat. Hist. Soc. 83: 32-48.

Koenig, C., F. Weick & J.H. Becking (1999): Owls, A

Guide to the Owls of the World. Pica Press,

Sussex, pp. 309-310 + pi. 33.

Paynter, R.A. (1970): Species with Malaysian affinities

in the Sundarbans, East Pakistan. Bull. Brit. Orn.

Club 90(5): 118-119.

Stevens, H. (1915): Notes on the Birds of Upper Assam,

Part 2. J. Bombay nat.. Hist. Soc. 23: 547-570

(Malay Fish Owl: 560).

26. A NOTE ON THE CIRCUMORBITAL SKIN COLOUR

OF INDIAN GREY HORNBILL OCYCEROS BIROSTR1S

On August 8, 2000, I was attracted by the

calls of Indian grey hornbills Ocyceros birostris

(Scopoli) from a large fruiting p i pal Ficus

religiosa Linnaeus, in my garden in Hyderabad.

This was the third consecutive day that I was

watching this trio of 2 males and 1 female. It

was obvious they were courting, for the males

displayed and postured with upright body and

bill pointed skyward. While doing so, the

feathers above their tarsi were puffed up, like

small balls of white cotton. Their white-tipped

tails were constantly and rhythmically pumped

in an arc (swung like a pendulum if the birds

were viewed in profile) from their normal

positions, inwards. Their loud calls resounded

through the garden. Males uttered a shrill and

quavering squeal, while the female’s call was a

clearer and louder, nasal “wheeeee”. The birds

also uttered a ratchety uk-k-k-k-k-k-krrr” (maybe

only male) as they moved upward, towards the

top of the tree. All three birds hopped and

clambered about in the tree, using their bills to

pull themselves up at times, plucking and eating

figs as they proceeded. Twice a male hopped

along a branch towards the female and acted as

if he was going to feed her, though no item of

food was passed.

I got a very good view of all three as they

moved about. I noticed almost immediately that

the circumorbitai skin on the female was a rich

dark orange and her irides were dark brown. Both

the males had black circumorbitai skin and

orange irides. While the colours of these soft and

bare parts of a male are recorded in published

literature (Ali and Ripley 1987, Baker 1927,

Kemp 1995, Roberts 1991), there is no mention

of the circumorbitai skin colour of a female

Indian grey hornbill (Kemp 1995).

On August 29, 2000, I was in Mumbai and

took the opportunity to check the specimens in

JOURNAL BOMBAY NATURAL HISTORY SOCIETY. 100(1), APR. 2003

141

MISCELLANEOUS NOTES

the collection of the Bombay Natural History

Society. I examined 11 females, of which 9 had

clear orange skin around the eyes. The

circumorbital skin colour was not clear on

specimen # 1 1447, collected by P.T.L. Dodsworth

on 1 3 .i. 1 9 1 3 from Karka, N.W. Himalayas. A

remark on the tag read, “Irides reddish-brown.”

Specimen # 1 1450, collected by V.S. Lapersonne

on 29. v. 1929 from Chitteri Range, Salem at

2,000, was marked as a female and had a quarter

moon-shaped orange area below the eye. This

specimen also had a relatively more prominent

casque than that of the others.

It is, however, possible that the colours of

bare skin and irides may vary according to the

emotional state of the bird and the resulting

endocrinal secretions. Outside of the breeding

season, probably the bare skin of both sexes may

well be dark grey to black, and that of the female

Refer

Ali, S. & S.D. Ripley (1987): Compact Handbook of the

Birds of India and Pakistan together with those of

Bangladesh, Nepal, Bhutan and Sri Lanka. 2nd

edition, Oxford University Press, Delhi. Pp. 292.

Baker, E.C. Stuart (1927): The Fauna of British India

including Ceylon and Burma. Birds. Vol. IV, 2nd

changes only during courtship and times of

excitement.

The shape of the casque, the extent of

yellow on the bill, and the colour of the irides

are used to separate sexes of the Indian grey

hornbill in the field. These field-marks can now

be supplemented by the differences in their

circumorbital skin also.

Acknowledgements

I thank S.A. Hussain, T.J. Roberts and R.

Kannan for their comments on these observations.

March 23, 2001 AASHEESH PITTIE

8-2-545 Road No. 7, Banjara Hills,

Hyderabad 500 034,

Andhra Pradesh, India.

Email: [email protected]

E N C E S

edition. Taylor and Francis, London. Pp. 301-302.

Kemp, Allan (1995): The Hornbills. Oxford University

Press, Oxford. Pp. 157-159.

Roberts, T.J. (1991): The Birds of Pakistan. Vol. 1.

Regional Studies and non-Passeriformes. Oxford

University Press, Karachi. Pp. 532-534.

27. SIGHTING OF MALABAR PIED HORNBILL ANTHRACOCEROS CORONATES

IN SANJAY GANDHI NATIONAL PARK, MUMBAI

On July 30, 2000, during a walk in the

Sanjay Gandhi National Park (SGNP), Mumbai,

at 0845 hrs we stumbled upon a noisy flock of

Indian grey hornbill Ocyceros birostris. There

were four individuals to our left, coursing

restlessly through the thick canopy about 40 m

away. We were at an elevation and the birds,

though about 1 0 m from the ground were exactly

level with us. While we were contemplating the

reason for such aggressive behaviour, another

much larger and darker bird emerged from the

thick cover. It was still partly hidden and difficult

to identify, but as soon as it alighted from a tall

Adina cordifolia tree, the white trailing edge on

its predominantly black wings and white outer

tail feathers disclosed its true identity. The bird

being mobbed by the four Indian grey hornbills

was the Malabar pied hornbill Anthracoceros

coronatus. Soon, A. coronatus was chased away

by the four birds and the flock disappeared into

the canopy.

However, within five minutes these noisy

hornbills returned to the same area, but perched

slightly closer to us. This time there were two

A. coronatus. One bird was identified to be a

female from the absence of the black patch at

the posterior end of its casque, while the sex of

the other bird could not be determined. The four

O. birostris always kept a close vigil, but never

advanced too close to the larger species.

142

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

MISCELLANEOUS NOTES

O. birostris has been observed to nest in this area,

due to the presence of huge trees, and its

aggression towards its conspecifics could be

justified. However, both birds breed between

March and June in their respective ranges (Ali

and Ripley 1987), and a fight over a nesting site

is quite unlikely in July. A detailed study is

required to comment conclusively on it.

A. coronatus was never reported from

Sanjay Gandhi National Park until February 6,

2000, when one bird was sighted by a group of

bird watchers in the same valley, called the

Pongam Valley due to a profusion of Pongamia

pinnata. Subsequently, there have been a few

confirmed and unconfirmed records of the bird

from various parts of the Park.

According to Ali (1996), the bird is found

in south and central India: north to southeast

Uttar Pradesh, Bihar and Orissa. Grimrnett et

al. (1999) mention its distribution in the Western

Ghats and east India.

Anthracoceros coronatus is mainly

frugivorous, but can also subsist on small reptiles,

mice and juvenile birds as has been reported by

Ali and Ripley (1987). The present sighting of

A. coronatus is almost six months after its first

record from SGNP. It is evident, therefore, that

the birds, which presumably strayed or were

released into SGNP (a Tropical Moist Semi-

deciduous Forest) outlived the harsh summer and

acclimatized to their newly found home. Our next

step should be to examine whether the species

attempts to breed here, provided that both sexes

are present.

Additional Note from first author

I was at Sanjay Gandhi National Park on

March 9, 2003 and was fortunate to spot a female

A. coronatus with a flock of O. birostris in

precisely the same spot as the first sighting. Over

three years have passed since A. coronatus was

first sighted, and since then, a female bird has

been regularly seen by many bird watchers

including the first author near the Pongam Valley,

moving boldly with a flock of O. birostris. It is,

therefore, clear that the bird has adapted to the

Moist Semi-deciduous Forest of SGNP. The most

striking feature of the sighting is that the larger

A. coronatus seems to be enjoying a congenial

relationship with the smaller, but more numerous

hornbill O. birostris. I have seen the two species

in other forests of India as well, sometimes even

sharing the same tree, but have never observed

any significant interaction between the two.

Thus, this three-year association between the two

related yet distinct species is unique and worth

mentioning — A. A.

October 16, 2000 ANISH ANDHERIA

2, Sagar Building,

VP. Road, Andheri (West),

Mumbai 400 058,

Maharashtra, India.

Email: [email protected]

SUPRIYA JHUNJHUNWALA

Bombay Natural History Society,

Hornbill House, S.B. Singh Road,

Mumbai 400 023, Maharashtra, India.

Email: [email protected]

PARITOSH KHANVILKAR

5-Parimal, 1 7th Road,

Khar (W), Mumbai 400 052,

Maharashtra,

India.

References

Ali, S. & S.D. Ripley (1987): Compact Handbook of the

Birds of India and Pakistan together with those of

Bangladesh, Nepal, Bhutan and Sri Lanka. 2nd edn,

Oxford University Press, Delhi, pp. 737 + 104

plates.

All S. (1996): The Book of Indian Birds (12th Edn).

Bombay Natural History Society, Mumbai.

Grimmett, R., C. Inskipp & T. Inskipp ( 1 999): Birds of the

Indian Subcontinent. Oxford University Press, New

Delhi.

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 1 00(1), APR. 2003

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MISCELLANEOUS NOTES

28. NOCTURNAL FEEDING BY WHITE-BELLIED DRONGO DICRURUS CAERULESCENS

Van Ashram situated on the outskirts of

Sawai Madhopur city, has a Forest Rest House

and a Range Office. A forest nursery is also

present on the campus.

On May 14, 1999, while I was observing

nocturnal animals in and around Van Ashram,

at c. 2030 hrs, I came across three white-bellied

drongos Dicrurus caerulescens hovering near an

electric light bulb, in front of the Forest Rest

House, devouring insects. All the three birds were

perched on a wire. One more bird, feeding near

another bulb, was perched on a bough of

Adansonia digitata. I observed all the four birds

up to 2200 hrs. When I left, they were still

feeding. The following night, 1 saw perhaps the

same birds feeding on flying insects.

Nocturnal feeding by black drongo

Dicrurus adsimilis , inside human habitation, has

been reported by Khan (1990), Nameer (1990)

and Sharma (1991). The white-bellied drongo is

more arboreal in nature than the black drongo,

and it is a forest bird. It generally avoids human

habitation. The gradual destruction of forests

around the periphery of the expanding city and

reduction in the food sources is perhaps forcing

this bird to change its behaviour. Night light

sources of the city attract huge insect populations

from surrounding forest areas, with perhaps a

resultant fall of insect density in surrounding

forests. This in turn may lead to forest birds like

the white-bellied drongos following their food

resource for nocturnal feeding to the city.

February 6, 2001 SATISH KUMAR SHARMA

Phulwari Wildlife Sanctuary,

Kotra 307 025, District Udaipur,

Rajasthan,

India.

References

Khan, A.R. (1990): Feeding habits of the Black Drongo. Newsletter for Birdwatchers 30(7 & 8): 9.

Newsletter for Birdwatchers 30(10 & 11): 11. Sharma, S.K. (1991): Nocturnal feeding by Black Drongo.

Nameer, RO. ( 1 990): Midnight feeding by Black Drongo. Newsletter for Birdwatchers 31(3 & 4): 8.

29. TOTAL ALBINISM IN A LARGE GREY BABBLER TURDOIDES MALCOLM /

On July 29, 2000, while I was surveying

the biodiversity of Kumbhalgarh Wildlife

Sanctuary, a milk-white large grey babbler

Turdoides malcolmi caught my attention near

Nandeshma village, on the outskirts of the

Sanctuary. It was an adult bird with completely

white plumage. The legs and bill were lighter

than normal individuals. It was the only albino

in a flock of eight birds.

When I approached the flock to get a

photograph, the albino took off and hid in the

foliage of a nearby mango tree, while the rest of

the flock remained on the ground in a semi-alert

posture. I waited motionless behind a Butea

monosperma tree for the bird. When it came

down after a few minutes, I tried going near, but

this time too it took refuge in the foliage of

another tree. After a few minutes, it alighted on

a rock near its flock and this time I could

photograph it [photographic evidence given by

the author — Eds]. I observed this bird for nearly

15 min and found it to be quite shy compared to

normal individuals.

Albinism has been reported in crows

(Mahabal 1991; Abdulla 1997), doves (Javed

1992; Pandya 1994), redwattled lapwing (Soni

1992), Asian koel (Shyamal 1990), little grebe

(Bharos 1 996), coot (Paf ashary a et a/. 1 996), red-

vented bulbul (Soni 1992; Joshua 1996), lesser

whistling-duck (Chatterjee 1995) and common

144

JOURNAL , BOMBAY NATURAL HISTORY SOCIETY 100(1), APR 2003

MISCELLANEOUS NOTES

myna (Jha 1994). The large grey babbler is an

addition to the list of albino birds, hence worth

placing on record.

Acknowledgement

I thank R.G. Soni, PCCF and CCF (WL),

Rajasthan for giving me an opportunity to study

Refer

Abdulla, E.V. (1997): White jungle crow. Newsletter for

Birdwatchers 37(5): 91 .

Bharos, A.M.K. (1996): Albino little grebe Tachybaptus

ruficollis. J. Bombay nat. Hist. Soc. 93(2): 293.

Chatterjee, S. (1995): Occurrence of albino lesser

whistling teal Dendrocygna javanica (Horsfield).

J. Bombay nat. Hist. Soc. 92(3): 417-418.

Javed, S. (1992): Albinism in little brown dove. Newsletter

for Birdwatchers 32(3 & 4): 12.

Jha, Samiran ( 1 994): An albino myna Acridotheres tristis

(Linnaeus). J. Bombay nat. Hist. Soc. 91(3): 455.

Joshua, .1.(1 996): An albino redvented bulbul Pycnonotus

cafer. J. Bombay nat. Hist. Soc. 93(1): 586.

the biodiversity of Protected Areas of southern

Rajasthan.

October 16, 2000 SATISH KUMAR SHARMA

Phulwari Wildlife Sanctuary,

Kotra 307 025, District Udaipur,

Rajasthan,

India.

; N C E s

Mahabal, A. (1991): Cases ofalbinism in house and jungle

crows. Newsletter for Birdwatchers 31(9 & 10):

14.

Pandya, V. (1994): Sighting of albino dove. Newsletter

for Birdwatchers 34(4): 97.

Parasharya, B.M., R.B. Chaijhan & A.G. Sukhadia

(1996): A white coot at Kanewal, Gujarat.

J. Bombay nat. Hist. Soc. 93(1): 586.

Shyamal, L. ( 1 990): Partial albinism in a koel ( Eudynamys

scolopacea). Newsletter for Birdwatchers 30(10

& 11): 11.

Soni, R.G. (1992): Albinism in Birds. Newsletter for

Birdwatchers 32(3 & 4): 13.

30. YELLOW-RUMPED FLY CATC PIER FICEDULA ZANTHOPYG1A IN KERALA

On January 30, 1996 along the Mangala

Devi trail at Periyar National Park, Kerala, I

encountered a female yellow-rumped flycatcher

Ficedula zcmthopygia . The bird spent most of

its time between 5-10 m up in the mid-canopy of

trees on the edge of the forest . I watched the bird

for about 20 min ranging at c. 8 m, I was familiar

with the species from China and Thailand,

identified it quickly and confidently and,

knowing that it was a vagrant to India, made a

quick sketch and a few hurried field notes. Some

of the notes are reproduced here.

It was a distinctive bird that could perhaps

only be confused with the female of the Chinese-

breeding form of Narcissus flycatcher

F. narcissina elisae. However, elisae can be

distinguished, among other features, by the lack

of the conspicuous yellow rump of the aptly

named yellow-rumped flycatcher.

Description

Estimated to be about 10% larger and

noticeably bulkier, more plump, proportionately

larger headed and heavier billed than a red-

throated flycatcher F parva.

The upperparts were a fairly uniform dull

olive-green; the nape was subtly greyer and the

rump, undoubtedly the bird’s most striking

feature, was bright lemon yellow. This was visible

at rest, when the wings were frequently held

drooped, but was most conspicuous during the

bird’s short flights. The upper border of the rump

fell level with the tip of the bird’s shortest tertial

while the uppertail coverts (and tail) were

blackish and contrasted sharply.

The head was quite well marked. The ear-

coverts were slightly greyer than the crown and

nape and there was a narrow off-white band

JOURNAL . BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

145

MISCELLANEOUS NOTES

connecting the pale lores across the lower

forehead. This, combined with the rather broad

off-white eye-rings, created a bespectacled facial

pattern reminiscent of the plumbeous vireo Vireo

plumb eus.

The underparts were pale yellowish-white,

somewhat scruffy in that the yellow tone was not

evenly distributed but patchy. The vent and

undertail coverts were ciean white and there was

some slight olive mottling, almost scaling, on

the sides of the breast. The three innermost

greater coverts had conspicuous blackish centres,

broad white tips and narrower white fringes on

the outer webs. Two, or possibly three, median

coverts were similarly patterned with dark centres

and whitish edges. The longest tertial also had a

prominent white fringe.

The bill was entirely black, stout and

heavy-looking, the legs a paler plumbeous grey,

and the eye dark and appearing large.

The bird called several times during the

observation — a dry, hollow sounding ‘ trrrrt\

that was mellower and softer than the similar

contact calls of the red-throated flycatcher.

This was the second documented record

of the yellow-rumped flycatcher from the

Subcontinent. The first, a male in the Melghat

Sanctuary in Maharashtra on the April 30, 1989,

was documented by Haribal (1991). Interestingly,

a bird answering the description of a female

yellow-rumped flycatcher was apparently also

present at Melghat in the winter of 1994-95. It

was described to K. Kazmierczak by the local

Forest Department staff who had been unable to

identify it and were unaware of the previous

record. There is another more recent report — a

male observed by Steve Rooke and Deepal

Warakagoda et al. in Sri Lanka on the March 7,

1999. (Steve Rooke pers. comm.)

ACKNO WLEDG EM ENT

I thank Krys Kazmierczak for useful

information regarding the earlier sightings and

the encouragement to publish this one.

November 25, 2000 PAUL I. HOLT

New Laund Farm,

Greenhead Lane,

Fence, Burnley,

Lancashire, BB12 9DU, UK.

Email: [email protected]

Reference

Haribal, M. (1991): Yellowrumped flycatcher Ficedula (Muscicapa) zanthopygia (narcissina): a new addition to the

avifauna of the Indian subcontinent. J. Bombay nat. Hist. Soc. 88: 456-458.

31. BREEDING RECORDS OF THE ASIAN BROWN FLYCATCHER

MUSCICAPA DAUURICA IN SOUTHERN INDIA

Commenting on the status and distribution

of the Asian brown flycatcher ( Muscicapa

dauurica), Ali and Ripley (1983) mention: “A

partial migrant having a disjunct breeding range.

Movements imperfectly understood.” More

specifically of its status in southern India, they

say that the bird is a scarce breeder in the

southern parts of the Western Ghats at about

900 m in North Kanara, Coorg, the Palni Hills

and Cardamom Hills. It is also suspected that

the flycatcher may breed in the Chitteri Range

(Eastern Ghats) where it has been obtained in

June, and in the Biligirirangan Hills (obtained

in July). Post-dispersal of southern birds takes

place as early as August. Ghorpade (1973) has

mentioned it as a resident species in Sandur

(E. Karnataka). Recently, Pittie (2000) recorded

a young bird in June in Hyderabad.

My own records for the last two decades

indicate that the Asian brown flycatcher could

146

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

MISCELLANEOUS NOTES

be resident in small numbers in the

neighbourhood of Chennai city, Tamil Nadu and

in the Rishi Valley area (Chittoor district, Andhra

Pradesh), I have also seen a nest and a juvenile

in the Peechi-Vazhani Wildlife Sanctuary in

Trichur district, Kerala. I present the details

below.

In Chennai, I have 1 2 records of the bird

in the Guindy National Park and Theosophical

Society estate during June-August (1984-1990).

Usually single birds were noticed. Juvenile birds

with spotted plumage were seen on at least two

occasions — on July 29, 1984 and July 3, 1990

at the Guindy Park. On the latter occasion, an

adult bird was also seen.

In Rishi Valley ( c . 700 m), I have more

than 20 records of the bird in June-August, when

more than one bird was present on at least six

occasions during these months. Juvenile birds

were seen in July-August 1999 and in June-July

2000.

At the Peechi-Vazhani Wildlife Sanctuary

(c. 100 m), a nest was seen on March 2, 1993,

atop a dead tree at c. 17 m inside a hollow. On

August 26, 1991, 1 had seen a juvenile with spots

at Mannuthy close to the Sanctuary.

These records show that the Asian brown

flycatcher may nest in suitable localities in small

numbers in peninsular India, and more

information can be collected by birdwatchers by

careful observations.

March 23, 2001 V. SANTHARAM

Institute of Bird Studies & Natural History,

Rishi Valley Education Centre,

Rishi Valley 517 352,

Chittoor district, Andhra Pradesh,

India.

References

Ali, S. & S.D. Ripley (1983): Compact Handbook of the Ghorpade, K.D. (1973): Preliminary notes on the

Birds of India and Pakistan together with those of Ornithology of Sandur, Karnataka. ./. Bombay nat.

Bangladesh, Nepal, Bhutan and Sri Lanka. Oxford Hist. Soc. 70(3): 499-53 1 .

University Press, Delhi. Pittie, A. (2000): Birding Notes. Pitta No. 112: Pp. 7.

32. CRIMSON SUN BIRD AETHOPYGA SIPARAJA SEHERIAE IN MADHYA PRADESH

The crimson sunbird Aethopyga siparaja

appears to have a patchy distribution in the Indian

subcontinent. The subspecies seheriae , according

to the handbook (Ali and Ripley 1987) and the

synopsis (Ripley 1982), is a common resident

from Kangra east to Sikkim (Himalayan foothills,

subject to vertical movements) and thence south

to the plains of eastern Bihar, north Bengal, Uttar

Pradesh, hills of northeastern Peninsula and

Orissa. The southernmost record for this

subspecies is Balaghat in Madhya Pradesh.

The record of Aethopyga siparaja seheriae

for Madhya Pradesh (Balaghat) is based on a

single male specimen shot by D’Abreu at

Laugher Ghat (c. 589 m) in the Balaghat district

on April 1, 1913 (D’Abreu 1913). Since then,

nobody has reported this bird from Madhya

Pradesh.

I have been staying in the buffer zone

(village Mocha) of Kanha National Park (22°

17' N, 80° 30' E) since November 1986 and my

first sighting of this bird was in February 1992.

It was a male (easily identifiable), hopping busily

about the fire bush Woodfordia fruticosa, which

was in full bloom. I was quite surprised to see

this bird here, as there were no previous records

of the species from around Kanha. Within a few

days of my first sighting the male, I saw a female

of this species which was not very difficult to

distinguish from the female purple sunbird

Nectaririia asiatica , that happens to be the only

other resident sunbird species.

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

147

MISCELLANEOUS NOTES

Though Laugher Ghat in Balaghat district

is not very far from where I saw the crimson

sunbird pair, I thought this was another case of

strayed birds as there were no subsequent records

from Madhya Pradesh after 1913.1 observed this

pair for a month or so in the early morning hours,

mainly around Woodfordia fruticosa shrubs, the

flowers of which were heavily laden with nectar.

The Woodfordia shrubs stopped flowering by

March end, and the sunbirds too disappeared.

To my surprise, a male crimson sunbird returned

to my garden in February 1993, which coincided

with the flowering of Woodfordia fruticosa. The

bird disappeared in March. Since then, a male

crimson sunbird has been visiting my garden

almost every year, arriving in February and

leaving around the last week of March or April.

In 1995, a male arrived in the last week of

January and stayed on till the end of March. I

have never seen a female of this bird since

February 1992. I am not sure whether the male,

which has been arriving here is the same

individual or not. In February 1999, once again

a single male arrived and was going about its

business as usual. What surprised me was that

the bird did not return after the Woodfordia

fruticosa stopped flowering. The bird was seen

all through the hot summer months the last

sighting being on June 4. The bird was not seen

throughout the day, but was seen every evening

returning to roost on a small mango tree near

my house.

The regular sighting of this bird around

Kanha (in winter months) definitely proves .that

the bird winters here and probably stays till

mid-summer before returning to the foothills of

the Himalaya where it breeds. Another point is

that perhaps it visits more areas in eastern

Madhya Pradesh and the range of the bird

definitely includes parts of the state other than

the one single spot as recorded by D’Abreu in

1913.

October 16, 2000 E.P. ERIC D’ CUN H A

Kanha National Park,

Village Mocha,

P.O. Kisli 481 768,

Dist. Mandla, Madhya Pradesh, India.

References

Ali, S. & S.D. Ripley (1987): Compact Handbook of the

Birds of India and Pakistan together with those of

Bangladesh, Nepal, Bhutan and Sri Lanka. 2nd edn,

Oxford University Press, Delhi, pp.737 + 1 04 plates.

D’Abreu, E.A. (1913): The occurrence of the Himalayan

Yellowbacked Sunbird, Aethopyga seheriae in the

Central Provinces and extension of its habitat.

J. Bombay nat. Hist. Soc. 22(2)\ 392-393.

Ripley, S.D. (1982): A Synopsis of the Birds of India and

Pakistan together with those in Bangladesh, Nepal.

Bhutan and Sri Lanka. Bombay Natural History

Society, Bombay.

33. INCUBATION PERIOD OF CRIMSON SUNBIRD AETHOPYGA SIPARAJA

The crimson sunbird Aethopyga siparaja

is a resident and fairly common bird in Konkan,

Maharashtra. We see this brilliantly coloured

sunbird in its restless flight in gardens, groves

and dense forest regularly. According to Ali and

Ripley (1999), the nesting period of this bird is

April to July, but we have seen a female of this

species building a nest on August 12, 2000 in

Chiplun, Ratnagiri district, Maharashtra. The

nest was built just outside a house wall in an

urban area.

The pendant nest was entangled to a coir

string, which was tied to a rafter of the house,

96 cm above the ground. This pear-shaped nest

was 13 cm long, 7.5 cm wide and with an 8 cm

awning over the entrance. The entrance was

3.5 cm in diameter.

We observed the nest a couple of times

every day. On August 21 at 0700 hrs, there was

only one egg in the nest. Two more eggs were

added to the nest in the next two days. The female

started incubation only after the third egg was

148

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

MISCELLANEOUS NOTES

laid. The male was never seen incubating the

eggs.

The first egg hatched on September 6, 2000

at 0700 hrs, while the second hatched at

1400 hrs on the same day. The third egg hatched

two days later in the morning. The incubation

period for each egg was 16, 15 and 16 days

respectively. Both the parents fed the chicks, but

most of the work was done by the female, while

the male took very little part in it.

All three chicks fledged on September 21 ,

2000 at 0800 hrs.

March 23, 2001 VISHWAS KATDARE

ROHAN LOVALEKAR

AMEYA MQDAK

Sahyadri Nisarg Mitra

Near Laxminarayan Temple,

Chiplun 415 605, Ratnagiri district,

Maharashtra, India.

Reference

Ali, S. & S. Dillon Ripley (1999): Handbook of the Birds of India and Pakistan together with those of Bangladesh,

Nepal, Sikkim, Bhutan and Sri Lanka. Revised 2nd edn. Vol 10. Oxford University Press, Delhi.

34. THE SPANISH SPARROW PASSER HISPANIOLENSIS FOUND NESTING

IN HANUMANGARH DISTRICT, RAJASTHAN

On March 26, 1999, we were at Badopal

Lake, Hanumangarh district, Rajasthan watching

migratory birds on passage. Large mixed flocks

of Spanish sparrow Passer hispaniolensis and

house sparrow Passer domesticus were

conspicuous in the area, mainly around the ripe

wheat fields and Acacia tortilis plantations along

the road. The flocks of Spanish sparrow were

exceptionally vocal, calling incessantly and

seeming very restless. The whole area was alive

with their noisy and restless behaviour.

After watching the birds on the lake in the

morning, we retired to the shade of the plantation

for lunch, where the Spanish sparrows were

calling feverishly. While watching the birds, we

noticed a few nests on Acacia tortilis trees. A

few males in breeding plumage were loudly

calling “ che die che ....” while displaying with

their tails cocked up and moving actively around

the nests. Two males were noticed mounting

females.

We counted 22 nests on six Acacia tortilis

trees. These nests were about 6-9 m above the

ground, made up of fresh straw collected from

the nearby wheat fields. The nests were c. 25-30

cm in diameter, matching the dimensions given

by Gavrilov (1963), and almost spherical in shape.

Fresh straw was used on the outer surface of the

nests. Most of the nests were on the biggest Acacia

tortilis tree, located on its terminal branches.

On a second visit to the site on April 18,

1999, ten more nests were found. Two males were

displaying with their tails cocked up, and six

males were observed carrying some downy

material (probably cotton from the nearby fields)

and entering the nests.

The Spanish sparrow is a winter visitor to

the northwestern part of the Subcontinent (Ali

and Ripley 1987; Roberts 1992). It breeds

extralimitally in Central Asia westwards to the

Caucasus and eastwards throughout Kazakhstan.

The birds start the return flight from their winter

quarters to the native areas about the beginning

of March. The establishment of colonies in

breeding areas usually takes place a few days after

the beginning of mass arrivals, during the middle

or end of May. The birds usually nest in very

large colonies (Gavrilov 1963). In contrast, the

nesting colony at Badopal was very small and

established as early as March.

The sparrow nests almost exclusively in

cultivated areas, living in the proximity of man.

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149

MISCELLANEOUS NOTES

The availability of grain crops is one of the

necessary conditions for these birds, and they

seldom nest at any distance away from

cultivation (Gavrilov 1963). Development of

extensive plantations and agriculture during the

last forty years or so in northern Rajasthan seems

to have provided the Spanish sparrow with

excellent living conditions conducive to

establishing the first nesting colony in the Indian

subcontinent.

May 30, 2001 MANOJ KULSHRESHTHA

B-33, Sethi Colony,

Jaipur 302 004, Rajasthan,

India.

HARKIRAT SINGH SANGHA

B-27, Gautam Marg,

Hanuman Nagar, Jaipur 302 021,

Rajasthan, India.

Note from the Referee:

The spread of grain cultivation and tree

plantation in Rajasthan has nothing to do with

this unusual nesting. Such favourable conditions

have existed for nearly half a century over most

of the Spanish sparrow’s migration route. What

is significant is the fact that a species known to

be highly gregarious and colonial in nesting, has

not done this before. Similar examples of greater

flamingos and rosy starlings have been recorded

from time to time, usually of abortive attempts

by small colonies to nest in unusual locations. I

was fortunate during a holiday trip to Kazakhstan

two summers ago, to sit under a thorn tree

roadside plantation in an otherwise treeless

Refer

Ali, S. & S.D. Ripley (1987): Compact Handbook of the

Birds of the India and Pakistan together with those of

Bangladesh, Nepal, Bhutan and Sri Lanka. 2nd edn.

Oxford University Press, Delhi, 737 pp. + 104 plates.

Gavrilov, E.I. (1963): The Biology of the Eastern Spanish

steppe grassland region, where above my head

about 500 pairs of Spanish sparrows were

nesting, so I can easily visualise what the authors

saw in Hanumangarh district. Dissection of rosy

starlings on passage in April/May has shown that

their gonads are fully developed for breeding,

and early writers like Ticehurst often speculated

why they did not breed within the confines of

India. Like the Spanish sparrow, small rosy

starling colonies are often opportunistic, e.g. they

have nested in the crevices between a stack of

logs, and don’t show great site fidelity for

choosing their nesting site. The main motivation

seems to be synchronous breeding, as is the case

with the Spanish sparrow also. When the colony

decides to leave, late fledglings are often left to

starve in their nests! Huge colonies of over 1 ,000

birds do exhibit site fidelity, for obvious logistical

reasons, but the central Asian breeding

population of Spanish sparrows does have small

offshoot breeding colonies, despite what Gavrilov

wrote in describing huge breeding colonies in

the Journal. The one 1 saw in Kazhakhstan was

alongside a huge field bearing a crop of Safflower

( Carthamus tinctorius ), which is like a thorny

thistle and then still in flower, so not suitable

food for a sparrow. Otherwise, that colony had

to depend entirely on small grasshoppers and

wild grass seeds for feeding their nestlings. There

were no cereal grain crops within miles and

miles. This is why I feel it does not add to our

knowledge when the authors speculate that

favourable habitat was the reason for breeding

in Rajasthan. The observation teaches us more

about the needs of colonial and synchronous

nesting species — T.J. Roberts.

:nces

Sparrow, Passer hispaniolensis transcaspicus, in

Kazakhstan. J. Bombay nat. Hist. Soc. 60(2): 301-

317.

Roberts, T.J. (1992); The Birds of Pakistan. Vol. 2. Oxford

University Press, Karachi.

150

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MISCELLANEOUS NOTES

35. TERMITE REMOVAL FROM NEST MATERIAL AND REPAIR OF DAMAGED NEST

BY WHITE-RUMPED MUNIA LONCHURA STRIATA

This refers to the note “Termite attack on Piper nigrum and its supporting tree Erythrina

nest material leading to desertion of eggs by

birds” by K.V. Srinivas and S. Subramanya

(JBNHS 97(1): 145).

I have seen in Mangalore (during 1996-

1999), usually old nests of spotted dove

( Streptopelia chinensis), jungle babbler {Turdoides

striatus ) and white-rumped munia ( Lonchura

striata ) being attacked partially or fully by termites

after the birds have left the nest. The nests were

built on small trees at a height of c. 1-3 m. During

the late- and post-monsoon period, when the tree

bark and nest materials dry up, termites from the

ground reach the nest via the tree bark, which they

cover with mud while tunnelling.

1 also saw, on several occasions, live nests

of white-rumped munia (built in thick bushes of

36. BIRDS OF GOA — SOME

Further to "Birds of Goa’ by Heinz Lainer

(JBNHS 96(2): 203-220; 96(3): 405-423), I had

conducted a brief survey of the birds at three

wildlife sanctuaries (Cotigao, Bondla and

Bhagwan Mahaveer) between November 29 and

December 9, 1 995, as part of a survey of the great

black woodpecker. These comments are based

on the observations made then.

Besra sparrowhawk Accipiter virgatus

2 seen in Cotigao Wildlife Sanctuary (CWS);

1 seen in eucalyptus plantation, attacking smaller

birds. All the earlier reports of this species are from

Bhagwan Mahaveer Wildlife Sanctuary (BMWS).

Marsh sandpiper Tringa stagnatilis

1 seen at BMWS (Tambdi Surla area) on

December 7. Reported as winter visitor in small

numbers in the coastal belt.

Nilgiri wood-pigeon Columba elphinstonii

2 birds at CWS on November 30.

Considered a scarce, erratic visitor.

sp.), partially attacked by termites. Interestingly,

the white-rumped munia feasted on the termites,

a protein rich diet, in the nest material and nearby

regions. Moreover, they repaired the damaged

nest (as termites continue to attack the nest

during night time and may damage part of the

nest by early morning). The birds brought fresh

nest material and successfully raised their

offspring.

Oct. 16, 2000 R. SHYAMA PRASAD RAO

Centre for Ecological Sciences,

Indian Institute of Science,

Bangalore 560 012, Karnataka, India.

Email:[email protected]

sprasad 1 0 1 @hotmai 1 .com

SUPPLEMENTARY NOTES

Brainfever bird Hierococcyx varius

1 seen at CWS and 1 at Bondla Wildlife

Sanctuary (BWS). Said to be usually present from

mid-March to early November.

Drongo cuckoo Surniculus lugubris

1 seen in a mixed hunting party in CWS.

Earlier recorded as an uncommon monsoon

visitor — from late May to early October.

Small green-billed malkoha

Pit aenicophaeus viridirostris

A pair and a single bird seen at CWS. Said

to avoid the Western Ghats strip.

Eurasian scops-owl Otus scops

4-5 birds heard calling at BWS. This bird

has not been listed by Lainer (op.cit.).

Long-tailed nightjar Caprimulgus macrurus

1 heard at BWS. No recent records after

the late 1970s.

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

151

MISCELLANEOUS NOTES

Speckled piculet Picumnus innominatus

1 seen in a mixed flock in BWS and a pair

noticed (also in a mixed flock) at BMWS.

According to Lainer, this bird has been noticed

just once earlier.

Indian pitta Pitta brachyura

2-3 heard at CWS. Said to be found from

end April to early September and a few passage

migrants seen between early January and mid-

March.

Spangled drongo Dicrurus hottentottus

10-15 birds in flight at CWS; a pair at

BMWS. Said to be a scarce winter visitor in small

numbers.

Ashy woodswallow Artamus fuscus

5-6 in CWS; 1-2 heard at BWS and 4-5 in

flight at BMWS. Said to be absent from large tracts

during the dry season, being a monsoon visitor.

Scarlet minivet Pericrocotus flammeus

A juvenile female seen begging for food at

BWS.

Indian scimitar-babbler

Pomatorhinus horsfieldii

Calls heard at CWS. Recorded on the crest

of the Sahyadri and, occasionally, down to an

altitude of c. 70 m.

Grey-headed flycatcher

Culicicapa ceylonensis

1 seen at BWS. Has been recorded only

twice earlier.

Acknowledgements

I thank WCS, New York, who funded my

study on the great black woodpecker and the

Forest Department, Goa, for granting permission

and providing facilities.

November 25, 2000 V. SANTHARAM,

Institute of Bird Studies &

Natural History,

Rishi Valley Education Centre,

Rishi Valley 517 352,

Chittoor district, Andhra Pradesh,

India.

37. BIRDS OF TABO: A LESSER KNOWN COLD DESERT

IN THE WESTERN HIMALAYA

The cold desert area of Tabo Valley (30°

05' N, 78° 28' E) lies c. 3,500 m above msl in

the Spiti district, Himachal Pradesh State in

northern India. This broad, flat valley is

intersected by high, craggy, vertical cliffs of gray

and brown sand. The higher slopes of the valley

are covered with undulating alpine grasslands,

which rise to the mountains, often up to the

snowline or peaks (5,500-7,000 m). The boulder-

strewn valley is also traversed by the River Spiti

that enters through narrow gorges from the west,

and flows down eastwards. The soil is alkaline

(ph 7.2), poor in nutrients (K, Na, P) and organic

matter (0.08%), and is silty, clayey as well as

loamy (Singh and Gupta 1990).

The climate is cold and dry with heavy

snowfall (c. 80 cm) in winter (December-March).

Rainfall occurs only during September and

October, and is scanty (10-15 mm). The

temperature fluctuates from a maximum of

32.5 °C in August to a minimum of -32 °C in

January (Kapoor and Bhagat 1990). High

velocity winds blow throughout the year.

The vegetation of the area is xerophytic

— dry alpine scrub, associated with soil moisture

especially in the ravines and near water springs.

The dominating species are Hippophae

rhamnoides , Rosa webbiana , Ephedra

gerardiana , Caragana versicolor , Poa prate ns is,

Myricaria prostrata , Chenopodium hybridum.

152

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MISCELLANEOUS NOTES

Cotoneaster falconeri, Lonicera spinosa,

Capparis himalayana , Agropyron repens ,

Eragrostris sp., and Trigonella sp., (Singh and

Gupta 1990; Aswal and Mehrotra 1994).

A few trees of Juniperus macropoda and

J. semicarpifolia grow naturally along the Spiti

river and other watercourses. Plantations of

Popidus alba , Salix fragilis , S. alba and Robinia

pseudoacacia are also present along the roadside

in Tabo, raised under the ‘Desert Development

Programme’ in 1985 (Negi 1985). The principal

crops, cultivated only in flat areas having

irrigation, are barley, millet, buckwheat

(. Fagopyruin sp.), potatoes and wheat.

As there was no record of birds from Tabo

Valley, a survey was carried out from December

5-12, 1992. A total of 10 sites were selected from

the entire Tabo valley (5-6 sq. km). Observations

were recorded daily from 0900 to 1 600 hrs. Data

on the overall species richness, abundance, food,

feeding habits and behaviour of birds in the area

were collected. Winter was delayed that year, and

as a consequence there was no snowfall in Tabo,

making it favourable for birds (maximum day

temperature 14 °C; minimum night temperature

-18 °C).

Only 13 species of birds belonging to

9 families were observed in the study area.

Golden eagle Aquila chrysaetos

(Accipitridae): Two adults and an immature

bird observed flying and calling near high cliffs

on the mountainous slopes north of Tabo

village. In the afternoon, an adult was noted

successfully preying on a woolly hare Lepus

oiostolus.

Bearded vulture Gypaetus barbatus

(Accipitridae): Two adults flying low over River

Spiti, between rocky gorges. 3 km west of Tabo

village, along the road to Kaza.

Chukor Alectoris chukar (Phasianidae):

A small flock of five feeding on the ground under

Salix alba plantation at Tabo. All birds hiding

in a pile of boulders after sensing a red fox Vulpes

vulpes montana. They came out of hiding after

12 min when the threat was over. Second sighting

of 7 birds feeding in fallow terraced fields in the

company of more than 17 blue sheep Pseudois

nayaur.

Hill pigeon Columba rupestris

(Columbidae): A small flock of nine feeding on

wheat grains dropped on the road between Lari

and Tabo, 3 km west of Tabo village.

Yellow-billed chough Pyrrhocorax

graculus (Corvidae): Five individuals once

observed flying and circling over Tabo village.

Red-billed chough Pyrrhocorax

pyrrhocorax (Corvidae): Observed daily, three

birds seen perching on rocky slopes near Tabo

village.

Dark-throated thrush Turdus ruficollis

(Turdinae): Eight birds observed feeding on

insects on the ground under a Robinia

pseudoacacia plantation along the road.

Guldenstadt’s redstart Phoenicurus

erythrogaster (Muscicapidae): A unique

sighting of a large, congregation (>25

individuals), all males. These birds inhabited a

flat rocky area with stones and boulders beside

a stream, with many fruiting bushes of

Hippophae rhamnoides , 4 km west of Tabo

village along the road to Kaza. Each individual

feeding on the fruit and guarding its bush against

intruders by flying over it and displaying its

territory. One bird also observed on a Salix tree

at Tabo village.

Grey-backed shrike Lanius tephronotus

(Laniidae): A single shrike observed perching

on Salix alba tree in an open, boulder-strewn,

plantation. Seen searching for overwinter-

ing lepidopterous immatures in crevices of

boulders.

Common great rosefinch Carpodacus

rubicilla (Fringillidae): Small parties observed

daily, feeding on seeds (pods) of Robinia

pseudoacacia in a roadside plantation. A total

of 28 birds seen.

Hodgson’s mountain-finch Leucosticte

nemoricola (Fringillidae): Common (>30 birds)

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

153

MISCELLANEOUS NOTES

around Tabo village and on rocky slopes.

Rufous-breasted accentor Prunella

strophiata (Pruneilidae): Small flocks, common

in fields around Tabo village. Altogether 5 birds

seen.

Tibetan snowfinch Montifringilla adamsi

(Passerinae): Two birds once seen feeding on

the ground in the vicinity of Tabo village.

The birds were identified using Alt and

Ripley <1983, 1989) and Porter et al. (1981).

Refer

Ali, S. & S.D. Ripley (1983): A Pictorial Guide to the

Birds of the Indian subcontinent. Bombay Natural

History Society, Bombay.

Ali, S. & S.D. Ripley (1989): Compact Handbook of the

Birds of India and Pakistan, together with those of

Bangladesh, Nepal, Bhutan and Sri Lanka. 2nd edn,

Oxford University Press, Delhi, pp. 737 + 104

plates.

Aswal, B.S. & P N. Mehrotra (1994): Flora of Lahaul-

Spiti. Bishan Singh and Mahendra Pal Singh Publ.

Dehra Dun.

Kapoor, K.S. & S. Bhagat (1990): Resource potentials of

March 23, 2001 ARUN P. SINGH

Entomology Department,

Forest Research Institute,

Dehra Dun, Uttaranchal, India.

Email: [email protected]

Permanent Address: c/o Col. R.S. Verma

25-D, New Cantt. Rd,

Hathi Barkala, Dehra Dun,

Uttaranchal 248 001, India.

EN CE S

Spiti: Cold mountain desert of Himachal Pradesh.

Annals of Arid Zone 29: 243 -250.

Negi, J.P. (1985): Developmental perspectives in the

Himalayan mountain desert of Spiti (Himachal

Pradesh). Greening of Mountain Deserts. 72nd

Session of INS A, Lucknow.

Porter, R.F., S. Christensen, I. Willis & B.P. Neilsen

(1981): Flight identification of European raptors.

3rd edn. Poyser, Carlton.

Singh, R.P. & M.K. Gupta ( 1 990): Soil and vegetation study

of Lahaul -Spiti cold desert of western Himalayas.

Indian Forester 116: 785-790.

38. OCCURRENCE OF TRIOPS GRANARIUS (LUCAS),

CRUSTACEA: NOTOSTRACA, FROM MADURAI, TAMIL NADU

(With one text-figure)

Tadpole shrimps, considered as ‘living

fossils’, are widely distributed in all continents

except Antarctica (Whitehead 1990). The genus

Triops has been reported from isolated localities

in the Indian subcontinent (Packard 1871; Sars

1901; Kemp 1911; Walton 1911; Gurney 1925;

Chacko 1950; Tiwari 1951, 1952; Longhurst

1955; Shanbhag and Inamdar 1968; Sanjeeva Raj

1971; Ghate and Shetty 1997). It is known for

its intraspecific morphological variations that is,

within the same species in different localities,

hence it has been described under different

specific names, adding to the taxonomic

confusion within the genus.

This study reports the occurrence of Triops

from Madurai, Tamil Nadu. About 20 mature

live tadpole shrimps were collected near

Thirumangalam (near Latibos India), Madurai

(9° 58' N, 78° 10' E) during our survey on fairy

shrimps in October 1996. Sizes of both male and

female range from 1.8 to 3.9 cm. Species of this

group have an elongate body, oval carapace

covering the head, thorax and a variable portion

of the abdomen (Fig. la). Shape varies from

round to triangular, with rounded anterior

margin between eyes, whereas the posterior

margin is straight and slightly emarginated.

Lateral edges of carapace bear small spines and

edge of rear notch has short prominent spines.

Sulcus is triangular with rounded emargination

and wide base. Head bears an eye on the dorsal

surface and reduced second antennae. The dorsal

154

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MISCELLANEOUS NOTES

a

5 mm

b-

4 mm

Fig. la-b: a. Diagrammatic representation of the notostracan Triops granarius (Lucas):

b. Higher magnification of the dorsal eye

D - dorsal shield, abs - abdominal segment, t - telson, 2A - second antennae,

CE - compound eye, e - dorsal eye, f - furca, ne - naupliar eye, N - neck organ, phy - phyllopods, s - sulcus

eye possesses a reduced naupliar eye and a neck

organ between the kidney-shaped compound eyes

(Fig. lb).

Abdomen has 35 movable somites, about

15 of which are exposed beyond the carapace.

Nine apodal segments possess prominent

chitinised brown dorsal and ventral spines.

Telson is broader and consists of 4 dorso-medial

spines, 3 strong setal spines and small denticles,

6 posterior marginal spines, 4-5 lateral spines

and 5 furcal spines. Furca is slender and longer

than carapace, with serrated spines. Eleven pairs

of thoracic appendages, namely phyllopods, one

on each of the eleven body segments are used for

locomotion. First pair consists of two segments

separated by a joint chitinous cuticle, which

allows bending of one segment upon the other;

they have a sensory function and are used for

capture of prey. There are 50 pairs of legs,

gradually decreasing in size from the genital

apertures; they are known as “abdominal

appendages” and are used both for feeding and

locomotion. Reproductive mode varies on a

geographical basis, northern forms being

hermaphroditic while southern populations are

bisexual. In general, males are rare in die genus

Triops. Gurney (1925) reported males

outnumbering females in his collection (=Apus

asiaticus ) and Tiwari (1951) also agrees with

Gurney’s findings.

Notostracans are detritus feeders and

predators. In the present study, it was collected

along with anostracans and conchostracans.

T. orientalis appears to be omnivorous and it has

been observed to feed on bacteria, protozoa,

Daphnia , copepods, small oiigochaetes and also

on Strepiocephalus and Leptestheriied forms

(Shanbhag and Inamdar 1968). Triops exhibits

cannibalism, and even a small individual can

easily eat a larger one. Triops is recorded as a

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

155

MISCELLANEOUS NOTES

pest of rice cultivation in different parts of

the world such as Kashmir (Kemp 1911, Walton

1911), Spain (Font de Mora 1923), California

(Rosenberg 1946) and Japan (Takahashi 1977).

Fryer (1987) reported natives of the Federal

District, Mexico using tadpole shrimps as

food.

The genus is well known for its

discontinuous distribution. Linder (1952) and

Longhurst (1955) recognized only about 11

species, of which seven have wide geographic

distribution. This classification has long been

accepted 'even attaining the status of dogma’

(Sassaman et al. 1997).

According to our study and other published

records, only two species of Triops ,

Refer

Chacko, P.I. ( 1 950): Occurrence of the fairy shrimp Apus

in a temple tank in Tirunelveli District, Madras.

J. Bombay nat. Hist. Soc. 49: 571.

Font de Mora. R. (1923): Un destructor de planteles de

arroz. Bol. Soc. Esp. Hist. Nat. Madrid. 23: 313.

Fryer, G. (1987): A new classification of the branchiopod

Crustacea. Zool. J. Linn. Soc. 91: 357-383.

Ghate, H. V. & Nagaraj Shetty ( 1 997): Record of Triops

(Crustacea: Branchiopoda: Notostraca) from Pune,

Maharashtra. J. Bombay nat. Hist. Soc. 94: 588-

589.

Gurney. R. (1925): Some Asiatic species of Apus. Rec.

Indian Mus. 27: 439-442.

Kemp. S. ( 1 9 1 1 ): Notes on the occurrence of Apus in Eastern

Asia. 11. Notes on Major Walton's specimens and

on others from Kashmir with a list of previous

records fi om Eastern Asia. Rcc. Indian Mus. 6: 353-

357.

Linder. F. (1952): Contributions to the morphology and

taxonomy of the Branchiopoda Notostraca. with

special reference to the North American Species.

Proc. U.S. Nat. Mus. 102: 1-69.

Longhurst, A.R. ( 1 955): A review of the Notostraca. Bull.

Brit. Mus. Nat. Hist. (Zool.) 3: 3-57.

Packard, A.S. (1871): Preliminary notice of new North

American phyllopoda. Am. J. Sci. Arts. Set: 3(2):

108-113.

Rosenberg, L.E. (1946): Fairy shrimps in California.

Science 104: 111.

Sanjeeva Raj. P.J. (1971): Triops granarius (Lucas)

T. cancrifonnis (Bose) and T. granarius (Lucas)

are known to occur in the Indian subcontinent.

The former was reported in northern localities

and the latter in the rest of India, particularly in

the southern parts.

The authors are deeply indebted to the late

Mr. Anthony Basil, Technical Officer, Madurai

Kamaraj University, Madurai for accompanying

them during the collections.

July 18, 2001 C.S. VELU

*N. MUNUSWAMY

Department of Zoology,

University of Madras, Guindy Campus,

Chennai 600 025, Tamil Nadu, India.

*Email: [email protected]

E N C E S

(Crustacea: Branchipoda) from Tamil Nadu and a

review of the species from India. J. Bombay nat.

H ist. Soc. 68: 161-168.

Sars, G.O. (1901): Or. the crustacean fauna of Central

Asia. Ann. Mus. St. Petersburg 6: 130-164.

Sassaman, C.. M.A. Simovich & M. Fugate (1997):

Reproductive isolation and genetic differentiation

in North American species of Triops (Crustacea:

Branchiopoda: Notostraca). Hydrobiologia 359:

125-147.

Shanbhag, S.V. & N.B. Inamdar (1968): On the

occurrence of Triops mavliensis (Tivvari),

Notostraca (Crustacea) in the Okhamandal region

of Saurashtra (India). J. Bombay nat. Hist. Soc.

65: 408-417.

Takahashi, F. (1977): Triops spp. (Notostraca: Triopsidae)

for the biological control agents of weeds in rice

paddies in Japan. Entomophaga 22: 35 1 -357.

Tivvari, K.K. (1951): Indian species of the genus Apus

(Crustacea: Branchiopoda) with description of two

new species. Rec. Indian Mus. XLIX: 1 97-205.

Tivvari, K.K. ( 1 952): On sex ratio and variability of Apodal

segments in Apus (Phyllopoda, Crustacea).

J. Bombay nat. Hist. Soc. 52: 641-644.

Walton, S.J. (1911): Notes on the occurrence of Apus in

Eastern Asia. 1. On the occurrence of Apus Latreilie,

in the United Provinces of India. Rec. Indian Mus.

6: 351-352.

Whitehead. P. ( 1 990): Systematics: an endangered species.

Svst. Zool. 39: 179-184.

156

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MISCELLANEOUS NOTES

39. OCCURRENCE OF RARE JUMPING SPIDER HARMOCHIR US BRACHIATUS

(THORELL) (FAMILY: SALTICIDAE) IN THE BANANA AGRO-ECOSYSTEM OF

VADODARA, GUJARAT

While studying the spider diversity of the

banana agro-ecosystem of Vadodara, we came

across a rare jumping spider Harmochirns

brachiatus (Thorell). It was first reported by

Tikader (1976) from Pune, Maharashtra, India.

There is no report of its occurrence thereafter. It

is an addition to the spider fauna of Gujarat.

Adult male is c. 2.00 mm in total length,

Carapace 1.1 mm long and 0.8 mm wide. Abdo-

men 0.8 mm long and 0.7 mm wide. The tibia

and femur of the first pair of legs swollen and

first leg always kept up in a defensive position.

Because of the unique structure of the first pair

of legs and their peculiar way of movement, this

spider can be easily distinguished from other

salticids. A closer view of the 1st pair of legs,

shows a row of fringed hairs on the ventral and

dorsal sides of the tibia, 3 and 2 pairs of ventral

spines present on tibia and metatarsi respectively.

They move in the banana fields, on the

pseudostem of the banana plant, and among the

dry and decaying leaves of banana. Uncommonly

seen in the pre-monsoon season (January-May).

April 30, 2001 SILIWAL MANJU

* DOLLY KUMAR

Division of Entomology,

Department of Zoology,

Faculty of Science,

M.S. University of Baroda,

Vadodara 390 002, Gujarat, India.

*Email: [email protected]

Reference

Tikader, B.K. (1976): Redescription of a jumping spider Harmochirns brachiatus (Thorell) with a new record from

India. J. Bombay nat. Hist. Soc. 73(2): 410-4 1 1 .

40. DESCRIPTION OF FEMALE AMYCIAEA FORTICEPS (CAMBRIDGE),

ARANEAE: THOMISIDAE, WITH A REDESCRIPTION OF ITS MALE

FROM KERALA, INDIA

(With one text-figure)

Amyciaea forticeps (Cambridge) is an ant-

mimicking species of crab spider reported from

India, Holland, Africa, Burma (=Myanmar) and

Malaysia. Its resemblance to Oecophylla

smaragdina (Fabr.) was studied by Mathew

(1954). Tikader (1963) gave a description of this

species in fauna of india based on a single male

specimen collected from Pune. However,

taxonomic literature regarding A. forticeps

remains largely incomplete due to the absence

of the description of a female. During our study

of spiders of Ernakulam district, Kerala we came

across several specimens of A. forticeps. On the

basis of these specimens, a description and

illustration of A. forticeps is given below.

Collection and preservation of the spider

samples were done following Tikader. The

material was studied using a Stereozoom

binocular microscope; model Leica MS 5. All

measurements are in millimetres, made with an

eyepiece graticule.

Amyciaea forticeps (Cambridge)

(Fig. la-h)

1873 Amycle forticeps Cambridge, Proc.

loot. Soc., Lond. 1873: 122

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MISCELLANEOUS NOTES

Fig. 1: Amyciaea forticeps (Cambridge); a. Dorsal view of the female, b. Lateral view of the female;

e. Front view of the face, d. Sternum with Labium and Maxillae, e. Epigyne, f. Internal genitalia,

g. Palp - Ventral view. h. Palp - lateral view.

158

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MISCELLANEOUS NOTES

1885 Amyciaea forticeps: Simon, Bull.

Soc. Zool. Fr. 10: 447.

1 963 Amyciaea forticeps : Tikader, J. Univ.

Poona Sci. & Tech. 24: 52.

1980 Amyciaea forticeps : Tikader, Fauna

of India: Spiders (Araneae: Thomisidae) 1(1): 169.

Female: Measurements (in mm): Total

length 5.5 L, Carapace 2.2 L & 1.3 W; Abdomen

3.2 L & 1.5 W. Legs I: 8.24, II: 7.92, III: 4.33,

IV: 6.87.

Cephalothorax longer than wide, cephalic

region strongly elevated and sloping in front,

anterior margin of cephalothorax straight.

Thoracic area sloping gradually, wider behind

PLE, lateral sides steeply sloped. Cephalothorax

reddish brown or pale orange, with a few body

hairs. Fovea shallow and inconspicuous. Eyes

recurved, in three rows, PLE and ALE encircled

by inner white ring and outer black patch.

Median eyes smaller than laterals, PME smallest,

PME at the middle of two eye rows, ocular

quadrangle wider behind. Eyes black, eye

diameter ALE = PLE - 0.5 mm, AME = 0.25

mm, PLE = 0.1 mm. Clypeus high, twice the

height of AME. Sternum heart shaped, 1.3 times

longer than wide, anterior margin straight,

clothed with minute hairs, pointed posteriorly,

reddish brown or light orange in colour. Labium

reddish brown, longer than wide, maxillae

similar in colour to labium, scopulae present on

the inner margin. Chelicera moderately strong,

similar to clypeus in colour and height, with a

few hairs on the outer margins, seven small teeth

on outer margin and two large teeth on inner

margin. Palp as in Fig. lg-h. Legs long and

slender, tarsus with two claws provided with

minute hairs. Leg formula 1243. Legs resemble

ant legs.

Abdomen longer than wide, anterior

portion narrower, middle portion widest, two

black eye-like spots on the dorsum on the

posterolateral sides. A conspicuous inverted kY’

shaped dark brown marking on the anterior half.

Ventral side pale reddish-brown. A black spot at

the anterior lateral end of the abdomen. Pedicel

long. Abdomen clothed with fine hairs.

Male: Measurements (in mm): Total length

5L, carapace 2L & 1 . 1 W, Abdomen 3L & 1 .5 W.

Cephalothorax longer than wide, reddish

brown, narrowed anteriorly, broadest behind

PLE Cephalic region strongly raised and anterior

margin slightly sloped. Thoracic area sloping

gradually, posterior margin smoothly curved.

Fovea long, shallow and inconspicuous. Clypeus

moderately high, a little less than the separation

of ALE. Ocular area as in female. Sternum

reddish brown, longer than wide with truncate

anterior end and pointed posterior end. Labium

longer than wide, reddish brown. Maxillae

similar to labium in colour. Chelicera moderately

strong. Legs long, slender and ant-like, reddish

brown in colour, tarsus and distal end of the

metatarsus lighter. All segments uniformly

clothed with minute hairs, few large bristles on

femur, tarsus with two claws. Leg formula 1243.

Pedipalp as long as tibia of leg I, MBA one,

cymbium long straight and conical.

Abdomen longer than wide, reddish brown

but lighter than carapace. Anterior end narrower,

broadest behind the middle. Dorsum marked with

an inverted ‘Y’ shaped dark brown marking and

three to four transverse dark brown stripes

Table 1 : Measurements of leg segments (in mm) of $ Amyciaea forticeps (Cambridge)

Leg Coxa Trochanter Femur Patella Tibia Metatarsus Tarsus Totai

I 0.43 0.18 1.98 0.43 2.16 1.98 1.08 8.24

II 0.36 0.29 1.62 0.54 2.09 1.87 1.15 7.92

Hi 0.29 0.11 1.08 0.29 1 08 1.01 0.47 4.33

IV 0.36 0.18 1.98 0.32 1.69 1.62 0.72 6.87

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159

MISCELLANEOUS NOTES

posteriorly. A black eye-like irregular spot on the

anterolateral end near the pedicel. Spinnerets

subequa! in length. Dorsum has two hump-like

prominences on the anterior and posterior end,

with the middle being depressed, resembling an

ant’s abdomen. Ventral side pale reddish brown.

Materials examined: 2$?, location:

Ernakulam 3.xii.2000, Coll: Samson Davis;

2 9 9, location: Cochin 2.xi.2000, Coll: Sunil

Jose, K.; 2dd, location: Paravur, Ernakulam

15.xii.2000, Coll: Sudhikumar, A.V.

Natural History: Collected from leaves of

Mangifera indica in the same habitats as that of

Oecophylla smaragdina (Fabr.)

Distribution: India: Ernakulam (Kerala

State), Pune (Maharashtra); Burma (=Myanmar);

Malaysia; Holland; Africa.

Remarks: According to Tikader, an eye-

like spot is present on the posterolateral end of

the abdomen in male, whereas this is absent in

our specimens. Similarly, in the palp the embolus

is coiled two and half times, whereas it was only

one and half times according to Tikader. The

black irregular spot present on the anterolateral

end of the abdomen in both sexes in our specimen

is also absent in Tikader’s description. Since the

specimens are similar to Tikader’s specimen in

all other characters, we believe this difference

may be a geographical variation or due to an

omission in description.

Acknowledgement

We thank the Principal, Sacred Heart

College Thevara, Rev. Fr. George Koyikara

C.M.I. for providing facilities.

February 7, 2001 K. SUNIL JOSE

SAMSON DAVIS

A.V. SUDHIKUMAR

P.A. SEBASTIAN

Department of Zoology,

Sacred Heart College, Thevara, Kochi,

Kerala 682 013, India.

Reference

M athew, A.R (1954): Observations of habits of two spider mimics of the Red Ant Oecophylla smaragdina (Fabr.).

J. Bombay nat. Llist. Soc. 52: 249-263.

41. RARE SIGHTING OF OGRE-FACED SPIDER D1NOP1S GOALPARAENSIS, ARANEAE:

DINOPIDAE, IN THE BANANA AGRO-ECOSYSTEM OF VADODARA, GUJARAT

( With two text-figures)

Dinopis is commonly known as the ogre-

faced spider. Review of literature shows that

this spider is poorly known from India. There is

a single report of the occurrence of Dinopis

goalparaensis by Tikader and Malhotra (1978)

from Jamduar, District Goalpara, Assam. This

note records the occurrence of Dinopis

goalparaensis from Gujarat for the first time.

To study spider diversity in a banana agro-

ecosystem an extensive survey was carried out

in different banana fields situated in a 20 km

radius of Vadodara city. Spiders were hand picked

and preserved in 70% alcohol. The ogre-faced

spider, attached to dry leaves of a banana plant,

was collected from its web. These spiders have a

flattened carapace, elongated abdomen and long

slender legs. Because of these characters this

spider superficially resembles the juvenile of

Eucta sp (Family: Tetragnathidae), however, it

is easily differentiated from Eucta sp. by the

following characters:

1 . Posterior median eyes larger than the rest,

black in colour, anterior median eyes

smallest and anterior laterals present on

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MISCELLANEOUS NOTES

Posterior

Lateral Eye

Posterior

Median Eye

Fig. I: Front view of face, showing arrangement

of eyes

tubercles which are pointing downwards

(Fig. 1). In Eucta sp. eyes are smaller and

equal in size.

2. Subadult female measured about 4.4 mm

in total length, carapace 1.2 mm long and

1.0 mm wide, abdomen 3.2 mm long and

0.8 mm wide. However, adult female

measures about 13.3 mm in total length

(Tikaderand Malhotra 1978), nearly equal

to that of Eucta sp.

3. Abdomen long but not pointed at the

posterior end (unlike Eucta sp.).

4. Legs long but very delicate as compared

to Eucta sp.

Dinopis goalparaensis rests in its web,

stretching its legs like Eucta sp., anterior legs

extended forward and posterior legs extended

backward.

It is a nocturnal weaver and constructs two

types of webs: Orb web and Actual Prey capture

web (Fig. 2)

Fig. 2: Prey capture web held between the first two

pairs of legs

The Orb web of Dinopis is similar to that

of other orb weavers and it is not basically used

for catching prey but for resting. The prey capture

web is rectangular (like a tennis court net) and

is held between the front legs by the spider, it

consists of sticky silk threads.

Acknowledgement

We thank Dr. B.K. Biswas, Zoological

Survey of India, Kolkata, for confirming the

identity of the species and providing relevant

literature.

July 3, 2001 SILIWAL MANJU

* DOLLY KUMAR

Division of Entomology,

Department of Zoology, Faculty of Science,

M.S. University of Baroda,

Vadodara 390 002, Gujarat, India.

*Email: [email protected]

Reference

Tikader, B.K. & M.S. Malhotra ( 1 978): A new record of rare spider of the Family Dinopidae from India with description

of a new species. Proc. Indian Acad. Sci. 87B(6): 1 57- 1 59.

42. OBSERVATIONS ON BAUHINIA MALABARICA ROXB, LEGUMINOSAE:

CAESALP1NIOIDEAE, SHAPE OF CALYX IS NOT CORRELATED WITH SEXUAL

NATURE OF FLOWERS

Roxburgh while commenting on Bauhinia 1832) said, “This very distinct species is

malabarica Roxb. (in Carey ed., FI. Ind. 2: 321. remarkable for the regularity of its five-parted

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

161

MISCELLANEOUS NOTES

calyx. . .”; de Wit (in Reinwardtia 3(4): 533-534.

1956) stated “...the calyx splits in Malaysian

specimens in the upper part into two lobes, one

consisting of two sepal-tops and the other of

three. In some cases, the five tops become free.”

He felt that the dimorphism might be connected

with the sexual nature of the flowers, but he had

not been able to demonstrate that the shape of

the calyx was correlated with the sex of the

flower. He further stated, “It is just possible that

in India the tops of the sepals become always

free and that this is connected with the flowers

being male, which is confirmed by a few

specimens from India which 1 was able to

examine”. In the course of my study. 1 have

observed that in the Indian specimens, the calyx

is five-lobed in the upper part in the female

flowers too. Thus, the shape of the calyx is

not correlated with the sexual nature of the

flowers.

In this connection, 1 would like to mention

that fully developed male flowers are rarely found

in herbarium specimens (see also de Wit in

Reinwardtia 3(4): 533. 1956) because they

remain attached to the pedicels just for a night

and start falling from the next morning. Thus,

during the flowering period, numerous fresh male

flowers are found scattered under the tree,

particularly in the morning hours.

March 29, 200 1 S. BANDYOPADHYAY

Botanical Survey of India,

P.O. Botanic Garden,

Howrah 711 103, West Bengal, India.

43. REDISCOVERY OF CEROPEGJA EVANSII McCANN,

ASCLEPIADACEAE, FROM MAHARASHTRA

( With one plate and one text-figure)

Ceropegia evansii McCann

(Asclepiadaceae) is an endemic and threatened

plant species. The species is known to occur only

from the hill ranges of the Western Ghats of

Maharashtra i.e. Khandala and the neighbouring

Sakarpathar-Ambavane range of Pune district

(Ansari 1984, Jagtap and Singh 1999).

The species was first described by McCann

from Khandala (1945). Santapau and Irani (1958,

1962) reported the species from the same hill

ranges. The species was collected on July 27, 1964

by B.V. Reddi (93331) from Ambavane and

deposited at the Botanical Survey of India (BSI).

About the occurrence, Santapau (1953)

noted that the species is “one of the commonest of

the Ceropegias in Khandala and is found abundant

on the lower slopes beiow Duke’s Nose.”

The species has disappeared very fast

from its type locality because of anthropogenic

problems and habitat destruction. It has not

been collected again from its type locality and

other areas after 1964. This might be due to

anthropogenic pressures and habitat destruction.

Ahmedullah and Nayar (1968) kept this

plant under the rare and endangered category

because of its localized distribution. In the red data

book, Nayar and Sastry (1987) gave “vulnerable”

status to the species. Almeida and Almeida ( 1 990)

have listed it as a threatened and endemic species.

Singh and Karthikeyan (2000), Mishra and Singh

(2000) have treated the species as critically

endangered. According to the latter, the number

of mature individuals in the wild is below 50. They

have also reported that in 1 997, a few plants were

noted at Amba Ghat (Yadav, pers. comm.). Tetali

et al. (2000) have treated it as vulnerable.

During routine botanical explorations, we

have collected C. evansii from Rajgad, a hill fort,

located in the Velhe taluka of Pune district in

Maharashtra State (Fig. 1) at an altitude of

850 m. The present report is a rediscovery of

C. evansii from a new locality other than

162

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

MISCELLANEOUS NOTES

Tetali, P. et al. : Ceropegia evansii McCann

Plate 1

Fig. 1 : Ceropegia evansii McCann

a. Habit; b. Inflorescence; c. Single flower; d. Vertical section of flower showing corona

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163

MISCELLANEOUS NOTES

Fig. 1 : Distribution map of Ceropegia evansii McC.

the Ambavane hill ranges (14.viL200G Tetali,

s.m).

We found only one individual in the entire

locality, indicating extreme rarity of the species.

The present collection site is 100 km away from,

and southwest of the type locality.

The area where the specimen was collected

is a degraded hill slope. The plant was found

growing at the edge of a subtropical hill forest

among Lantana camara and Carissa congesta

bushes. The surrounding vegetation of the area

is dominated by a gregarious shrub Carvia

callosa. The area also seems to be the grazing

ground for village cattle. Dozens of stray cattle

were found grazing in the habitat. The cattle

eat the entire plant. Cowherds and stray cattle

appear to be a serious threat to the natural

populations.

Information with regard to ranges in

descriptions of certain morphological characters

observed in comparison to the earlier description

are as given below.

Twiners up to 3 m long (up to 2.7 m);

petioles up to 2.5 cm long (up to 1 cm long);

cymes consisting of few to many flowers, up to

13 (few flowered); peduncles up to 7.5 cm long

(long (sic))\ pedicels up to 1.3 cm long (up to

1 cm long); corolla 4.5 cm long (4 cm long).

Various parts of the plant are shown in Plate 1 to

facilitate identification.

The voucher specimens are deposited at the

Herbarium, Botanical Survey of India, Western

Circle, Pune (BSI), and Naoroji Godrej Centre

for Plant Research, Lawkim Ltd. Campus,

Shindewadi.

Acknowledgements

We are grateful to Mr. V.M. Crishna,

Director and Mr. D.G. Oak, Naoroji Godrej

Centre for Plant Research and also to the

Director, Botanical Survey of India and Deputy

Director, Botanical Survey of India, Western

Circle, Pune for facilities and support.

JOURNAL. BOMBAY NATURAL HISTORY SOCIETY, 100(1), APR. 2003

165

MISCELLANEOUS NOTES

March 29, 2001 P. TETALI

SUJATA TETALI

Naoroji Godrej Centre for Plant Research,

Gate No. 431, “Lawkim Ltd.” Campus.

Shindewadi, Shirwal Post 412 801,

Satara district, Maharashtra, India.

Refer

Ahmedullah, M. & M.R Nayar(1986): Endemic Plants

of the Indian Region., BS1, Calcutta. Pp. 120.

Almeida, S.M. & M.R. Almeida (1990): Threatened

endemic plants of Maharashtra. In: Conservation in

Developing Countries: Problems and Prospects.

Proceedings of Centenary Seminar of the Bombay

Natural History Society (Ed.: Daniel, J.C. and J.S.

Serrao). Pp. 544-551.

Ansarl M.Y. ( 1 984): Asclepiadaceae: Genus - Ceropegia.

Ease. FI. Ind. 16: 1-34 spl. 4, figs. 18. Botanical

Survey of India, Howrah.

Jagtap, A.P. & N.P. Singh (1999): Asclepiadaceae &

Periplocaceae. Fasc. FI. India 24: 1-332, photo pi.

24, figs. 53. Botanical Survey of India, Calcutta.

McCann, C. (1945): New species of Ceropegia and the

synonymy of the Indian species. J. Bombay, nat. Hist.

Soc. 45: 209-211 .

Mishra, D.K. & N.P. Singh (2000): Status survey of

Endemic species of Ceropegia L. in Maharashtra.

In: Fligher plants of Indian subcontinent (Ed.: Gupta,

B.K). Indian J. For. series 1 1: 17-32. Bishen Singh

• > ..

P LAKSHMINARASIMHAN

P.V. PRASANNA

B.G. KULKARNI

Botanical Survey of India,

Western Circle, 7 Koregaon Road,

Pune 411 001, Maharashtra, India.

ENCES

Mahendra Pal Singh, Dehra Dun.

Nayar, M.P. & A.R.K. Sastry (Eds.) (1987): Red Data

Book of Indian plants. Vol. 1, pp. 54-55. Botanical

Survey of India, Calcutta.

Santapau, H. (1953): The FloraofKhandalaonthe Western

Ghats of India. Rec. Bot. Snrv. India pp. xxvii+396

map. figs. 3. (3rcl Rev. Edn. 1967).

Santapau, FI. & N.A. Irani (1958): The genus Ceropegia

in Bombay. Bull. Bot. Soc. Beng. (Agharkar Mem.

Vol.) 12: 6-17, figs. 4.

Santapau. H. & N.A. Irani (1962): The Asclepiadaceae

; and Periplocaceae of Bombay. Univ. Bombay Bot.

Mem. 4: iv+1 1 8, tab. 12.

Singh, N.P. & S. Karthikeyan (Eds.) (2000): Flora of

Maharashtra State: Dicotyledons, Vol. 1

(Ranunculaceae to Rhizophoraceae). Pp. 87. 103.

Botanical Survey of India, Calcutta.

Tetali, P., Sujata Tetali, B.G. Kulkarni r.r al. (2000):

Endemic Plants of India (A Status report of

Maharashtra State). Pp. 31. Naoroji Godrej Centre

for Plant Research, Shindewadi.

Printed by Bro. Leo at St. Francis Industrial Training Institute, Borivli, Mumbai 400 103 and

published on July 17, 2003, by J.C. Daniel for Bombay Natural History Society,

Hornbill House, Dr. Salim Ali Chowk, Shaheed Bhagat Singh Road, Mumbai 400 023.

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Registered with the Registrar of Newspapers under RN 5685/57 ISSN 0006-6982

EDITORIAL

CONTENTS

SMITHSONIAN INSTITUTION LIBRARIES

1

BREEDING BEHAVIOUR OF THE GREATER ADJUTANT-STORK

LEPTOPTILOS DUBIUS IN ASSAM, INDIA

(With one text-figure and two plates )

By Hillaljyoti Singha, Asad R. Rahmani, Malcolm C. Coulter and Salim Javed 9

CURRENT STATUS OF THE GANGES RIVER DOLPHIN, PLATANISTA

GANGETICA IN THE RIVERS KOSI AND SON, BIHAR, INDIA

( With one text-figure)

By R.K. Sinha and Gopal Sharma 27

CROP DEPREDATION BY WILDLIFE ALONG THE EASTERN BOUNDARY

OF THE KALAKAD-MUNDANTHURAI TIGER RESERVE, SOUTHERN

INDIA

(With two text-figures)

By Punidan D. Jeyasingh and Priya Davidar 38

SMALL MAMMALS IN MONTANE ECOSYSTEMS OF THE NILGIRIS,

SOUTHERN INDIA: THEIR ECOLOGY AND NATURAL HISTORY

By Kartik Shanker 46

THE EFFECTS OF CATTLE GRAZING AND HABITAT ON HELMINTH LOADS

OF CHITAL (AXIS AXIS) IN THE MUDUMALAI WILDLIFE SANCTUARY,

SOUTHERN INDIA

By Guha Dharmarajan, M. Raman and Mathew C. John 58

DEMOGRAPHY OF LIONTAILED MACAQUE (MACACA SILENUS) IN AN

UNDISTURBED RAINFOREST OF SILENT VALLEY NATIONAL PARK,

KERALA, INDIA

( With two text-figures)

By Gigi K. Joseph and K.K. Ramachandran 65

NEW DESCRIPTIONS 72

OBITUARY 94

REVIEWS 96

*

MISCELLANEOUS NOTES 100

Printed by Bro. Leo at St. Francis Industrial Training Institute, Borivli, Mumbai 400 103 and

published by J.C. Daniel for Bombay Natural History Society, Hornbilt House,

Dr. Salim Ali Chowk, Shaheed Bhagat Singh Road, Mumbai 400 023.

website: www.bnhs.org; email: [email protected]

•jj

AUG. -DEC. 2003

C.ESWAKI

BOARD OF EDITORS

Editor

J.C. DANIEL

M.R. ALMEIDA

M.K. CHANDRASHEKARAN

B.F. CHHARGAR

R. GADAGKAR

SNDRANEIL DAS

A.J.T. JOHNSINGH

AJITH KUMAR

T.C. NARENDRAN

A.R. RAHMANI

J.S. SINGH

R. WHITAKER

Assistant Editor

GAYATRI WATTAL UGRA

Editorial Assistance : Divya Fernandez

Layout and cover design : V. Gopi Naidu

About the Cover

Front: The painting of the now extinct Caspian Tigress by C.E. Swan, a rare record from the JBNHS

Vol. 33 (1929), is a fine example of the valuable material contained in the BNHS’s database. The

holotype specimen of the Travancore Tortoise Indotestudo travancorica was recorded by Boulenger

in Vol. 1 7 (1 907). The other illustrations are the Great Indian Bustard Ardeotis nigriceps by H. Gronwold

for Game Birds of India, authored by E.C. Stuart-Baker, Vol. 21 (1912); Danaus genutia male, a butterfly,

by C.B. Williams, Vol. 40 (1938); Heavenly Morning Glory Ipomoea rubro-caerulea by Ganga Singh

for the serial Some Beautiful Indian Climbers and Shrubs by N. Bor and M.B. Raizada, Vol. 41 (1939).

The Scarlet-backed Flowerpecker Dicaeum cruentatum painted by Carl D’Silva appears in The Book

of Indian Birds by Salim Ali and the picture of the Buffstriped Keelback Amphiesma stolata by Isaac

Kehimkar appears in the Book of Indian Reptiles and Amphibians by J.C. Daniel.

Back: The cover of the Inaugural Vol. 1(1) (1886), surrounded by those of Vol. 96(1) (1999) Tiger in

Sundarban by Sudheer Agashe; Vol. 97(1) (2002) Asian Elephant by an anonymous contributor;

Vol. 98(1) (2001) King Cobra Ophiophagus hannah by Shekar Dattatri and Vol. 99(1) (2002) mass

flowering of Karvi Carvia callosa by Ashok R. Kothari.

These illustrations represent some of the most spectacular aspects of the biodiversity of the Indian

subcontinent.

any means, electronic or mechanical, including photocopying, recording or by any information storage

and retrieval system, without permission in writing from the Bombay Natural History Society. Enquiries

concerning reproduction outside the scope of the above should be sent to the Bombay Natural

History Society at the address below.

For Instructions to Contributors, please refer to our website <www.bnhs.org>

Bombay Natural History Society,

Hornbill House, Shaheed Bhagat Singh Road,

Mumbai 400 023.

Editors,

Journal of the Bombay

Natural History Society

VOLUME 100 (2&3): AUGUST-

CONTENTS

EDITORIAL : 167

TIGER ECOLOGY AND CONSERVATION IN THE INDIAN SUBCONTINENT

( With one text-figure and one plate)

By K. Ullas Karanth 169

BEAR CONSERVATION IN INDIA

( With four text-figures)

By A.J.T. Johnsingh 190

KUDREMUKH NATIONAL PARK, KARNATAKA: A PROFILE AND A STRATEGY

FOR THE FUTURE

( With two plates)

By S.A. Hussain 202

DEVELOPING RESPONSIVE INDICATORS FOR THE INDIAN BIOSPHERE RESERVE

PROGRAMME

By Anirban Ganguly, Yogesh Gokhale and Madhav Gadgil 214

PERSPECTIVES ON THE USES OF BIOCIDES: CONSERVATION STRATEGIES

FOR THE NEXT CENTURY

By Robert W. Risebrough 226

PERCEPTIONS OF THE DEVELOPMENT OF WILDLIFE CONSERVATION IN INDIA,

HIGHLIGHTING THE PAST QUARTER CENTURY, AND THE INPUT OF THE

U.S. FISH AND WILDLIFE SERVICE THROUGH THE BOMBAY NATURAL

HISTORY SOCIETY AND THE WILDLIFE INSTITUTE OF INDIA

By David A. Ferguson 240

CHRONOBIOLOGY, ECOLOGY AND BEHAVIOUR OF SOME INSECTIVOROUS BATS

OF SOUTHERN INDIA

(With twenty-three text-figures and one plate)

By M.K. Chandrashekaran 250

TAIL LENGTH IN ENIGMATIC NORTHEAST INDIAN MACAQUES AND PROBABLE

RELATIVES

( With four text-figures)

By Jack Fooden 285

STATUS, ECOLOGY AND CONSERVATION OF THE INDIAN WOLF CAN IS LUPUS

PALLIPES SYKES

( With two text-figures and one plate)

By Yadvendradev Jhala 293

POPULATION STRUCTURE, COMPOSITION AND ABUNDANCE OF ELEPHANTS

ELEPHAS MAXIMUS IN MINNERIYA NATIONAL PARK, SRI LANKA

( With six text-figures and one plate)

By Charles Santiapillai, S. Wijeyamohan, Chaminda Wijesundara

and Rajnish Vandercone 308

THE SOUTHERN KIANG EQUUS KIANG POLYODON

( With six text-figures and two plates)

By G. Neumann-Denzau and H. Denzau 322

TAXONOMY OF UNGULATES OF THE INDIAN SUBCONTINENT

By Colin Groves

341

DISAPPEARANCE OF THE WHITE- WINGED DUCK C A IRINA SCUTULATA FROM THE

PABLAKHALI WILDLIFE SANCTUARY: A SAGA OF LARGE-SCALE

DESTRUCTION OF MIXED EVERGREEN FOREST IN BANGLADESH

( With one text-figure and two plates)

By Mohammad Ali Reza Khan 363

THE INCUBATION MOUND AND HATCHING SUCCESS OF THE NICOBAR

MEGAPODE MEGA PODIUS NICOBARIENSIS BLYTH

( With four text-figures and one plate)

By K. Sivakumar and R. Sankaran 375

THE SONG OF N INOX SCUTULATA OBSCURA

( With one text-figure)

By Ben King 388

LEARNING ABOUT VOCAL COMMUNICATION IN BIRDS

By TJ. Roberts 390

BIRD SPECIES DIVERSITY ALONG THE HIMALAYA: A COMPARISON OF

HIMACHAL PRADESH WITH KASHMIR

(With two text-figures and two plates)

By Trevor Price, Jennifer Zee, Kartika Jamdar and Nitin Jamdar 394

FRUGIVORY, SEED DISPERSAL AND REGENERATION BY BIRDS IN SOUTH INDIAN

FORESTS

(With seven text-figures and one plate)

By P. Balasubramanian and B. Maheswaran 411

CROCODILE CONSERVATION, WESTERN ASIA REGION: AN UPDATE

(With three text-figures and two plates)

By Romulus Whitaker and Harry Andrews 432

GROWTH OF KNOWLEDGE ON THE REPTILES OF INDIA, WITH AN INTRODUCTION

TO SYSTEMATICS, TAXONOMY AND NOMENCLATURE

By Indraneil Das 446

HISTORY AND DEVELOPMENT OF FISHERIES RESEARCH IN INDIA

By E.G. Silas 502

HISTORY OF MARINE SCIENCES (EXCEPT ICHTHYOLOGY) IN INDIA

By B.F. Chhapgar 521

CONTRIBUTIONS TO THE BIOLOGY OF THE QUEENLESS PONERINE ANT

DIACAMMA CEYLONENSE EMERY (FORMICIDAE)

( With five text-figures)

By Vedham Karpakakunjaram, Padmini Nair, ThresiammaVarghese, George Royappa,

Milind Kolatkar and Raghavendra Gadagkar 533

MALARIA IN INDIA — IS AN ECOFRIENDLY SOLUTION POSSIBLE?

By Rachel Reuben 544

ZOOLOGICAL SURVEY OF INDIA AND ITS IMPACT ON THE STUDY OF FIELD

ZOOLOGY IN INDIA

By J.R.B. Alfred 553

GLIMPSES OF THE PHYTOGEOGRAPHY OF MAHARASHTRA

By M.R. Almeida, Suchandra Dutta and S.M. Almeida 559

ON THE DATES OF PUBLICATION OF THE JOURNAL OF THE BOMBAY NATURAL

HISTORY SOCIETY, VOLUMES I- 100 (1886-2003), AND OTHER MATTERS

By Aasheesh Pittie 589

ii

OBITUARY

HUMAYUN ABDULALi (1914-2001) 614

MISCELLANEOUS NOTES

MAMMALS

1. Dogs Canis familiaris hunting the Indian

porcupine Hystrix indica in the wild at

Jodhpur, Rajasthan

By Anil Kumar Chhangani 617

2. Sighting of the Indian wild ass Equus onager

in Rajasthan: a northward range extension

By Harkirat Singh Sangha 6 1 7

BIRDS

3. Ring recovery from great cormorants

Phalacrocorax carbo in India

By R. Suresh Kumar 621

4. Recovery of a ringed demoiselle crane Grus

virgo in Kutch

By S.N. Varu and M.H. Trivedi 624

5. First record of lesser florican Svpheotides

indica (Miller) from Keoladeo National Park,

Bharatpur, Rajasthan

By Ashok Verma and Brijendra Singh 625

6. On the longevity of the great pied hornbill

Buceros bicornis in captivity

By L.N. Acharjyo, Vinod Kumar and

S.K. Patnaik 626

7. Foraging associations and interactions in

woodpeckers

By V. Santharam 627

REPTILES

8. Exploitation of sea turtles along the southeast

coast of Tamil Nadu, India

By S. Bhupathy and S. Saravanan 628

AMPHIBIANS

9. Record of the painted kaloula Kaloula

taprobanica in Andhra Pradesh

By S. Sivakumar, Ranjit Manakadan and

Varad Giri 631

INSECTS

10. Predation of dragonfly Ictinogomphus rapax

(Rambur) (Odonata: Anisoptera) by robberfly

Stenopogon pradhani Joseph & Parui (Diptera:

Asilidae)

By R.M. Sharma and S.S. Talmale 632

1 1 . Congregations of Common Crow butterflies

Euploea core Cramer at Aralam Wildlife

Sanctuary, Kerala

By Vinayan P. Nair 632

OTHER INVERTEBRATES

12. Observations on the ecology of raft spiders

(Araneae: Pisauridae) in Madhya Pradesh

By Pawan Gajbe 634

BOTANY

13. Rediscovery of Aerva wightii Hook. f.

(Amaranthaceae), an endemic, presumed

extinct species, from Tirunelveli district, Tamil

Nadu, India

By M.B. Viswanathan and N. Ramesh 635

1 4. Ethnobotanical study and ex situ conservation

of Alpinia galanga Willd. — a promising

medicinal plant

By N.K. Verma, A. Sharma, P.K. Singh,

P. Kumar and D.K. Hore 638

ACKNOWLEDGEMENTS

We are grateful to the Ministry of Science and Technology,

Govt of India,

FOR ENHANCED FINANCIAL SUPPORT FOR THE PUBLICATION OF THE JOURNAL.

We acknowledge with gratitude a generous donation from the

Seth Purshotamdas Thakurdas and Divaliba Charitable Trust

IN AID OF THE CENTENARY JOURNAL SEMINAR, 2003.

Editorial

The Journal of the Bombay Natural History Society was first published in 1886 as one of

the major activities of the Society founded in 1883 to “stimulate lovers of nature to record

and communicate their observations.” The editorial to the first issue states, “In accordance

with the character this Society has assumed from the beginning, the aim of its Journal will

be as far as possible, to interest all students of nature, ever remembering that there are

many naturalists, in the highest sense of the term, who have not such a technical knowledge

of any particular branch of the science as to be able to enter with interest into questions of

nomenclature and the discrimination of closely allied species.” In the 117 years of its

existence, the Journal has surpassed its modest objective and become one of the main

sources of information on the biodiversity of the Indian subcontinent. In a publication

history straddling three centuries and two millennia, it has achieved a remarkable standard

of excellence comparable to other journals of international repute. For a natural history

journal published by a private Society largely out of revenues derived from its membership

subscriptions and with minimal support from the Government and other sources, it is indeed

a praiseworthy achievement. The Journal, in its first fifty volumes, records the achievements

of its amateur membership, remarkable in the study of the general ecology of the fauna of

the Subcontinent and the taxonomy of both the fauna and flora of the Subcontinent. This

establishes the Journal as required reading for the study of the Subcontinent’s biodiversity.

The Society’s members through their contributions to the Journal established the credibility

of the Society as a source for information on the conservation needs of the Subcontinent

and the laws that were framed for the conservation of the Subcontinent were based on the

contributions of the Society’s members to the Journal on the status of wildlife and the need

for their conservation.

In the second fifty years of its existence, the Journal has been dominated more by

professional biologists rather than amateur naturalists and this is perhaps a reflection of

the status of the study of natural history in the Subcontinent.

To quote Salim Ali from in his editorial to the 50th Volume of the Journal:.

“The membership of the Society does not consist of scientific men alone, nor does

it consist of naturalists pure and simple, nor altogether of persons who look upon

natural history merely as an amusing pastime. It is a conglomerate of all these

types. And this is not all, for while readers of the Journal include some who are

mainly interested in large game animals, the interest of others centres chiefly on

plants or snakes or butterflies or birds. Every branch of the study of animal or

plant life, moreover, has its devotees among them. Some are interested in problems

of evolution or systematics and taxonomy, others in field study and ecology, others

in morphology and laboratory experiments, others in economics and applied

biology or some other line of study, and yet others in nature photography. The

effort to cater for all these polyglot tastes makes the task of editing the Journal one

of absorbing interest, but by no means easy. The problem always is to maintain the

golden mean, and the difficulties involved in the effort are such as the casual

reader can have but a vague conception of. Since the Journal does not pay for

contributions, it is seldom in a position to pick and choose material to any large

extent; nor is it always possible to publish articles strictly in rotation as they are

received. Such delay sometimes causes dissatisfaction among contributors, and in

some cases, where, for instance a new species is described, deferred publication

may even constitute a genuine grievance. It may happen that one particular issue

contains a preponderance of articles on birds, or plants or fish or what have you, or

it may contain more articles on systematics or morphology — ‘dry-as-dusf as they

are commonly dubbed — than perhaps the average reader or field student cares to

be inflicted with. A howl goes up immediately. One member complains that the

Journal is getting much too ‘high brow’ for a simple nature lover like him and

therefore writes in to please accept his resignation from the Society! Another

member complains of a following issue that the Journal has descended to the level

of a story-telling magazine and is no longer a truly scientific publication, therefore

he feels constrained to dissociate his good name from it! And so it goes on. One

finds in the Journal too much of fish and too little of birds; another too much

morphology and too little natural history; and the charges of similar excesses and

deficiencies levelled at the editors are without end. Our sins of omission and

commission are indeed bewildering!”

But they are not new, and that the Journal has survived them during the last 117

years, and even grown from strength to strength to enjoy the high esteem of scientific

workers throughout the world would seem abounding proof that it has, on the whole, been

conducted along the right lines.

J.C. DANIEL

168

JOURNAL . BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003

JOURNAL

OF THE

BOMBAY NATURAL HISTORY SOCIETY

August-December 2003 Vol. 100 No. 2&3

TIGER ECOLOGY AND CONSERVATION IN THE INDIAN SUBCONTINENT

( With one text-figure and one plate )

K. Ullas Karanth1

Key words: Tiger, Panthera tigris , India, ecology, conservation, predation,

social organisation, evolution

The tiger has served as an effective umbrella species in conserving many forms of biodiversity in the

Indian subcontinent. During the last three decades, scientific research employing modern methods

has generated reliable information on tiger ecology in a range of habitats in the Indian subcontinent.

These studies show that tigers evolved as solitary predators of large ungulates, and their social

organisation pivots around breeding females that try to maintain and defend home ranges. Across

the Subcontinent, tiger population densities vary from a low of<l tiger/100 sq. km to a high of 20

tigers/1 00 sq. km, depending primarily on densities of ungulate prey. Although over 300,000 sq. km

of potential tiger habitat still exists in southern Asia, breeding ‘source’ populations for wild tigers

are primarily confined to effectively protected reserves that occupy less than 2% of the overall

landscape, the rest of which acts as a population ‘sink’. Tiger demography is characterised by both

high productivity and mortality. Consequently, the depletion of their prey base due to human over-

hunting appears to be a major threat to tigers, besides habitat loss and poaching. After being

persecuted for centuries and pushed to the verge of extirpation, tigers received official protection

over the last thirty years. However, their future is still not secure because of newly emergent

misplaced priorities in conservation policies. Protecting viable tiger populations in reserves and

buffering them against incompatible human uses of their habitats must continue to be at the core of

the conservation strategy if tigers are to survive this century and beyond.

Introduction

Saving tigers: a landscape species approach to

biodiversity conservation

Through centuries, the tiger Panthera tigris

has been a predominant cultural icon in the Indian

subcontinent (Jackson 1990, Karanth 2001). At

the same time, however, commoners, kings and

'Wildlife Conservation Society (India Program),

26-2, Aga Abbas Ali Road (Apt: 430),

Bangalore 560 042, Karnataka, India.

Email: [email protected]

colonial adventurers have ruthlessly persecuted

tigers. Thereafter, since the early 1970s, several

South Asian countries have tried to use the tiger

as an umbrella species for wildlife conservation

through species recovery plans like India’s

ambitious ‘Project Tiger’ (Karanth 2001). As a

result, in the Indian subcontinent, about 13,181

sq. km temperate forest, 9,043 sq. km wet evergreen

forest, 1 3,736 sq. km moist deciduous forest, 1 9,360

sq. km dry deciduous forest, 6,927 sq. km alluvial

grassland and 873 sq. km mangrove forest have

been proclaimed as protected nature reserves

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003

169

TIGER ECOLOGY AND CONSERVATION

(Dinerstein et al. 1 997), thereby benefiting a whole

range of other species and the overall biodiversity

in general. Thus, in the Indian subcontinent,

survival prospects for biodiversity in its myriad

forms are inextricably linked to effective

conservation of tiger habitats. The anxiety that

the focus on tigers is some form of benign neglect

of other species is thus clearly not justified.

Beginning with Jim Corbett, conserva-

tionists have repeatedly predicted the imminent

extinction of tigers in the Indian subcontinent for

the last 70 years (Karanth 2001). However, these

predictions have failed to materialise because of

the ecological resilience of the tiger (Sunquist et

al. 1999) and due to timely conservation

interventions (Karanth 2001). On the other hand,

governments have also exaggerated their

conservation successes (Panwar 1987) based on

“tiger numbers” they generated using

demonstrably failure-prone techniques (Karanth

1987, 1999; Karanth et al. 2003). Such widely

divergent perceptions about the fate of the tiger

arise from inadequate scientific understanding of

tiger ecology and conservation issues. The

purpose of this paper is to briefly review tiger

ecology in the Indian subcontinent based on

recent and ongoing scientific studies and to

explore future directions for tiger conservation.

Generating a knowledge base for tiger

conservation

Many hunters and naturalists have

produced anecdotal accounts of tiger biology

during the past two centuries (see Thapar 200 1

for some examples): accounts by Brander (1923),

Champion ( 1 929), Corbett ( 1 944), Singh ( 1 984) and

Thapar (1 989) can be cited as good examples from

India. Although qualitative in nature, they

provided useful insights into tiger biology.

Despite the widespread interest in tigers

among hunters of the past, and more recently

among conservationists, and despite the massive

efforts and investments made to recover tiger

populations, knowledge about the species

remains scanty, even within the conservation

community. Most popular literature published

even today repeats the same old flawed cliches:

that there are five ‘subspecies’ of surviving tigers;

that the largest wild tiger population in the world

exists in Sundarbans; that white tigers have

conservation value; that there is a need for

releasing captive-bred tigers into the wild to save

the species; that we reliably know the tiger

numbers in specific reserves, regions or even over

the entire country; and above all, that human

beings and tigers had lived in harmony during

some past golden age, and therefore, special

reserves that try to curtail incompatible human

uses of tiger habitats are unnecessary now.

Although ecological studies of tigers within

a modern scientific framework began forty years

ago with George Schaller’s pioneering work in

Kanha (Schaller 1967) and have advanced

tremendously thereafter as a result of research by

other scientists, much of this new knowledge

appears to have escaped the notice of wildlife

managers and conservationists in the Indian

subcontinent. Therefore, there is an urgent need

to summarise this knowledge that can provide a

foundation for effective action to save tigers.

Major scientific advances in understanding

tiger ecology were made in the 1973-1985 period

through radio telemetry studies in Chitwan, Nepal,

under the Smithsonian Tiger Ecology Project

(Seidensticker 1976, Sunquist 1981, Smith et al.

1987, Sunquist and Sunquist 1988, Smith 1993,

Seidensticker and McDougal 1993). During

thel990s, long-term ecological studies in

Nagarahole (Karanth and Sunquist 1992, 1995,

2000, Karanth et al. 1999), Panna (Chundawat et

al 1999) and other areas of India (Karanth and

Nichols 1998, 2000) that employed modern

techniques such as radiotelemetry, camera

trapping, dietary analyses and prey density

estimation, generated substantial new knowledge

about wild tigers. At the same time, new studies

by taxonomists, geneticists, evolutionary

biologists and biogeographers (Wentzel et al.

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1999, Kitchener 1999, Kitchener and Dugmore

2000) generated fresh insights into issues of

evolutionary origin, radiation, and classification

of tigers. In the following section, I will try to

provide a brief overview of this new knowledge

about tiger biology in the Indian subcontinent.

Biology of the Tiger

Morphology

The tiger is the largest of all living wild cats.

Its body is adapted for stalking and ambushing

ungulate prey up to five times its own size. Its

strong but light skeletal structure and powerful

muscles permit speedy short rushes, leaps and

grappling. The jaw muscles and long canines

enable a strong bite necessary to quickly kill

struggling prey. Although quite spectacular, the

tiger’s coloration and stripes camouflage it well in

the forest brush (Karanth 2001).

The standard body measurements (Riney

1 982) and weights of tigers in South Asia (Pocock

1929; Sunquist 1981; Karanth, unpubl. data) are

as follows: body mass of 175-260 kg for males and

1 00- 1 60 kg for females; total length of 270-3 1 0 cm

for males and 240-265 cm for females inclusive of

an 85-1 10 cm long tail. The height at the shoulder

is 90-110 cm. Contrary to earlier perceptions,

measurements obtained from tigers captured for

radiotelemetry studies in the Indian subcontinent

(Sunquist 1981; Karanth, unpubl. data) show that

they are not smaller than tigers captured in the

Russian Far East (Dale Miquelle and John

Goodrich, unpubl. data).

Tigers possess 30 teeth, with 6 upper and 6

lower incisors, 2 upper and 2 lower canines, 6

upper and 4 lower premolars and 2 upper and 2

lower molars. The upper canines are 50-60 mm

long, and the lower ones 40-50 mm. Like other

cats, tigers have five front toes (only four leave

prints) and four hind toes. The toes have sheathed

claws that can be extended for grasping. Their

large, round eyes possess excellent night vision

and ability to detect movement, but appear to have

poor colour discrimination capability. Their hearing

is acute and is used to locate prey, but the sense

of smell is used primarily for detecting scent from

the ground or vegetation. The cat’s sense of

“touch”, with its padded feet as well as long

vibrissae, is critical for silent movement through

dense cover. Tigers possess several scent glands

around their cheeks, toes, tail and the anal region,

like other cats do (Ewer 1985).

Evolution, radiation and taxonomy of tigers

Tiger evolution has been studied using

fossil evidence and molecular genetic techniques

(Hemmer 1987, Herrington 1987, Kitchener 1999,

Kitchener and Dugmore 2000). Tigers belong to

the family Felidae and the genus Panthera, within

which they branched off as a distinct species even

before lions ( Panthera led), leopards ( Panthera

pardus) and jaguars (. Panthera onca) and were

widely distributed over China and Southeast Asia

even about 2 million years ago. Tigers had

managed to expand their range northwards into

Russia, Japan, the Bering land-bridge, and south

and westwards into the Indian subcontinent and

the Caspian regions by about a million years ago.

By the beginning of the Holocene (about 10, 000

years ago), tigers were found on the islands of

Java and Bali, but not in Sri Lanka (Kitchener and

Dugmore 2000).

The climatic changes that shaped the

expansion of tiger range across Asia during the

Pleistocene and Holocene primarily operated

through changes in connectivity of land-bridges

and landscapes, which in turn was driven by

changes in sea level and vegetation patterns.

These environmental factors led to the evolution

and radiation of large ungulates, particularly

several species of deer (Cervidae) and wild cattle

(Bovinii), opening up an ecological niche for a

large, solitary, forest predator (Sunquist et al. 1 999).

Although tigers tolerate high ambient

temperatures up to 48 °C in northern India, they

are not adapted to the arid, water-scarce

environments in which lions and leopards still

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survive. However, tigers can tolerate severe cold

climate (-35 °C) in the Russian Far East. They live

at altitudes ranging from sea level to 3,000 m

sometimes crossing Himalayan passes at 4,700 m.

Tigers occur in the cold Temperate Zone forests

of northeastern Asia as well as in the hot, humid,

wet or dry forests further south. In the Indian

subcontinent, they are found in Tropical Dry and

Moist Deciduous Forests, Evergreen and

Mangrove Forests, Riparian Grassland-forests of

the terai and in mixed subtropical forests of

Himalayan foothills. Their distribution seems to

be determined primarily by availability of large

ungulate prey rather than by vegetation types

(Sunquist et al. 1 999, Karanth 200 1 ).

The tigers that successfully evolved and

adapted to these varied environments and prey

types, now ‘look’ somewhat different in different

parts of their range. Based on such perceived

morphological differences, taxonomists had

classified tigers into four subspecies ( tigris in

South Asia, virgata in the Caspian region, altaica

in Russia, sondaica in Java) by the 19th century.

Four more tiger subspecies were described in the

20th century: ( amoyensis in southern China,

balica in Bali, sumatrae in Sumatra and corbetti

in mainland Southeast Asia). These traditional

“eight tiger subspecies” were segregated based

on body measurements and pelage details

(Hemmer 1987, Herrington 1987) obtained from

very few museum specimens (Kitchener 1999).

However, more recent syntheses (Wentzel

et al. 1999, Kitchener 1999, Kitchener and

Dugmore 2000) of the genetic, morphological and

biogeographic evidence suggest that this

traditional classification of “eight subspecies” of

tigers is not reliable. New data suggest that

morphological variation in tigers occurs along a

gradient, rather than at the level of discrete

subspecies. Therefore, the more plausible current

models of tiger evolution lump all mainland Asian

tigers into one or two subspecies that are distinct

only from the other subspecies from the Sundaic

Islands. Tigers now surviving in Russia, China,

Indo-China and southern Asia, all appear to

belong to one subspecies that is distinct only

from the island subspecies surviving in Sumatra.

Such studies also highlight the critical importance

of representatively conserving the much wider

range of ecological and behavioural variations in

wild tigers as adaptations to their specific habitats,

rather than merely trying to save traditional

“subspecies” (Wikramanayake et al. 1998,

Karanth 2001) that have little basis in reality.

Communication and social behaviour

Communication and sociality.

Because tigers are solitary animals that live

at very low densities (Sunquist 1981, Sunquist et

al. 1999), communication between individuals,

either to seek out or to avoid one another, is crucial

for maintaining their social organisation. The

tigers’ communication system involves an

elaborate repertoire of chemical, visual and vocal

signals. Common chemical and visual signals

include exuding scent mixed with urine and scats,

leaving visual marks by scraping or rolling on the

ground and clawing trees (Smith et al. 1 989, Smith

1993). A variety of long and short-range vocal

signals, including roars, grunts, growls and purrs

are also used. Among all these modes of

communication, spraying scent mixed with urine

is perhaps the most effective one overall (Sunquist

1981, Smiths/ al. 1989).

Using a combination of such signals, tigers

communicate information on their individual

identity, time of passage, social status, sexual

receptivity and range-ownership to other tigers,

thus enabling contact or avoidance, depending

on the social context. With such a communication

system, tigers maintain their social organisation

with relatively few aggressive encounters that can

have fatal consequences for the animals involved.

Sexual behaviour.

Tigers appear to mate throughout the year

in the Indian subcontinent (Sunquist 1981; Smith

1993; Karanth, unpubl. data). Tigresses advertise

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their oestrous status through increased bouts of

roaring and scent marking that help male tigers

find them. The mating session lasts from 2-7 days

and involves dozens of copulations of about 15

seconds duration every day. Mating tigers

indulge in a lot of aggressive play. The tip of the

male’s penis has backward pointing ‘spines’,

which may provide stimulation to induce

ovulation in the female (Ewer 1985). Once the

mating period is over, the two animals go their

own separate ways.

Relationship between mother and cubs :

Following a gestation of 102-108 days,

litters of 2-5 (usually 3) cubs are dropped in a

secluded hideout. The cubs are born blind and

helpless, and, are aggressively protected by the

tigress. However, rarely, through a temporary

hormonal imbalance, a tigress may kill or even eat

her newborn cubs. The cubs are nursed on milk

for the first two months and effectively hidden

from other predators and even other tigers.

Because the tigress must nurse and guard her

cubs closely, her home range shrinks to a fraction

of its usual size (Sunquist 1981, Smith 1 993). When

the cubs are 2-3 months old, the tigress starts to

take them to her kills. At about 12 months old,

they accompany the mother over her entire home

range (Schaller 1967, Sunquist 1981 , Smith 1993).

Cubs learn to hunt by watching their mother.

Sometimes, several prey animals may be killed

within a few hours, when the tigress teaches her

cubs to kill effectively.

Sometimes, the adult male that sired the cubs

may temporarily associate with the tigress to share

a kill or even play with the cubs (Schaller 1967,

Thapar 1989). A tigress with cubs may sometimes

share kills with her older daughter, who may have

her own litters, leading to temporary associations

of 7-8 related tigers. On the other hand, the tigress

zealously guards her cubs from strange males,

which often exhibit the infanticide behaviour

common to many mammalian species (Smith 1 993;

Karanth, unpubh data).

Between 12-18 months age, tiger cubs

acquire their permanent teeth and become

proficient killers. They learn to search, stalk,

capture and kill potentially dangerous prey. By

the age of 1 8 months, juvenile males make forays

away from their mother’s range to begin a

transient life. Juvenile females stay close to their

mother, but eventually disperse by 20-28 months,

as their mother becomes increasingly aggressive

towards them (Smith 1993). By then, the tigress

would have come into oestrus again and mated.

Population structure and social organisation:

The structure of the typical tiger

population can be described in terms of the sex

and age categories of its members. Tigers of both

sexes can be categorised into demographic

stages (Karanth and Stith 1999) such as cubs

(less than 1 year old), juveniles (1-2 years old),

post-dispersal floaters or transients (over 2 years

old) and breeding adults (3-12 years old). A few

of the transients may also be old or incapacitated

breeders evicted from their ranges by more

vigorous successors.

Tiger social organisation pivots around

breeding females that maintain fixed home ranges

within which they try to raise cubs. These

tigresses acquire ranges by evicting previous

residents, or by ‘inheriting’ a part of their

mother’s range (Smith et al. 1987, Smith 1993).

Normally, in a good habitat stocked with enough

prey, a tigress starts to breed at 3-4 years. Her

normal residential tenure is of 5-7 years, after

which she loses her range to a competitor. The

degree of spatial overlap (or lack of it) between

neighbouring female ranges varies, depending

on prey density and other ecological factors

(Sunquist 1981, Smith et al. 1 987, Miquelle et al.

1999, Karanth and Sunquist 2000). Adult males

have larger ranges that overlap ranges of several

breeding females, the average being about three.

The degree of exclusiveness of male ranges is

variable, depending on factors that are as yet

unclear. The land tenures of breeding males are

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TIGER ECOLOGY AND CONSERVATION

shorter (2-4 years) than those of females (Sunquist

1981, Smith 1993).

Although the ranges of neighbouring

female tigers were exclusive in Chitwan (Sunquist

1981, Smith et al. 1 987, Smith 1 993), there appears

to be some overlap in Nagarahole (Karanth,

unpubl. data). In Chitwan, male ranges were

mutually exclusive, but this was not clear in

Nagarahole. In Chitwan and Nagarahole, the size

of breeding female ranges varied from 13-30 sq.

km and that of males from 40- 1 00 sq. km (Sunquist

1981, Smith 1 993, Karanth and Sunquist 2000). In

Panna Reserve in central India, where tiger

habitats are just beginning to be restored through

protection, the range sizes were 243 sq. km for a

male and 27 sq. km for a female, respectively

(Chundawat et al. 1999). While female range size

seems to be determined primarily by prey

abundance, the size of a male’s range appears to

be a function of the number of female ranges that

he manages to cover.

At about 1 8-28 months age, juveniles either

leave their natal ranges or are evicted by their

mother to become transients. Such floaters,

particularly males, criss-cross several breeder

ranges and even disperse away into new areas.

These transient tigers have large ranges, which

may cover an entire ecological unit. During a six-

month period, two transient males had ranges

of 99 and 77 sq. km in Nagarhole and the

latter’s range shrank to 44 sq. km when he

acquired a breeding range (Karanth and Sunquist

2000). Radiotelemetry studies in Chitwan (Smith

1993) showed 10 dispersing males travelled an

average distance of 33 km, and four females a

distance of 10 km, before establishing their own

ranges. However, occasionally such dispersers

travel great distances of 100 km or more (Smith

1993).

In productive tiger populations, there is

intense competition for breeding ranges.

However, breeders of both sexes tolerate the

passage of their transient offspring within their

ranges and transient siblings seem to tolerate

each other. However, when a breeder male is

replaced, the new male systematically kills cubs

of resident tigresses within his range. Through

such infanticide, the tigresses are induced to come

into oestrus again and mate, thus conferring an

evolutionary advantage to the male through

propagation of his genes (Smith 1993).

Predatory behaviour and ecology

Prey types and prey selection :

The need to hunt alone and kill large

ungulate prey has been the driving force behind

evolution of tigers (Seidensticker and McDougal

1993, Sunquist et al. 1999). Although tigers kill

prey ranging in size from frogs to adult gaur ( Bos

gaurus) — the bulk of their requirement must

come from deer, pigs, and wild or domestic cattle,

that weigh between 20-1,000 kg (Sunquist et al.

1999, Karanth 2001). Apart from livestock, the

principal wild prey of tigers in the Indian

subcontinent include: wild pig Sus scrofa , sambar

Cervus unicolor , barasingha Cervus duvaucelii,

red deer Cervus elaphus , chital Axis axis , hog

deer Axis porcinus , muntjac Muntiacus muntjak ,

nilgai Boselaphus tragocamelus, chousingha

Tetracerus quadricornis, chinkara Gazella

bennettii , blackbuck Antilope cervicapra , gaur,

wild buffalo Bubalus bubalis, takin Budorcas

taxicolor, goral Naemorhedus goral , serow

Naemorhedus sumatraensis , and, occasionally,

elephant Elephas maximus and rhino Rhinoceros

unicornis calves. Tigers also opportunistically

kill and eat other carnivore species such as sloth

bears Melursus ursinus, leopards and dholes

Cuon alpinus.

Activity and hunting behaviour.

Tigers hunt primarily after dark, when their

superior vision confers an advantage (Sunquist

1981, Karanth and Sunquist 2000). Tigers tend to

be active at the same times of the day when their

prey are also active because they can more easily

detect and home in on the latter. Consequently,

human activities such as forest product collection

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and hunting that compel prey species to become

nocturnal, also compel tigers to do the same.

Usually tigers become active at dusk, and

remain so through the night until dawn. During

hot parts of the day, they rest under shade, often

lying up in water (Schaller 1967, Thapar 1989).

Radiotelemetry locations in Chitwan and

Nagarahole (Sunquist 1981, Karanth and Sunquist

2000) showed that tigers moved around a lot more

at night (80% of locations) compared to midday

(10% of locations). They typically remain active

for about 6-8 hours in a day. The linear distances

between radio-locations on successive days were

1-8 km (Sunquist 1981, Karanth, unpubl. data)

although the actual distance walked overnight

was more (5-25 km).

Tigers pad along forest trails and locate

prey through hearing (in dense cover) or visually.

Occasionally, they lie in ambush at localities

favoured by prey, like water holes, clearings or

salt licks (Karanth and Sunquist 2000). Some

observers (Schaller 1967, Thapar 1989) guess that

tigers have a 5-10% success rate while hunting.

Because of difficulties in observing an unbiased

sample of hunts, it is difficult to validate such

estimates. However, success rates are likely to

strongly depend on probabilities of encountering

prey, and, therefore, on prey densities in an area

(Karanth and Sunquist 1995).

After locating the prey, the tiger stalks it

silently to get within range for a final rush of 1 5-

30 m (Seidensticker and McDougal 1 993, Karanth

and Sunquist 2000), or longer across open

clearings or waterbodies (Thapar 1989). The tiger

usually attacks the prey from the flanks or rear,

and knocks it down by the impact of its

momentum and by grappling with its forelimbs.

Simultaneously, it tries to bite the prey animal’s

throat or nape to immobilise it (Seidensticker and

McDougal 1993). The tiger tries to keep away

from the flailing hooves and horns of large

ungulate prey. With dangerous prey such as adult

gaur or buffalo, if the initial attempt to knock down

and immobilise does not succeed, tigers may

even give up the attempt. Sometimes, tigers get

injured or killed by such quarry. After the prey

animal is brought down, it is killed by

strangulation or by rupturing of the cervical

vertebrae, spinal chord, brain case or major blood

vessels. Tigers deliver the lethal bite to the throat

of larger prey animals, whereas smaller prey, such

as pigs or chital may also be killed with a nape

bite.

Feeding ecology.

Tigers in Nagarahole dragged their kills

over distances ranging from 0-350 m (with an

average of 5 1 m, for a sample of 1 33 kills), hiding

the carcasses in dense cover unless the prey

was too heavy (Karanth and Sunquist 2000).

Tigers consume 20-35 kg of meat in their first

meal, and unless disturbed, stay close by to guard

their kills from other tigers and scavengers.

Depending on the kill size and the number of

tigers feeding, they stay with the kill for 1-7 days,

eating two thirds of the kill including some fairly

putrid meat. The remaining one third, comprising

of larger bones, rumen contents and intestines is

normally discarded (Karanth and Sunquist 2000).

When hungry, tigers scavenge kills made by other

tigers or by other predator species.

Killing and cropping rates'.

Tigers are provisioned 1 ,825-2, 1 90 kg meat/

year in captivity (Sunquist 1 98 1 ). A female tiger

kills about 40-45 ungulate prey/year, consuming

about 2,000 kg of meat (or about 3,000 kg of live

prey), just for maintenance. The quantity of live

prey consumed by adult males is higher (4,000

kg/year) and by juveniles and cubs is less. A

tigress raising three cubs has to kill 60-75 ungulate

prey/year (Schaller 1967, Sunquist 1981, Sunquist

et al. 1999). Thus an ‘average’ tiger can be

estimated to take about 50 prey animals or 3,000

kg of live prey annually (Schaller 1967, Sunquist

etal. 1999).

Tigers may crop roughly 10-15% of

available prey in an area annually, depending on

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how much prey is additionally killed by other

predators such as leopards, dholes and human

hunters. Considering the natural reproductive

rates of ungulate prey, a 10% cropping rate

translates into the requirement of a standing

prey base of about 400-500 ungulates to support

a single tiger through one year. Therefore, sizes

of tiger home ranges depend on prey densities.

Ranges of breeding female tigers vary from a low

of 10-20 sq. km in prey-rich habitats to 200-300

sq. km or more in poor quality habitats (Sunquist

et al. 1 999, Miquelle et al. 1 999). Therefore, more

tigers can ‘pack’ into an area at higher prey

densities, although social spacing behaviours

may set an upper limit on tiger numbers. In the

semi-arid area of Panna, availability of water and

prey distribution (rather than prey abundance

alone) influences tiger home range sizes

(Chundawat et al. 1999, R.S. Chundawat, unpubl

data).

Population ecology

Population densities'.

Over most of their range, tigers coexist with

other predatory carnivores such as leopards and

dholes. The relative densities of each predator

species in such guilds appear to be determined

by the relative abundance of different size classes

of prey in the assemblage (Karanth and Sunquist

1995, Karanth and Nichols 1998). Furthermore, as

noted earlier, densities of tigers appear to be

primarily a function of prey densities (Schaller

1 967, Sunquist 1981, Seidensticker and McDougal

1993, Karanth and Sunquist 1995, Karanth and

Nichols 1998, Sunquist et al 1999, Chundawat et

al. 1999).

As primary predators of large ungulates,

tigers cannot sustain themselves or reproduce in

the absence of such prey in sufficient numbers,

even if smaller prey are quite abundant (Schaller

1 967, Sunquist 1981, Karanth and Sunquist 1 995,

Sunquist et al. 1999). Recent studies that

estimated tiger and prey abundance using

rigorous methods (Karanth and Nichols 1998,

2000, Sunquist et al. 1999) clearly show a strong

positive relationship between abundance of large

ungulates and tiger densities.

As prey densities get lower, female ranges

become larger, reducing the number of such

breeders the area can support. Lower prey

densities also appear to result in lower densities

of transient tigers (Karanth and Nichols 1998,

2000). Because survival rates of cubs and

juveniles are also likely to be lower at lower prey

densities (Karanth and Stith 1 999), the numbers

of tigers in these two demographic stages will

also be lower. Therefore, while other habitat-related

or managerial factors may influence tiger density

at a given site, prey abundance appears to be the

most critical determinant.

However, accurately estimating tiger

densities is difficult (Karanth and Chundawat

2002). The Indian Government’s official pugmark

censuses have often yielded unreliable results

(Karanth 1987, 1988, 1999, Karanth etal. 2003).

Camera trap sample surveys within a formal

Capture-Recapture modelling framework has

proved to be a good method for obtaining reliable

tiger density estimates in well-protected study

areas, particularly at higher tiger densities

(Karanth and Nichols 1998; Plate 1, Figs 1, 2).

Mean densities of tigers (excluding cubs <1 year)

derived using the photographic Capture-

Recapture approach in some typical tiger habitats

in India were: Pench (Madhya Pradesh) 4.9 tigers/

100 sq. km, Kanha 11.7 tigers/100 sq. km,

Nagarahole 11.9 tigers/ 100 sq. km, Kaziranga

16.8 tigers/ 100 sq. km (Karanth and Nichols

1998).

The above cited density estimates show

that alluvial grassland-forest mosaics of the

Himalayan foothills and moist-deciduous forests

of peninsular India potentially support the highest

densities of tigers anywhere in the world (15-22

tigers/100 sq. km, including cubs). At the other

end of the ecological scale, in the Russian Far

East, tiger densities are as low as 0.5- 1.5 tigers/

100 sq.km (Miquelle et al. 1999).

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Karanth, K.U.: Camera trapping of Panther a tigris

Plate 1

Fig. 1: Camera trap being set up for studying tiger density in Nagarahole National Park

Fig. 2: A tiger ‘photo-captured' by the camera trap

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TIGER ECOLOGY AND CONSERVATION

Fecundity and mortality rates :

In good habitats, tigresses appear to have

an oestrous cycle of 20-30 days, beginning at

around 2 years of age or earlier, but they conceive

and produce cubs only at 3-4 years of age after

acquiring permanent home ranges. The sex ratio

is equal at birth. If cubs survive to dispersal age,

usually the tigress will produce the next litter after

2-2.5 years. If the cubs die in the interim, the tigress

comes into oestrus almost immediately. In prey-

rich habitats, the average cub production can be

estimated at roughly 1 cub/breeding female/

year (Karanth and Stith 1999). In such a productive

population of 20 breeding females, roughly a third

breed every year, adding about 20 new tigers to

the population.

However, this high productivity of tigers is

balanced by naturally high mortality rates. Tigers

in all demographic stages die from a variety of

causes. Male tigers that acquire new ranges try

to kill the cubs of the former resident breeder.

Other factors that cause mortalities among cubs

include: starvation, floods, forest fires, other

predators and human persecution. Hunting of

tigresses also leads to indirect mortalities of

dependent cubs (Sunquist 1981, Smith 1993,

Karanth 200 1 ). Juvenile tigers ( 1 -2 years) die from

starvation, hunting-related injuries and intra-

specific aggression.

During dispersal phase, transient tigers

move back and forth through larger ecological

units, criss-crossing boundaries of breeder

territories, nature reserves and even states or

countries (Smith 1993). Such transient tigers

suffer heavy attrition through starvation, intra-

specific aggression and human persecution.

Based on data from Chitwan and Nagarahole,

demographic models of tiger populations built by

Kenny et al. ( 1 995) and Karanth and Stith ( 1 999)

assumed the following approximate annual

mortality rates in healthy tiger populations: cubs

of both sexes 40%, juveniles of both sexes 10 %,

transient males 35%, transient females 30%,

breeder males 20% and breeder females 1 0%.

Factors that influence tiger population dynamics'.

Karanth and Stith (1999) recently built a

stochastic population model for tiger populations

under different ecological scenarios. Their model

of a productive (but insular) wild tiger population

with a ‘carrying capacity’ of 24 breeding females,

generated the following typical age-sex structure:

8 breeding males, 28 transients, 14 juveniles and

24 cubs. This population produces 24 cubs/year

on an average, and, would be balanced by annual

mortalities of the same magnitude. This model

suggests that even relatively small wild tiger

populations with only 12 breeding tigresses may

be demographically viable. The model explains

well how tiger populations could have survived

the heavy pressure inflicted by hunters and

poachers in the past. For example, between 1 860-

1960, approximately 93,000 tigers were legally killed

for sport or bounties in parts of British India and

some princely states (Rangarajan 1999).

The demographic model of Karanth and

Stith (1999) indicates that rebounding tiger

populations will quickly reach saturation densities,

and remain relatively stable thereafter, because

any further increase in reproduction rates is

balanced by increase in mortality rates or

dispersal. Therefore, in most productive tiger

populations, there is potentially a ‘doomed

surplus’, that perishes annually, without lowering

the tiger population density (Karanth 2001).

The Karanth and Stith (1999) simulations

also suggest that prey depletion (caused by

human hunters or competition with livestock)

reduces the numbers of breeding females and

transients, as well as depresses cub survival rates.

On the other hand, moderate levels of tiger

poaching may simply remove a part of an existing

annual surplus. In prey-rich habitats, a tigress

may produce 3-4 litters or about 9- 1 6 cubs during

her reproductive tenure (Sunquist 1981, Karanth

and Stith 1999), thereby producing a substantial

‘surplus’ of tigers. Consequently, human-induced

prey depletion may be a far more serious threat to

the viability of tiger populations than moderate

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levels of tiger poaching (Karanth and Stith 1 999,

Karanth2001).

Tiger Conservation in the Indian Subcontinent

Conservation status of wild tiger populations

In historic times, tigers were found all the

way from the Temperate Zone forests of the

Russian Far East to the tropical forests of

southwestern India. They ranged from Azerbaijan

and Iraq on the West all way through the Indian

subcontinent to parts of southern China, eastern

Russia and Southeast Asia. Their range covered

30 present day countries, stretching over 70

degrees of latitude and 100 degrees of longitude

on the earth’s surface (Seidensticker et al. 1999,

Karanth 2001).

Within the Indian subcontinent, tigers were

present in a diverse array of habitats: Tropical

Dry and Moist Deciduous Forests, Evergreen and

Mangrove Forests, terai grasslands and Mixed

Conifer-broadleaf Forests in the Himalayan

foothills. Availability of ungulate prey, water and

shade seem to determine their distribution

(Sunquist et al. 1999, Chundawat et al. 1999,

Karanth 2001). Tigers once overlapped with lions

in a wide region stretching across northwestern

India. Early human modifications of landscapes

such as development of water resources, and

extirpation of the more-easily hunted lions by

human societies might have benefited tigers,

allowing them to expand their range in drier

regions of northwestern India by moving into

newly-opened ecological niches (Karanth 2001).

Recent assessments (Wikramanayake et al.

1999) show that the current distributional range

of tigers in the Indian subcontinent is around

350,000-400,000 sq. km (Fig. 1). Even within this

reduced range, reproducing tiger populations are

now restricted to a few better-protected reserves

that may cover about 40,000 sq. km, or less than

one percent of the tigers’ historical range (Karanth

2001). The mountainous regions of the Himalaya,

the dense evergreen forests of northeastern India

as well as mangrove forests of India and

Bangladesh, are inherently poor quality habitats

for tigers. Lowland areas of Nepal and several

Indian states have high quality habitat patches

within the 50 or so protected areas that

sporadically occur in a matrix of human-dominated

landscapes.

Perhaps over 90% of their range, local tiger

populations cannot be sustained without periodic

immigration from breeding populations in

protected areas. In the overall landscape, such

protected areas might be the only ‘sources’ from

which tigers disperse and perish in the

surrounding ‘sinks’ (Karanth 2001, Karanth and

Chundawat 2002).

Furthermore, many of these ‘source’ tiger

populations are under threat from prey depletion,

tiger poaching, and, habitat degradation and

fragmentation. These threats arise from a variety

of factors linked to local rural uses as well as

economic development projects (Karanth 2001).

The potential erosion of genetic variability in wild

tiger populations as a result of habitat

fragmentation is considered to be a major threat

by some workers (Tilson and Christie 1999).

However, Caughley (1994) averred that the

smallness of animal populations should be viewed

as a consequence rather than a cause of animal

extinctions.

All the above threats to tigers are generally

recognised in scientific and popular literature

(Seidensticker et al. 1 999). Therefore, the essential

challenge now lies in setting appropriate priorities

in responding to these threats. Such priority

setting must necessarily be based on an objective

evaluation of past successes and failures in tiger

conservation.

A brief history of tiger conservation

It is necessary to have a brief overview of

the social context and history of tiger conservation

in the Indian subcontinent before exploring

current conservation issues. This outline is

necessarily brief, and has to be read in the context

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TIGER ECOLOGY AND CONSERVATION

Fig. 1 : Potential tiger habitat in the Indian subcontinent

BANGLADESH

POTENTIAL TIGER HABfTAT

300

Kilometres

600

o\

On

ON

<u

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ca

e

5

cn

On

O

C/O

W

«

o

3

O

O0

of more detailed historical analyses (Rangarajan

1999,2001).

Throughout the 1 9th century and during the

first half of the 20th, tigers were viewed as vermin

to be eradicated by giving bounties to local

hunters or as glamorous game animals to be

exclusively hunted by elite ‘sportsmen’. Both

colonial rulers and native chieftains adhered to

this view. Although some local cultures have

traditionally revered tigers as deities (Jackson

1 990, Karanth 200 1 ), in practice such reverence

made no difference to tigers as people persistently

hunted tigers and their prey, and encroached on

tiger habitat to convert it to farmland. However, in

a few “game reserves” established for the

sportsmen of the ruling classes who wielded

substantial social power, “native poachers” were

kept out. Consequently, tiger populations

survived despite heavy pressure from elite

hunters.

In the post-colonial period (after the 1950s)

the first systematic attempts at tiger

conservation were made under pressure from

hunter-naturalists (Rangarajan 2001). These

included the introduction of the first wildlife

protection laws and establishment of “game

reserves” all over India. However, due to the

weakness of these efforts, tigers continued to

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TIGER ECOLOGY AND CONSERVATION

be under pressure and were in decline into the

1960s. Only a few naturalists (Gee 1964, Daniel

1970) pleaded the case for more effective laws

and protected areas for tigers. Around the same

time, Schaller ( 1 967) published the first scientific

study of tigers, forcefully publicising their grim

conservation status in the Indian subcontinent.

Despite these early warnings, commercial tiger

hunting by foreign tourists as well as “sport

hunting” by the local elite prevailed, particularly

in the forests of Central India (Sankhala 1978).

During the late 1960s assessments by Sankhala

(1978) and a questionnaire survey by Daniel

(1970), both re-confirmed the tiger’s precarious

status in India, suggesting (without much

evidence) that tiger numbers in India could be

lower than 2000 animals. Such concerns led to

effective lobbying and support from international

conservation agencies (mainly IUCN and the

World Wildlife Fund) that resulted in the

Governments of India and Nepal initiating

stronger tiger conservation measures (Jackson

1990).

At the core of these fresh conservation

initiatives were the greatly strengthened wildlife

protection laws, special protected reserves for

tigers and increased funding for protective

infrastructures (Panwar 1987, Karanth 2001). The

criticisms that such tiger recovery measures were

based on a top-down perception of conservation

and did not sufficiently empower or involve local

people in tiger conservation (Saberwal 1997,

Rangarajan 2001) are valid. However, it is equally

true that in most protected areas where these “top-

down” conservation measures were implemented,

significant recovery of tiger populations, prey

base and habitats was observed (Karanth et al.

1999, Karanth 2001). These recoveries resulted

from measures that met the ecological needs of

tigers by reducing pressures of incompatible uses

of tiger habitats by local people as well as by

forestry departments. Had these unpopular

protective measures not been put in place, it is

very likely that tigers would have been extirpated

from even these protected areas, as indeed they

were from forests outside them.

Shifting conservation paradigms

Unfortunately, the tiger population

recoveries in the 1970s and ’80s in the Indian

subcontinent were not documented using

rigorous science (Karanth et al. 2003). As a result,

the conservation community at large did not draw

correct inferences from these large-scale

conservation “experiments”. Consequently, the

proposed alternative of a more “bottom-up”

conservation policy (Kothari et al. 1995)

downplays the importance of preservationist

measures. Instead it emphasises “sustainable

use”. This alternative conservation paradigm

appears to advocate “human-tiger coexistence”

and multiple use of even designated priority tiger

conservation areas (Kothari et al. 1995, Saberwal

1997). Although tigers did “coexist” with

subsistence-level human use of their habitats in

the past, they lost ground steadily as a result of

the ensuing conflict. The tiger’s distributional

range shrank by more than 95 percent in the

process, within a few centuries. How such

coexistence can now become beneficial to tigers

again remains undemonstrated.

Tigers are landscape animals that typically

live at low densities. The area needed to support

a wild tiger population with 25 breeding females

may range from 500 to 5,000 sq. km, depending

on prey density and other habitat parameters.

Being large-bodied carnivores, tigers readily kill

livestock and occasionally, humans. When tigers

lose their natural fear of humans, they can become

persistent man-eaters (McDougal 1987, Karanth

2001). Usually, increased human use of tiger

habitat depresses densities of principal ungulate

prey (Karanth et al. 1999) through hunting,

competition with livestock, and over-harvest of

vegetation. Consequently, productive tiger

populations cannot “coexist” with activities such

as agriculture, livestock grazing, minor forest

product collection and intensive logging,

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without serious human - tiger conflict. In the long

run, such conflicts inevitably lead to the

extirpation of tigers from areas of intensive human

use.

Therefore, the assertion that local

involvement in resource extraction from tiger

habitats, together with the political empowerment

of ‘local people’ are sufficient conditions to

recover tiger populations, rests on no solid

evidence either in ecological theory or in

conservation experience. While there could be

arguments over ‘who’ should manage tiger

reserves, there is no doubt that such special

reserves are needed, and, ‘someone’ has to

enforce the preservationist measures necessary

to maintain and perpetuate them.

Currently, as a result of increasing human

and livestock densities, higher rates of forest-

product harvests and hunting (all linked to

growing commercial markets), wild tiger

populations are declining. Intensification of

human use of tiger habitats, and economic

development projects, both now present major

threats to their ultimate survival (Karanth 2001).

Scientific knowledge of tiger biology as well

as our empirical knowledge of past conservation

practices (what works and what does not)

reviewed above, clearly show that reducing

human pressures on tiger reserves and buffering

them against incompatible land use practices

outside, and extending and increasing

connectivity among these reserves, are the best

ways to recover and sustain wild tiger

populations.

A renewed commitment to such effective

protected tiger reserves, under a new conservation

paradigm of maintaining ‘sustainable landscapes’

overall, rather than implementing ‘sustainable use’

practices everywhere, seems to be the most

fundamental paradigm change necessary to save

tigers in this century and beyond. However,

conservationists appear to be often distracted

from this central task by a variety of ‘surrogate

tiger conservation activities’.

Surrogate tiger conservation activities

Because the practical measures necessary

to recover wild tiger populations are socially and

politically complex and difficult to implement in

almost any specific local context, conservationists

often escape from them by taking up ‘surrogate

tiger conservation activities’. As a result,

currently, a substantial amount of goodwill,

concern, effort and financial resources meant to

support tiger conservation are being misdirected

at implementing such surrogate solutions. While

some of these activities simply divert scarce

resources and energy away from more immediate

needs, others adversely affect tiger conservation

in a more direct manner.

One example of surrogate tiger conserva-

tion is the disproportionate attention paid to

initiatives that involve captive breeding, assisted

reproduction and reintroduction of tigers,

ostensibly for augmenting wild populations or

maintaining genetic viability or for promoting

animal welfare. However, this approach is

fundamentally flawed for reasons explained in

the following paragraphs.

Wild tiger populations have been extirpated

through demographic causes (increased mortality,

decreased reproduction) that are driven by over-

hunting, prey depletion and habitat loss.

Reintroductions serve no purpose in replenishing

wild populations because the introduced tigers will

also be eliminated by the same causes. As to the

issue of genetic viability, leading conservation

geneticists and population ecologists (Lande 1 988,

Caughley 1 994) have argued that smallness of wild

populations is most likely an effect of demographic

factors and not their cause. Even if infusion of new

genes into an isolated tiger population is

scientifically demonstrated as a real need, it is more

efficient and practical to capture and translocate

dispersal-age individuals between wild tiger

populations, rather than reintroduce captive animals

that have to be reared and trained at great cost.

Given what we know about the social

organisation of tigers, even in the unlikely event

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that an introduced animal establishes a range and

breeds, such an introduction will inevitably

involve the killing or eviction of an existing

resident breeder. It will also have the same

disruptive influence that the natural turnover of

resident breeders in a population involves.

Because the number and sizes of tigers an area

can support is limited by prey density, there will

be no net population gain even after a successful

introduction. Even in a rare (and expensive) case

of successful reintroduction, the new animal

simply replaces some other less fortunate

individual in the population. Therefore, tiger

reintroduction can neither be logically justified

as a population augmentation tool, nor even be

defended as an animal welfare measure.

On the other hand, if the reintroduced tiger

fails to establish a home range, as is more likely,

there are high probabilities of the cat turning into

a ‘problem animal’ that kills livestock or even

humans. Such an event will further aggravate the

antipathy local people already feel towards tigers

and exclusive reserves that protect them, thus

making the job of genuine tiger conservationists

even more difficult than it already is.

Another example of surrogate conservation

involves diversion of tiger conservation monies

to other worthy social objectives like providing

schools, hospitals and other services to people

living around tiger habitats. Such activities indeed

should be funded, but out of the much larger pool

of money earmarked for developmental and social

causes. Given that resources earmarked for tiger

conservation are meagre, and the demonstrated

immediate needs are for improved protection,

habitat consolidation, conservation monitoring

and conservation education — all currently

under-funded activities — diversion of

conservation funds to meet social objectives that

have no immediate impact on wild tigers is also a

form of surrogate tiger conservation.

An extreme example of the above approach

are the current “eco-development projects” of

various kinds being implemented with the

assistance of the World Bank-GEF combine and

other multilateral aid agencies, and with

enthusiastic support from a large number of Indian

officials and conservationists. These projects

have diverted huge amounts of money, energy

and attention away from the core issue of

tiger protection towards largely wasteful

“developmental” activities leading to further

deterioration of tiger protection (Karanth 2002).

Another oft-promoted surrogate activity is

‘eco-tourism’, a term often misapplied to the

expensive corporate sponsored form of wildlife

tourism practised in India. It is true that well-

managed tourism involving charismatic animals

like tigers can generate substantial revenues for

the reserves and help protect tigers directly. Such

schemes can also potentially generate incomes

and revenues for the local people around the tiger

reserves, thus engendering additional public

support for tigers. Rarely, such projects can also

lead to land use changes outside the reserve that

are favourable to tigers (Dinerstein et al. 1 999).

However, unfortunately, the high-revenue tiger

tourism practised in India is singularly devoid of

any of these positive features. It simply

constitutes a large net drain on the park budgets

and engenders mostly apathy or even hostility

among local residents whose access to and use

of tiger habitats has been curtailed to meet tiger

conservation needs. Therefore, in the Indian

subcontinent, despite its vast potential, eco-

tourism has been turned into another surrogate

tiger conservation activity that confers little

benefit to the target species.

Beyond the pugmark census: ecological

monitoring of tigers

We need to monitor tiger populations for

three fundamental reasons. Firstly, to objectively

evaluate the success or failure of past conservation

interventions, so as to react adaptively to solve

problems. A second major goal is to establish

benchmark data that can serve as a basis for future

management. A third overarching goal of tiger

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monitoring is to improve our basic understanding

of tiger ecology and behavior to develop a body

of empirical and theoretical knowledge that can

potentially improve our predictive capacity to deal

with new situations (Karanth et al. 2002).

At this point, given the critical status of

tigers and the substantial investments made in

their conservation, wildlife managers and

conservation agencies need clear and reliable

answers to some basic questions given below

(Karanth et al. 2002, Karanth and Chundawat

2002), without which they cannot even begin to

evaluate the success or failure of tiger

conservation:

1 . What is the extent and range occupied

by different individual tiger populations?

2. Where are individual tiger populations

increasing their range, and, where are these ranges

fragmenting or shrinking?

3. Within the distributional range of tigers,

what is the proportion of the area occupied by

productive, breeding populations?

4. In important individual tiger reserves,

what are the tiger population trends? Are tiger

populations in such reserves holding steady,

declining or increasing?

The traditional approach to answering such

questions has been based on attempts to obtain

total counts of wild tigers all over the country

through ‘pugmark censuses’ (Choudhury 1970,

1972). Such ‘census-based5 approaches have

major biological and statistical weaknesses

(Karanth 1987, 1999, Karanth and Chundawat

2002, Karanth et al. 2003). Therefore, the need to

employ more reliable ‘population sampling-based’

methods tailored to suit a variety of practical

contexts is being increasingly realised (Anon.

1997, Karanth et al. 2002)

During the past 35 years, our knowledge of

tiger ecology has advanced significantly as a

result of several scientific studies. During the same

period, methodologies for assessing wildlife

population parameters have also developed

substantially. In particular, two conceptual

approaches to population sampling, Distance

Sampling and Capture-Recapture Sampling, have

advanced particularly rapidly (Thompson et al.

1 998, Williams et al. 2002). Such methods now offer

powerful tools for ecological monitoring of tigers

and other wildlife in India. The type of monitoring

feasible in each specific context can be determined

by considering the potential methods in relation

to available resources and local conditions.

The following guidelines from Karanth et

al. (2002) may be useful in choosing a monitoring

method appropriate to any specific local context:

1. It is almost impossible to estimate

absolute or even relative densities of tigers or

prey if trained manpower, equipment and other

resources are extremely limited, and, large regions,

states or countries have to be covered. Since most

of the distributional range of tigers in the Indian

subcontinent typifies such conditions, one can

only attempt sample surveys of presence of tigers

and prey species to estimate and map their

distribution at this large landscape scale.

2. Where adequately trained personnel are

available, measuring the relative density of tigers

from sample surveys of encounter rates with tiger

tracks or scats or the relative densities of prey

species from pellet or dung counts are options

(Karanth and Kumar 2002). It is likely that such

index-based surveys are feasible only in some

individual reserves.

3. If special equipment, trained personnel and

other resources are available, absolute densities of

prey species can be estimated from line transects

using distance sampling methods (Buckland et al.

1993, 2001). Even absolute densities of tigers can

be estimated using photographic capture-recapture

sampling from camera trap surveys (Karanth and

Nichols 1998, 2002a). Such advanced methods are

likely to be practical only in a few priority tiger

reserves or study sites.

If there is a mismatch between available

resources and the goals that tiger managers hope

to achieve, failure of monitoring is almost certain.

The goal (1) of monitoring tiger or prey spatial

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TIGER ECOLOGY AND CONSERVATION

distribution is a critically important first step in

implementing any landscape level conservation

programmes. Gradually, over the years, one can build

up the capacity and resources to try to meet

objectives (2) and (3) in priority conservation areas.

I emphasise that if the goal is to reliably

estimate parameters such as tiger densities,

survival and recruitment rates, there is no escape

from implementing advanced population

estimation methods. However, these methods can

be employed only where necessary skills and

resources are available. They cannot be applied

for routine population surveys over large regions.

However, the two most critical needs of tiger

monitoring in the Indian subcontinent, mapping

spatial distributions over large regions and

determining population trends in specific reserves

through indices, are widely attainable goals using

relatively simple methods (Karanth and Nichols

2002b). ’

Refer

Anon. (1997): Guidelines for estimating tigers. Letter

dated 28-04-1997 from Director, Project Tiger,

Government of India to Field Directors of Project

Tiger Reserves.

Brander, A.A.D. (1923): Wild animals in Central India.

Arnold, London, UK. 296 pp.

Buckland, S.T., D.R. Anderson, K.P. Burnham & J.L.

Laake (1993). Distance sampling: Estimating

abundance of biological populations. Chapman and

Hall, New York, USA. 446 pp.

Buckland, S.T., D.R. Anderson, K.P. Burnham, J.L.

Laake, D.L. Borchers & L. Thomas (2001):

Introduction to Distance Sampling. Oxford University

Press, Oxford, UK. 432 pp.

Caughley, G. (1994): Directions in conservation biology.

J. Anim. Ecol. 63: 215-44.

Champion, F.W. (1929): Tiger tracks. J. Bombay nat.

Hist. Soc. 33: 284-87.

Choudhury, S.R. ( 1 970): Let us count our tigers. Cheetal

14: 41-51.

Choudhury, S.R. (1972): Tiger census in India. Cheetal

15: 67-84.

Chundawat, R.S., N. Gogate & A.J.T. Johnsingh (1999):

Tigers in Panna: Preliminary results from an Indian

tropical dry forest. Pp. 123-129. In: Riding the tiger:

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Acknowledgements

I acknowledge the long-term support for

my tiger research and conservation activities by

the Wildlife Conservation Society (WCS), the

Ministry of Environment and Forests, Directorate

of Project Tiger and several State Forest

Departments. I am also indebted to the following

agencies which funded my work over the years:

U.S. Fish and Wildlife Service (Division of

International Conservation); Save the Tiger Fund,

of the National Fish and Wildlife Foundation; and

Global Tiger Patrol.

On the subject of tigers, I owe intellectual

debts to George Schaller, John Eisenberg, John

Seidensticker, James Nichols, Melvin Sunquist

and Fiona Sunquist. My fieldwork has been ably

supported over the years by K.M. Chinnappa,

N. Samba Kumar, Praveen Bhargav and D. V. Girish.

I am grateful to all these colleagues.

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BEAR CONSERVATION IN INDIA

( With four text-figures)

A.J.T. JOHNSINGH1

Key words: Himalaya, northeastern India, Ursus arctos isabellinus , U. thibetanus,

Helarctos malayanus, Melursus ursinus. Transfrontier Reserve, poaching, shifting cultivation

Of the eight bear species currently existing in the world, four are found in India: brown bear Ursus

arctos isabellinus , Asiatic black bear U. thibetanus , sun bear Helarctos malayanus and sloth bear

Melursus ursinus. The abundance of each species in India is determined by habitat type, its

location within the global distributional range of the species, its biology, the quantum and continuity

of habitat available, and the anthropogenic pressures it faces. All species are in Schedule-I of the

Indian Wildlife (Protection) Act 1972, the highest legal protection status.

India marks the southeastern end of the brown bear’s distribution range; the black bear in India

occupies a large portion of the southwestern part of the species’ range, sun bear the western end,

and sloth bear most of the species’ range. Populations at the periphery of the range of the species

are usually more exposed to pressures, and have greater chances of going extinct.

The habitat available to the largely herbivorous, hibernation-dependent brown bear in the Trans-

and Greater Himalaya is probably limited to c. 10,000 sq. km, and is disturbed by anthropogenic

pressures such as grazing and medicinal plant collection. There are about 23 protected areas in the

range of the brown bear, within which there are around 300 animals. The omnivorous black bear has

a vast forested habitat in the Outer and Greater Himalaya and northeastern India (c. 300,000

sq. km), with at least 56 protected areas and a minimum of 3,000 animals. Black bears in the higher

reaches of the Greater Himalayan ranges hibernate, while those in the Outer Himalaya and northeastern

India, where they range as low as 500 m, do not. Although disturbed, black bear habitat in the

Himalaya is still continuous and productive as a result of orchards and croplands. Poaching for gall

bladder, fat, meat and skin is a threat to the species. The sun bear is extremely rare in India, and is

confined to the forests along the Myanmar border. Hunting, shifting cultivation and capture of

young for trade, after killing the mother, are the threats across the species’ range. The

myrmecophagous sloth bear still has a vast habitat (250,000 sq. km), harbouring a minimum of

10,000 bears. Yet it is threatened by habitat degradation and fragmentation, poaching for gall

bladder, and capture of young to be trained as performing bears.

Long-term conservation of the brown bear in the Himalaya is possible only with: (i) cessation of

hostilities between India and Pakistan, leading to the formation of a Transfrontier Reserve protecting

the bear habitat, (ii) a total ban on the capture of young for bear baiting with dogs, and (iii) ensuring

that the protected areas are not unduly disturbed between May and October. Protection against

poaching and capture of cubs is a must for the conservation of the black bear and the sloth bear.

Weaning the people in the former range of the sun bear from shifting cultivation, and protection

from poaching, may revive the habitat and population of the sun bear.

Introduction

India is unique in having four of the eight

extant species of bear. They are the brown bear

Ursus arctos isabellinus , the Asiatic black bear

'Wildlife Institute of India, P.O. Box No. 18,

Chandrabani, Dehra Dun 248 001, Uttaranchal, India.

Email: [email protected]

U. thibetanus , the sun bear Helarctos malayanus ,

and the sloth bear Melursus ursinus. This

uniqueness is due to India’s geographic location

at the junction of the Palaearctic and Indo-

Malayan biogeographic zones, which enabled

Palaearctic species (brown and black bear) and

an Indo-Malayan species (sun bear) to range into

the Indian subcontinent. The sloth bear probably

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radiated from the ancestral stock of brown bear

during the mid-Pliocene (Kurten 1968), and

evolved within the Indian subtropical region.

The brown bear in Asia is the same species

as the North American grizzly, but the populations

inhabiting the Himalaya and the northern

mountain ranges of the Tien Shan and Altai have

been ascribed to a separate subspecies. This is

due to its smaller average size, generally whitish

claws, and the guard hairs often being pale at the

tip. The Asiatic black bear is considered the

ecological equivalent of the American black bear

( Ursus americanus) in terms of body form and

habits (Roberts 1997, Schaller 1977). The sun bear

is the smallest of the eight bear species, and the

only one inhabiting lowland tropical rainforests

throughout much of southeast Asia (Servheen

1999a).

Bear Habitats in India

Mainland India has been categorised into

nine biogeographical zones (Rodgers and Panwar

1988), eight of which are important for bears. The

vegetation descriptions given below are from

Champion and Seth (1968).

The northernmost Palaearctic zone, the

Trans-Himalaya, lies north of the main Himalayan

range in Jammu and Kashmir, and within the dry

inner Himalayan valleys of Himachal Pradesh.

Vegetation here consists primarily of dry steppe

and alpine scrub, dominated by xerophytic plants

that reflect the extremely dry and cold climatic

conditions. Typical throughout the region are the

streamside Salix , Populus and Hippophae

associations, and Juniperus and Betula woodlands

on mountain slopes, primarily in Himachal Pradesh.

The brown bear occasionally visits these areas.

The Himalayan Zone includes the

mountains of southwest Jammu and Kashmir,

Himachal Pradesh, Uttaranchal, Sikkim, West

Bengal and Arunachal Pradesh. It encompasses

a complex topography, and a mixture of vegetation

formations with distinctive altitudinal zonation.

Dominant forest types in the northwestern

Himalaya include the chir pine ( Pinus roxburghii)

forests in the Siwalik hills; subtropical evergreen

forests up to 1 ,000 m above msl, characterised by

Persea odoratissima , Olea glandulifera and

Syzygium operculatum\ moist temperate oak

(e.g. Quercus incana) and conifer (P roxburghii)

mixed formations; and mixed conifer and conifer-

deciduous associations with Abies , Cedrus ,

Taxus , Acer and Betula. Between 2,900 and

3.500 m, dry temperate forests dominated by

Pinus wallichiana replace the moist temperate

vegetation. Beyond this, subalpine patches of

Betula and Juniperus grade into shrublands of

Berberis, Lonicera and Rhododendron , and

finally scrub and herb up to c. 5,000 m.

The central and eastern section of the

Himalayan zone is more strongly affected by the

monsoon rains, and the vegetation is both denser

and richer. The tropical semi-evergreen forests

below 1,000 m above msl, are characterised by

Phoebe hainesiana, P. lanceolata, Terminalia

myriocarpa, Bischofia javanica and Schima

wallichii. Above this, are broad-leaved forests

up to 2,000 m, with various Quercus species. Sub-

alpine forests, beginning at elevations >3000 m,

are dominated by Pinus , Betula , Rhododendron

and Acer species up to 4,500 m, where they

give way to sparse scrub extending as high as

5.500 m on southern aspects. The higher altitudes

(>3,000 m) of these habitats are used by brown

bear in Sikkim and middle and lower altitudes (800-

2.500 m) by black bear. Below 1 ,000 m, black bear

and sloth bear habitats may overlap.

The North-East India Zone includes the

Brahmaputra valley ( Shorea assamica,

Cephalanthus cetrandra , Glochidion hirsutum

and Phragmites karka) and Assam hills {Shorea

assamica, Dipterocarpus macrocarpus , Amoora

wallichii , Mesua ferrea and Dendrocalamus

hamiltonii). These areas include the habitat of

the black bear, sloth bear and sun bear.

The semi-arid tracts of Gujarat {Tectona

grandis , Capparis decidua , Carissa carandas and

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BEAR CONSERVATION IN INDIA

Zizyphus nummularia). Upper and Lower Gangetic

plain (Shorea robusta , Syzygium cum ini, Zizyphus

mauritiana, Glycosmis pentaphylla ), Deccan

peninsula ( Tectona grandis, Madhuca latifolia,

Diospyros melanoxylon, Z. mauritiana and Cassia

fistula), and the Western Ghats ( Artocarpus

heterophylla, Mangifera indica, Cullenia excelsa,

Macaranga peltata, Hopea parviflora , Mesua

ferrea, Ochlandra travancorica, Buchanania

lanzan, Phoenix humilis, Z. mauritiana and

Z oenoplia) are used only by the sloth bear.

Status of Bear Species

in the Indian Subcontinent

Brown bear

The southern limit of the brown bear in Asia

is the Himalaya, where it is largely confined to

rolling uplands and alpine meadows above the

timberline, thus ecologically separated from the

forest-dwelling black bear (Schaller 1977; Fig. 1).

The brown bear was once abundant in the

Himalaya, with Kinloch (1 892) having seen 28 in

one day. The brown bear has become rare in

Pakistan, and summarising the survey results of

Choudhry and Farooq (1995), T.J. Roberts (pers.

comm.) estimates about 40-50 brown bears in

Pakistan. Although Gee (1967b) reported the

shooting of a brown bear in Bhutan, there is no

report of brown bear in Bhutan in recent years.

Brown bear occurs in Upper Mustang in Nepal

(Anon. 1994a).

Sathyakumar (1999) gives an excellent

review of the occurrence of brown bear in India

where the populations are largely confined to the

western and northwestern Himalayan ranges in

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BEAR CONSERVATION IN INDIA

the states of Jammu and Kashmir, Himachal

Pradesh and Uttaranchal (Fig. 1). A small

population, possibly continuous with the

population in Tibet, occurs in Sikkim. The

subspecies in Tibet has been identified as Ur sinus

arctos pruinosus (Schaller 1977, Mallon 1985),

which may range into Sikkim and Nepal. Brown

bears may range into the alpine regions of the

Eastern Himalaya (Arunachal Pradesh), but no

confirmed reports are available. There are possibly

around 300 brown bears in India.

Black bear

In Asia, the distribution range of black bear

from west to east, once extended through Iran,

Afghanistan, north and northwestern Pakistan,

the higher altitudes of Nepal, Bhutan and the

Indian Himalaya, forested tracts of Myanmar,

Thailand, Indo-China, southern and northeastern

China, Taiwan, Far-eastern Russia and Japan

(Roberts 1997, Schaller 1977, Servheen 1999a;

Fig. 2). The habitat available to the black bear in

the Outer and Greater Himalaya and northeast

India could be about 300,000 sq. km. Although

disturbed, the black bear habitat in the Himalaya

is still relatively continuous and productive, as a

result of orchards and croplands. In India, black

bears have been reported to occur in 56 protected

areas covering 1 8,340 sq. km in the states of Jammu

and Kashmir, Himachal Pradesh, Uttaranchal,

Sikkim, West Bengal, Arunachal Pradesh,

Meghalaya, Mizoram and Tripura. Protected areas

with black bears range in size from 1 sq. km to

2,237 sq. km (an average of 330 sq. km). However,

information about black bear status is scanty in

most areas, and there have been no reports from

Manipur and Nagaland (Sathyakumar 1999).

Choudhury (1997) reports that black bears are

found in the hills of northeast India, with a few

records from the plains. However, during a four-

month study on elephant-human conflict in Garo

hills in Meghalaya in 1995, Christy Williams (pers.

Fig. 2: Distribution of Asiatic black bear in the southern part of its range (After Servheen et al. 1999b)

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BEAR CONSERVATION IN INDIA

comm.) did not find any evidence of black bear.

Similarly, T.R. Shankar Raman (pers. comm.) did

not find any evidence of black bear during his 120

days of fieldwork in Dampa Tiger Reserve in

Mizoram in 1 994- 1 995 . There are probably at least

3,000 black bears in India.

Sloth bear

The sloth bear is restricted to the Indian

subcontinent: India, Sri Lanka, Nepal and

Bangladesh. In all these countries, sloth bear

distribution has shrunk and become patchy over

the decades (Fig. 3). In Sri Lanka, a population of

about 400 is found largely in the northern and

eastern lowlands (Santiapillai and Santiapillai

1990). In Nepal, sloth bears are largely confined

to the lowland terai protected areas such as

Chitwan-Parsa and Bardia. Chitwan has the

highest density, and density in Bardia is reported

to be lower, as it does not have extensive alluvial

grasslands like Chitwan. East of Chitwan, the range

extends just short of Kosi Tappu Wildlife Reserve.

No evidence of sloth bear was seen in Royal

Suklaphanta Wildlife Reserve in 1 993-94 and sloth

bear numbers in Nepal are probably less than 500

(Garshelis et al. 1999). Once sloth bear occurred

in Suklaphanta (Spillet and Tamang 1967); Cliff

Rice {pers. comm. ) took a picture of a bear feeding

in a burnt terai grassland in December 1976. It is

difficult to understand why the sloth bear has

Fig. 3: Sloth bear distribution in the Indian subcontinent

(After Yoganand, K., C.G. Rice and A.J.T. Johnsingh)

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become extinct in Suklaphanta, which has

extensive alluvial grasslands supporting the best

population (c. 2,000 of the total 5,000) of swamp

deer Cervus duvaucelii duvaucelii (Gopal 1 995).

A recent report by the WWF-Nepal Program

(2002), however, indicates that “there may be only

a few individuals in and around the Reserve.”

Sloth bears may still occur in the remnant, Mixed

Evergreen Forests of Bangladesh. One sighting

of a sloth bear with a cub in Dampa Tiger Reserve

in December 1994 (T.R. Shankar Raman, pers.

comm.), and evidence in Balpakram National Park

(Christy Williams, pers. comm.) have been

reported.

In India, a population of about 10,000 sloth

bears still occurs, widely distributed over a habitat

of 250,000 sq. km, although the habitat is

discontinuous and highly disturbed in most places.

The sloth bear ranges throughout peninsular

India, up north to the foothills of the Himalaya,

inhabiting most low altitude, non-arid areas where

forest cover still remains. The forests of Western

Ghats and the Central Indian Highlands are

currently the two strongholds of the sloth bear

(Yoganand et al., unpubl.).

Sun bear

Servheen (1999b) summarises the historic

range and current distribution of the sun bear. Of

interest are the historic records of sun bears in

places like eastern Tibet and Sichuan, China

(Lydekker 1906), Manipur and Assam (Higgins

1932) and the upper Chindwin district in present

day Myanmar (Wroughton 1916), places where

the species is now extinct. The sun bear is now

found in Southeast Asia from Myanmar eastward

through Thailand, Indo-China and Malaysia. It is

also found on the islands of Sumatra and Borneo.

Although Ullas Karanth (pers. comm.) has

obtained a camera trap photograph of the species,

in Namdapha Tiger Reserve in Arunachal Pradesh

(Fig. 4), in the late 1990s, its occurrence in India is

very rare, and probably confined to the forests

along the India-Myanmar border.

Behavioural Ecology of Bears

Relevant for Conservation

The review on brown bear is largely based

on Roberts ( 1 997) and Schaller ( 1 977). The brown

bear inhabits high alpine mountain slopes and

Fig. 4: Range (south of Brahmaputra) of sun bear in India

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BEAR CONSERVATION IN INDIA

valleys above the tree line, from about 3,000 m to

5,000 m. It generally avoids coniferous forests,

although it will descend to cultivated valleys in

late summer to raid crops and trees in fruit. It is

shy and fearful of man, avoiding upland valleys

occupied by nomadic herders. It has rather poor

eyesight and hearing, but an extremely acute sense

of smell. It locates by smell and even digs out

high altitude Alticola (voles) from their burrows.

The brown bear is skilled in catching fish, and

alpine torrents on the Deosai in Pakistan that have

snow trout ( Dipticus maculatus) are favourite

hunting grounds for the bears. However, analysis

of bear scats and three years of research in the

Deosai have shown that 90% of their diet is made

up of vegetable matter, comprising various

sedges, grasses and especially starchy rhizomes

and bulbs of Carex (sedges), Juncus (rushes) and

Eremurus (foxtail lilies). Only 2 of 70 faecal

samples collected contained rodent or fish bones

(Anon. 1994b). Occasionally, an individual learns

to kill domestic stock, and such animals become

notorious, killing up to 30 sheep and goats in one

night if they get into a penned enclosure (Kruuk

1972). In early summer, brown bears dig

assiduously on grassy slopes in search of

succulent rhizomes and tubers, and turn over rocks

for insects and Crustacea lurking beneath. They

are strongly attracted to carrion and have been

observed feeding on the remains of ibex ( Capra

ibex sibirica), which commonly get killed in winter

avalanches. In Deosai, the radio collared bears

frequently foraged over a distance of 5 km during

the course of one day. Usually they forage only

for a few hours early in the morning and in the

evening, but when human disturbance is high,

they forage only at night.

Mating in brown bear takes place from late

June to end July, and there is delayed implantation

of the fertilised ova. The bears go into hibernation

in October. During hibernation, they frequently

emerge above ground in warmer weather. Usually

two cubs are produced during hibernation, and

they remain dependent on their mother for at least

two years, often accompanying her for up to three

years. Females generally start breeding at the age

of five and thereafter breed once in three years.

Adult males, which can be cannibalistic, are

avoided by a female with cubs.

Although the Asiatic black bear is an

omnivore, a major part of its diet is made up of

fruits, which often need to be collected from trees.

As an adaptation for climbing, it has long claws

on all paws. Other food items include leaf material,

insects and other animal matter in smaller

proportions. In areas where the bears hibernate,

their active period is limited to about six to eight

months. During the active period, feeding forms

the major activity. The study of feeding of black

bears in Dachigam National Park in Kashmir, from

early May to early October, by Manjrekar ( 1 989),

showed that fleshy fruits (soft mast) like mulberry

( Morus alba), cherry ( Prunus avium), peach

( P. persica) and raspberry ( Rubus niveus ), rich in

sugar and carbohydrates, contributed

significantly to the diet in summer. Schaller ( 1 969)

studied the feeding of black bears in Dachigam

for a short period before their hibernation, from

October 6 to 2 1 . The three major food items during

this period were the fruits of Celtis australis ,

walnut ( Juglans regia) and acorn ( Quercus

robur), all fat rich. Manjrekar (1989) also found

walnuts and acorns in the bears’ autumn (pre-

hibernation) diet.

Mating takes place between June and

August. Bromley (1965) estimated hibernation to

last between 128 and 170 days, starting from early

November. In years of good crop, bears go into

their lair earlier when fat deposition is adequate.

The first to enter the dens are adult females with

young. Other females, both barren and pregnant,

are the next to den, and finally the males. During

hibernation, bears move into lairs which are most

often hollows at the base of a tree, otherwise in

rock caves, under fallen logs or in ground dens

that are dug by the bears sometimes even a few

months before hibernation. Thermal stability,

snow accumulation and lack of human

196

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BEAR CONSERVATION IN INDIA

disturbance are the factors that determine the

selection of den sites (Hazumi and Maruyama

1983, 1986, Reid et al. 1991). There is evidence

that the black bear does not always undergo either

prolonged or deep hibernation, as it will

occasionally emerge to forage even during the

winter months (Roberts 1997). The emaciated

males are the first to emerge, followed by the

barren females and finally the females with cubs.

Bears in the foothills of the Himalaya, northeast

India, Myanmar, Thailand and Indo-China do not

hibernate.

Usually two cubs are bom between mid-

January and mid-February, after a gestation of

over six months (Bromley 1 965, Nowak 1 999). The

cubs are weaned at two to two and a half years,

and they become sexually mature at three or four

years of age. Like all bears, the Asiatic black bear

is usually solitary, and groups of three or more do

not occur often, except as mother-cub

associations, and as feeding associations

especially during periods of clumped food

resource availability. The tiger, leopard and wolf

are the potential predators of black bear cubs in

different parts of their range (Bromley 1 965).

Based on the morphology of strong claws,

jaws and long tongue, it can be concluded that

the sun bear is adapted to climbing trees and

extracting food such as insects, larvae or honey

from cavities (Servheen 1999b). The sun bear is

active at night, usually sleeping and sunbathing

by day on a tree 2-7 m above the ground. Tree

branches are broken or bent to form a nest and

look-out post. The diet is omnivorous and

includes figs and termites ( Prohamitennes

mirabilis) (Lekagul and McNeely 1988,

Fredrickson 200 1 , Siew Te et al. 200 1 ). Fredrickson

(200 1 ) observed the bears showing a preference

for unburnt forests where the density of

P. mirabilis was high. Birth may occur at any time

of the year. Gestation period for nine pregnancies

recorded from zoos varied between 95 and 240

days, evidently because of delayed fertilisation

(Nowak 1999).

The sloth bear exhibits several adaptations

to its subtropical and tropical habitat and diet.

To suit the tropics, it has no underfur; however,

it has a long coat that perhaps helps in defending

it from insect bites and also perhaps to exaggerate

its size to dissuade predators such as leopards

{Panthera pardus) and dholes ( Cuon alpinus )

(Yoganand et al ., unpubl.). The monsoonal

climate of the Indian subcontinent, and the

resultant seasonality of resource availability, may

have acted as selective pressures on the ancestor

of the sloth bear to evolve to its present form, as

suggested by Laurie and Seidensticker (1977).

They also suggested that the sloth bear’s

morphological adaptations (like large powerful

claws) and behavioural adaptations (like the

capability to suck out insects), that were mostly

driven by food hunting, are evolved for hard times

when food is limited. Abundant fruit is limited to

a few months, and there is also annual variability

in production of fruits. The sloth bear has to

subsist on other stable food resources like

termites and ants that are more or less available

year round. Although the sloth bear has diverged

towards a diet composed largely of insects, it

has retained the ability to use a variety of food in

conformation with its omnivorous ancestry.

Estimated sloth bear densities vary from 6 bears/

100 sq. km for a dry habitat like Panna Tiger

Reserve (Yoganand et al. , unpubl.) to 21 bears/

100 sq. km in the much more productive Chitwan

National Park, where ecological density during

the dry season can go up as high as 70 bears/ 100

sq. km (Joshi 1996). The reported adult sex ratio

is 1 : 1 (Laurie and Seidensticker 1 977, Joshi et al.

1999, and Yoganand et al. unpubl.). Mating

generally takes place between May and July, and

the cubs are born between November and

January. Litter size of two is most common (Laurie

and Seidensticker 1977, Joshi 1996, Yoganand et

al. unpubl.). Cubs are born either in a natural

cave or in a den dug by the mother. Females

seclude themselves in dens for 6- 1 0 weeks, hardly

coming out to forage, living on fat reserves and

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197

BEAR CONSERVATION IN INDIA

metabolic water during that period (Joshi 1996,

Yoganand et al. unpubl.).

Cubs are frequently carried on their mother’s

back, until they are about six months of age (Laurie

and Seidensticker 1977, Joshi 1996, Yoganand et

al. unpubl.). This is the cubs’ main defence against

attacks by predators or other bears. Cubs stay

with their mothers for one and a half or two and a

half years, becoming independent just before the

breeding season (Joshi 1996, Joshi et al. 1999).

Thus, females breed at either two- or three-year

intervals. The mother-young unit is the only

permanent social grouping exhibited by the sloth

bear (Eisenberg and Lockhart 1972, Joshi et al.

1999, Yoganand et al. unpubl.). They may also

gather at places with abundant food, such as a

dense patch of fruiting plants (Joshi 1996,

Yoganand et al. unpubl.). Tigers attack, kill and

feed on sloth bears occasionally (Joshi et al. 1 999,

Yoganand et al. unpubl.). Leopards, dholes and

even jackals may harass the bears. The anti-

predator strategy of the sloth bear is either to run

away or respond with a spectacular charge and

stand-up display. Its aggressive behaviour may

be a consequence of not being able to rely on

trees for escape, in a habitat that has predators

capable of climbing trees, making it advantageous

to live in fairly open habitat (K. Yoganand, pers.

comm.). Sloth bears probably consider humans

as predators. At close quarters they react to

human presence as they would to a predator

(Laurie and Seidensticker 1977, Yoganand, pers.

comm.). They roar and run away, or roar and attack

humans, before retreating. Bear attacks on

humans are common throughout the range where

bears and humans co-occur (Garshelis et al. 1 999,

Rajpurohit and Krausman 2000, Yoganand et al.

unpubl.).

Conservation: problems and prospects

The survival of bear species in India is

determined by their abundance, which in turn is

determined by habitat type, location of the Indian

range in the global distributional range of the

species, quantum of habitat available, species’

biology, and the anthropogenic pressures they

face. Populations at the extreme or periphery

(‘edge’) of the range of the species are usually

sparse (Hengeveld and Haeck 1982) and have a

greater chance of going extinct (Beddington et

al. 1976, Lawton 1995). The sun bear occupies

the western end of its range in India. Even in the

past, it had a limited range in northeast India,

confined to the hills south of the River

Brahmaputra (Gee 1967a). The population was

also possibly very small, since only 1 5 of the 1 ,389

bears killed in Assam between 1910 and 1917 by

bounty hunters, were Malayan sun bears (Higgins

1932). Hunting, shifting cultivation or jhuming ,

and capture of young for trade by killing the

mother, are the threats across the species’ range.

A forest subjected to jhuming gets re-colonised

by primates including gibbon, if it is left fallow for

a minimum period of 1 0 years, especially if it has

connectivity with primary forests with a source

population (Gupta and Kumar 1994). But so far no

study has attempted to find out at what age a

previously jhumed forest will become suitable for

the arboreal sun bear. Jhumed forests with a 25

year fallow period have frugivorous and omnivorous

bird species similar to forests left fallow for 1 00 years

(Raman et al. 1998).

As the brown bear in India occurs in the

southeastern end of the species’ distributional

range, it is also limited by the ‘edge effect’

described in the previous paragraph. The major

decimating factors faced by brown bear are

hunting [in the past, e.g., Kinloch (1892) killed 7

of the 28 bears he saw], and capture of cubs by

local tribes like the Qalanders in Pakistan to train

them for the cruel practice of bear baiting with

dogs. A survey by Choudhry and Farooq (1995)

found that Qalanders held no less than 2 1 5 brown

bears in captivity, all trained for bear baiting with

dogs. T.J. Roberts (pers. comm.) reports that

between 35 and 80 brown bear cubs were sold

each year to the Qalanders, and in the process of

securing cubs, often twins, the mother is killed

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BEAR CONSERVATION IN INDIA

and frequently the weaker of the cubs also dies

before reaching adulthood. It is possible that cubs

from India may also be smuggled out for this

nefarious entertainment on which a ban has been

imposed by the Pakistan Government, supported

by WWF-International. But the practice

reportedly continues clandestinely, sustained by

a few rich landowners. Continued border

skirmishes between India and Pakistan would also

affect the population occupying the border areas.

In addition, the habitat of the brown bear is

disturbed by grazing and medicinal plant

collection.

The major problem faced by the black bear,

which in India occupies a large portion of the

southwestern part of the species’ range, is

poaching for its gall bladder, fat, meat and skin.

While the first two are believed to be of medicinal

value, the last is for trophy or ornamental

purposes. Some black bear cubs may also be

smuggled to Pakistan from India, as the survey

by Choudhry and Farooq (1995) recorded about

300 black bears with Qalanders.

When compared with other range countries,

the status of the sloth bear is relatively good in

India, with India covering most of the species’

range. Nevertheless, the sloth bear is threatened

by habitat degradation and fragmentation in the

dry tracts of its range, in the states of Bihar,

Jharkhand, Chhattisgarh, Madhya Pradesh,

Rajasthan, Gujarat, Maharashtra and Andhra

Pradesh; also by poaching for gall bladder, bear-

human conflicts, and stealing of cubs, by killing

the mother, to be trained as performing bears

(Yoganand et al. unpubl.).

Conservation of the brown bear would

require the combined efforts of the Governments

of India and Pakistan, by cessation of hostilities

Refer

Anon. (1994a): Biodiversity Conservation Data Project.

Final report submitted to King Mahendra Trust for

Nature Conservation, Kathmandu, Nepal.

Anon. (1994b): Deosai Brown Bear Project. Final report.

Himalayan Wildlife Project, Islamabad, Pakistan.

and creation of a Transfrontier Reserve covering

the wildlife habitats across the Line of Control. In

addition, all the protected areas in the range of

the brown bear should be freed of disturbances

from May to October, the crucial period for them

to recover from the loss of physical condition

due to denning, and to put on sufficient fat

reserves for the next hibernation. Illegal trade in

bear cubs should be stopped.

To help the sun bear re-occupy its former

range in India, a strict control over poaching, and a

long fallow period (> 25 years) for jhumed forests

will be needed. Landscape planning in the bear range

should include a mosaic of primary forests, >25

year old jhumed forests, and corridors of jhumed

forests of other age classes. Conservation of black

bear and sloth bear would require stringent

measures to control poaching and illegal trade in

bear parts.

Given the magnitude of enormous biotic

pressures and the low level of motivation of the

officers and the staff assigned to the job of

controlling poaching, the long-term future of

brown bear and sun bear in India looks bleak.

However, things have to change for the better,

addressing all the conservation problems listed

above, if the bear species, their habitats and other

fascinating species inhabiting their habitat, are to

be ensured a future in a populous country like

India.

Acknowledgements

Mr. Sunil Banubakode provided information

on the sloth bear in Maharashtra. Mr. M.M. Babu

checked the plants listed for the different habitats

in India. Dr. G.S. Rawat, Dr. S. Sathyakumar, Dr. Nima

Manjrekar and Mr. K. Yoganand read through the

manuscript. All are thanked.

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201

KUDREMUKH NATIONAL PARK, KARNATAKA:

A PROFILE AND A STRATEGY FOR THE FUTURE

( With two plates )

S.A. Hussain1

Key words: Kudremukh, langur, liontailed macaque, mining, Karnataka, Western Ghats

The 600 sq. km Kudremukh National Park straddles the Western Ghats adjoining the districts of

Chikmagalore, Udupi and Dakshina Kannada. Rich deposits of magnetite and haematite in the Park

has led to extraction of iron ore by the Kudremukh Iron Ore Company Limited (KIOCL) since

1980. The vegetation is of Western Tropical Evergreen Forest, with higher slopes and rolling hills

having typical sho la-grass land mosaic. Over 35 species of amphibians, 400+ species of birds and 38

species of mammals occur here. Flagship mammals include the liontailed macaque and a small

isolated hitherto undescribed population of langurs, small in size, dark brown and shy, sharing the

habitat of the liontailed macaque, the Malabar civet, and the brown palm civet. The flagship birds

are the great pied hornbill, Nilgiri wood-pigeon, Wynaad laughingthrush, grey-breasted

laughingthrush, blue-winged parakeet, grey-headed bulbul, white-bellied treepie and white-bellied

blue-flycatcher. The KIOCL mining operations have caused some disturbance in the eastern part of

the Park. KIOCL mining will close down soon and the infrastructure left behind needs to be put to

good use without violating National Park rules. This paper suggests the appropriate course of action

to address this issue.

Introduction

The Western Ghats forests comprising of

Tungabhadra State Forest, the South Bhadra State

Forest of the revenue district of Chikmagalur as

well as the Naravi Reserve Forest and the Andar

Reserve Forest of the revenue districts of

Dakshina Kannada (DK) and Udupi (12°-16° N

latitude) are now collectively designated as the

Kudremukh National Park (KNP), notified on

September 4, 1987 under Section 35(1) of the

Wildlife Protection Act. It is under the control of

Kudremukh Wildlife Division of the Karnataka

State Forest Department. The Division looks after

KNP, Someshwara and Mukambika Wildlife

Sanctuaries in Udupi and DK districts. The

Division Headquarters is located at Karkala Town.

It has a Deputy Conservator of Forest (DCF) as

the head, with three Assistant Conservators of

‘Biodiversity Initiative Trust,

Hussain Manzil, Anekere Road,

Karkala 574 104, coastal Karnataka, India.

Email: [email protected]

Forest (ACF) located at the Headquarters, the

Kudremukh mining township, and the Mukambika

Wildlife Sanctuary, Kundapur Taluk, respectively.

The KNP having a total area of about 600 sq. km

comprises lowland as well as highland Tropical

Evergreen Forests, shola-grassland-savanna and

a mosaic of Mixed Semi-evergreen Forests and

plantations in the peripheral areas (Plate 1 , Fig. 1).

The altitude ranges from 300 m in the lowlands to

l, 892 m at the highest peak of Kudremukh. The

average altitude of the highland hills is about 1 ,000

m. The area receives good rainfall during monsoon

months (June-October; range = 1 ,778-6,350 mm,

highest recorded was 10,000 mm, in 1984).

Precipitation and runoff regime is roughly

defined at the crest of the ghat which also forms

the western and eastern facies of the landscape.

Thus, the watershed west and south of the divide

is very steep, contributing to rapid runoff of short-

span river systems that drain into the Arabian

sea. The watershed east and north of the divide

caters to river systems that meander through

gentler slopes and gullies and flow eastward

202

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, I00(2&3), AUG.-DEC. 2003

KUDREMUKH NATIONAL PARK

Hussain, S.A.: Kudremukh National Park

Plate 1

To Karkaia'

avathi Rive?

HABITATIONS 1

ROAD

RIVER

DENSE FOREST

SHOLA & GRASS LANDS

▲

HILL PEAKS

t Narasimhaparvatha

2. Vaiikunja

3. Gangadika!

4. Kuranjal

5. Kudremukh Peak

6. Kyathanamakki

Fig. 1: Map of Kudremukh National Park

JOURNAL , BOMBAY NATURAL HISTORY SOCIETY, 100(2 &3), AUG.-DEC. 2003

203

KIND COURTESY: DY CONSERVATOR OF FORESTS, WILDLIFE DIVISION, KARKALA

KUDREMUKH NATIONAL PARK

Hussain, S.A.: Kudremukh National Park

Plate 2

Fig. 2: Liontailed macaque Macaca silenus, one of the flagship species

of Kudremukh National Park

Fig. 3: A view of the mining township, from the mining area,

within Kudremukh National Park

O

204

JOURNAL , BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003

PICS: S.A. HUSSAIN

KUDREMUKH NATIONAL PARK

towards the Bay of Bengal. Two major rivers, the

Tunga and the Bhadra originate from the dense

Bhagavati Forest. While Tunga flows due north-

northeast towards Sringeri, Bhadra flows due east

passing through the leasehold of the public sector

enterprise, Kudremukh Iron Ore Company Ltd

(KIOCL) mining area, both join up near Bhadravati

town to form Tungabhadra river, and eventually

merge with River Krishna.

The KNP is very rich in biodiversity. The

vegetation is typical of the tropical evergreen

forests. Though there are inventories and listings

of prominent hardwoods, shrubs and herbs, no

comprehensive data is available on the total plant

species of the entire area.

Over 35 species of amphibians occur here

and their density and diversity have been

recorded. Over 400 species of birds have been

recorded for the KNP area. The KNP hosts 38

species of mammals belonging to 28 genera. Four

species are endemic to the Western Ghats; three

of them being the liontailed macaque Macaca

silenus (rare and endangered; Plate 2, Fig. 2), the

Malabar civet Viverra civettina and the brown

palm civet Paradoxurus jerdoni (Hussain et al.

1999). The fourth is a small isolated population of

langur species that has recently been noticed

sharing the shola habitat with the liontailed

macaque. This langur differs a great deal from the

common langur Semnopithecus entellus but is

closer to the Nilgiri langur Trachypithecus johni.

It is possible that this may be a distinct species of

langur, and more study is needed to ascertain it.

These four are the flagship species of ; ' ; region.

Vegetation Structure

Kadambi (1942a, b) has distinguished 3

types of vegetation structure in the

Poeciloneuron forests of South Bhadra and

Tunga, namely Mixed Evergreen, Semi-pure

Evergreen and Pure Evergreen Forests,

corresponding to increasing concentrations of

Poeciloneuron. Rai (1981) has studied the

structure and primary productivity in South

Bhadra RF (Reserve Forest) corresponding to the

gregarious Poeciloneuron stands.

The extensive lowland forests comprising

Naravi RF and Andar RF and the adjacent

Someshwara Wildlife Sanctuary are included

under the KNP. These forests comprise mainly of

Tropical Evergreen and Moist Deciduous types.

Some areas have been extensively planted with

teak Tectona grandis.

Fauna of Kudremukh National Park

Ornithology

From the distributional ranges given in

available literature, it appears that the total number

of bird species occurring in the region could be

400 ±20. These were based on actual sightings in

the highland/lowland forests as well as the

secondary forests and urban areas of the coastal

belt. The tropical forests of the southern Western

Ghats (including the coastal areas) harbour a

lower diversity of bird species in comparison to

similar habitats in African and Neotropical forests.

Daniels (1984) estimated that in the evergreen

forests of southwestern India, a maximum of 1 50

species of birds are present in an area less than 1 5

sq. bn, whereas 478 species have been recorded

in Ecuador (S. America) and 365 species in Gabon

(Africa). The recent rapid changes in the

landscape may have caused the disappearance

of some species, while some others may have

recently replaced these in the changed habitats.

The flagship species of tropical forest birds

which are either endemic to the southern Western

Ghats or have a patchy distribution of small

populations, are represented in the KNP region

by great pied hornbill Buceros bicornis , Nilgiri

wood-pigeon Columba elphinstonii , Wynaad

laughingthrush Garrulax delesserti , grey-

breasted laugh ingthrush G. jerdoni, blue-winged

parakeet Psittacula columboides, grey-headed

bulbul Pycnonotus priocephalus , white-bellied

treepie Dendrocitta leucogastra , white-bellied

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KUDREMUKH NATIONAL PARK

blue-flycatcher Cyornis pallipes and Nilgiri

flycatcher Eumyias albicaudata. Some other

species such as black eagle Ictinaetus malayensis

and Ceylon frogmouth Batrachostomus moniliger

also occur.

Though the KIOCL mining sector was

devoid of any species, surprisingly small patches

of remnant shola forests within the actual mining

area harboured small populations of habitat

specialists. Kudremukh township with a mosaic

of urban ornamental plantations intermingling with

patches of dense secondary scrub hosted yet

another diverse bird population. Lowland forests

of KNP also hosted a good species assemblage

typical of the area. Two censuses were carried

out, at each site in the KNP by the author, results

of which are given in Table 1 .

Mammals

In the KNP area, endemics are represented

by primates (2 spp.) and small carnivores (2 spp.)

(Table 2). Except the Nilgiri tahr Hemitragus

hylocrius , all other endemics of the Western Ghats

are found here. The KNP hosts perhaps the single

Table 2: Number of genera and species

and endemic mammals in KNP

largest population of the flagship species, the

liontailed macaque, north of Silent Valley in Kerala.

It is estimated that over 200-300 of them occur in

the KNP belt with a few other small populations

scattered in the lowland forests of Someshwara

and Mookambika Wildlife Sanctuaries located

north of KNP.

As mentioned earlier, the distinct langur

population of the shola forests needs further field

Table 1: A quick analysis of the results of the censuses at Kudremukh National Park (including mining area)

area.

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KUDREMUKH NATIONAL PARK

and DNA research to ascertain its specific status.

The other species reported recently is the rare

and endangered Malabar civet ( Viverra civettina),

small populations of which have been frequently

reported from Mookambika and Someshwara

Wildlife Sanctuaries as well as lowland and

highland forests of KNP.

The Indian gaur Bos gaurus is the largest

common ungulate occurring in both lowland and

highland forests of KNP, followed by sambar

Cervus unicolor and chital Axis axis. These are

the most common animals in the area. However,

large-scale forest clearing for agriculture and

human settlements in the past 100 years,

particularly in the lowland forests, has fragmented

their populations into smaller herds, frequently

bringing them into conflict with human

settlements. The populations of their natural

predator, the tiger Panthera tigris has also

reduced due to anthropogenic pressures.

However, in 2001, a tigress took residence very

close to the mining area and raised two cubs. As

everywhere else in the region, the leopard

Panthera pardus is still holding its own,

particularly in the secondary mixed forests and

mosaic vegetation.

It is estimated that over 150 liontailed

macaques are present in the KNP, most of which

are confined to heavy rainfall areas of shola/

grasslands of Bhagavati Forest, Tungabhadra

State Forest (Kerekatte) and steep western slopes

of the ghats and lowland forests of Andar and

Naravi Reserve Forests (more than 200 m above

msl). A total of ten troops were encountered by

the author. These are listed in Table 3.

The occurrence of the unidentified langurs

needs further observation. This population

which may have been overlooked so far by

earlier workers needs to be studied to determine

whether they are a distinct and new species. The

animals are small, with very dark and thick coat

and pale brown head, slow moving in the canopy

and confined to dense highland forests of

KNP.

Table 3: Troops of liontailed macaques

Elephants Elephas maximus are very rare in

KNP, though there have been occasional individuals

straying in the periphery of the Park. In 200 1 , three

individuals came up the pipeline road and stayed

within the Park. This population is more or less

resident now. Several species of bats occur in both

lowland and highland forests.

Resource Evaluation

The Western Ghats of Karnataka possess

some of the most pristine lowland and highland

forests in the entire Western Ghats chain. Though

these forests have gradually been fragmented into

smaller disjointed stretches of Evergreen, Moist

Evergreen, Moist Deciduous, mixed scrub, and

regenerating woodlands, the diversity of

mammalian fauna still holds out on its own in

smaller pockets. The transitional belt of lowland

forests of the coastal zone and highland forests

of steep hill slopes, culminating in the specialised

vegetation of hill crests which then gradually

stretch eastwards into a rolling mosaic of forests

mingled with plantations, host a number of species

including some rare endemics. The rich wildlife of

the area is underscored by the fact that two

national parks and several wildlife sanctuaries

straddle the Western Ghats.

The Kudremukh National Park serves both

eastern and western regions of the Western Ghats.

The major portion of Malnad area and the entire

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207

KUDREMUKH NATIONAL PARK

district of Udupi and Dakshina Kannada depend

on the water regime emanating from the

Kudremukh area. The districts in the immediate

vicinity served are Shimoga, Chikmagalur and

Hassan in the east and Udupi and Dakshina

Kannada in the west. Several perennial and

seasonal streams/ rivulets/ rivers exist in the KNP

area. Among the east flowing rivers, Tunga and

Bhadra and their tributaries Varada and Hagari

have their origin within KNP with a catchment

area of 684.4 sq. km and total length of 528 km.

The two rivers join up after Bhadravati town to

become the Tungabhadra river.

The west flowing, perennial Sita, Suvama,

Udyavar, Mulki, Gurpur and Nethravathi rivers

serve the Udupi and Dakshina Kannada districts

and their total catchment area is over 8,326 sq. km

and total spread area is 833,595 ha. The average

annual rainfall here is 4,420 mm. The extent to

which the villages around KNP depend on these

freshwater sources — in terms of irrigation,

fishing, daily consumption, number of people

served — is given in Table 4. The following are

the crops that depend on the freshwater sources:

A. North, east and southwards: Tea, coffee,

cardamom, pepper, arecanut and paddy in the

immediate surroundings. Paddy, sugarcane,

banana, cereal crops, etc. in the down river basin

and command areas of Tunga and Bhadra dams

and other regions right up to the Bay of Bengal.

B. Westwards: Coconut, rubber, arecanut,

paddy, sugarcane, and other crops.

Human Impact Evaluation

The area has no legally permanent human

settlements, except for a few communities that

have existed here for a long time. Even in the

periphery of the Park, particularly along its eastern

and northern boundaries, the human population

is sparse and widespread among plantations and

small holdings of cultivated fields. Unlike other

national parks and sanctuaries, anthropogenic

pressures on the vegetation for fuel and other

Table 4: The figures showing the dependence of

Dakshina Kannada and Udupi on the rivers

flowing out from KNP

Total agricultural land

Groundwater -

Domestic consumption

Irrigation

Total estimated consumption

Annual replenishment

River outflow -

annual average

Coastal fisheries -

annual average catch

Population of the districts

(Udupi &D.K.)

Density

286.000 ha

43.7 million cu. m/year

427 million cu. m/year

471 million cu. m/year

1 ,245 million cu. m/year

29,561 million cu. m/year

143.000 tonnes

2.69 million (in 1961)

319 per sq. km

Compiled from various sources

needs are minimal in KNP. However, as is

happening in other developing countries of the

tropics, there are many problems faced by the

KNP ecosystem. There have been recent cases of

large-scale encroachment of Park forests in the

periphery by rich landholders for illegal extension

of coffee and other plantations. The KIOCL

township is the only and the most populated

human habitation in the KNP (Plate 2, Fig. 3). The

total human population of the township was

estimated to be 10,636 (1991 Census). Though

actual figures are not available, it is estimated that

350-450 privately owned cattle (mostly cows and

a few buffaloes) exist in the Park area. A list of the

enclosures in the Park is given in Table 5.

Encroachment of forest land by marginal farmers,

large plantations, as well as cattle graziers is

becoming a serious problem.

The major activity in the Park ecosystem

is of iron ore mining by the public sector

Kudremukh Iron Ore Company Ltd (KIOCL).

The company has been operating here since

1980 in the leased area. Most of the actual

mining is in grass topped bare hills (370 ha)

with scattered small sholas.

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KUDREMUKH NATIONAL PARK

Table 5: Details of occupied land within

Kudremukh National Park

a:

Range No. of No. of Total land

enclosures families holding

(in ha)

b:

1 . Total extent of land leased to KIOCL

2. Extent of forest land leased

3. Extent of forest land converted

inside leased area

4. Extent of forest land converted for

pipeline outside the leased area

5. Extent of forest land likely to

submerge outside the leased area

due to raising of dam height

6. Total forest land being utilised by

KIOCL

4,605.02 ha

3,203.55 ha

1.452.74 ha

30.00 ha

340.00 ha

1.822.74 ha

Areas that have some level of threat to wildlife

and its habitat

Of the entire 600 sq. km area of the

Kudremukh National Park (KNP), some level of

threat to wildlife and its habitat is present at its

extreme eastern border where the KIOCL has its

operations. The mining concession and its

surroundings are a hilly terrain, mostly covered

with grasslands and small stunted shola patches.

These grasslands were originally the traditional

grazing (gomal) lands for the cattle of the villages

nearby. In fact, right in the Park premises there are

a couple of private holdings rearing about 200-

300 cattle which freely graze around.

On the whole, the mining concession with

its establishment, mines, processing area and

township is located very close to the eastern and

traditionally exploited border of the Park. Tea,

coffee, cardamom, pepper and areca plantations

as well as paddy cultivation are the major

agricultural activities in this area. Due to mosaic

vegetation (some good forest patches and tree

cover interspersed with villages and cultivation),

there is a good bird and insect diversity, with

some wild animals such as wild boar, sambar,

barking deer and other smaller mammals moving

in and out. These have adapted well to local

conditions.

The major impact of mining is on the

Bhadra river system which passes through the

immediate vicinity of mining fields and is

subjected to a heavy silt load, particularly during

the first onslaught of monsoon rains in June-

August. Incidentally, this is the period when most

of the hill stream fish, amphibians and other

microfauna of the terrestrial system begin to

regenerate and a very critical period for the

biodiversity of not only local but also major

downstream river systems. The main grouse of

the villagers and farmers downstream is that the

silt load renders the water unfit for drinking and/

or cultivation, particularly in the monsoon period.

Cumulative effect over the years may have far

reaching consequences in the Bhadra river

system.

Heavy grazing by village cattle, ill-advised

plantation on true grasslands in the KIOCL lease

area as well as other grasslands in the Park with

exotic Acacia auriculiformes , Eucalyptus ,

Casuarina , etc., and frequent setting of fire to

grasslands without any scientific study on

regeneration profile (which has led to colonisation

of grasslands by weed-like fern species, depleting

the grass available as food for wildlife, particularly

gaur and sambar), are some of the major problems

faced here.

Fortunately, the bulk of the dense pristine,

Tropical Wet Evergreen and the Shola Forests as

well as the grasslands are well away from the

KIOCL and preserved by the sheer hilly terrain.

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KUDREMUKH NATIONAL PARK

Except for the road from Mangalore to KIOCL,

the area is comparatively free of human presence.

The core area of the KNP is dense and well

preserved due to its inaccessibility'. This area,

which comprises the central and western portion

of the KNP, is home to the liontailed macaque,

gaur, sambar, and many other smaller mammals,

amphibians, reptiles and birds. Some scattered,

old, legal settlements are present in this area but

these are not very obvious, and as such there is

not much disturbance to wildlife. These

settlements need to be taken into account in the

management plan of KNP as a long-term policy. A

newly created national highway passes through

the dense forests of the western region of KNP.

Some strict measures have to be taken to see that

no widening of the road and/or illegal

encroachments take place here.

The creation of a separate Kudremukh

Wildlife Division with its headquarters nearby is

a major step taken by the State Government.

However, like many of the government

departments, this division is under-equipped and

under-staffed with meagre budget allotments. The

Division controls the gates, carries on fire

protection measures, has established and

maintains medicinal plant preservation plots in

the Park besides a nature camp with visitor

facilities, and carries out anti-poaching patrolling.

The current DCF-Wildlife is a dedicated

conservationist with a no-nonsense approach to

forest protection. However, a scientifically

designed management plan for the KNP has not

yet been prepared and there is an immediate need

for this. Local conservationists and experts

should be involved by the Kudremukh Wildlife

Division to prepare a comprehensive management

plan for the KNP.

KIOCL must be made to pay full

compensation and royalty to the Wildlife

Department for the exploitation they have carried

out so far and are continuing. This royalty could

be given out of the profits the KIOCL earns from

exploiting the resources. This could be anywhere

between 8-10 crores per annum and the amount

can be utilised for research, conservation and

maintenance expenses of the KNP. This is only

the compensation for past and existing

exploitation, and should be considered as a

penalty. New areas should never be given to

KIOCL for mining.

Possible Future Course of Action

Rationale

Sooner or later, the KIOCL may have to close

down the mining operations at Kudremukh and pull

out their establishment from the area. This will

happen either due to pressure from environmental

concerns and the Supreme Court directives or by

the sheer economics of mining in the area. For one,

the deposits will be exhausted very soon and there

is very little chance of permission being given for

additional mining concessions in either nearby

Nellibeedu or at Gangrikal which is in the ‘sanctum

sanctorum’ of the KNP.

The township (area 320 ha, population

c. 1 0,000 in 1 99 1 ) currently maintained by KIOCL

has all the modem facilities including water supply

and sewage treatment/disposal (see Appendices

I and II). The layout of the town is well designed

in an undulating landscape. Most of the original

shola/grassland mosaic remains untouched

between various housing sectors, giving it a

natural look. An indicator of the naturalness is

the presence of a large number of birds of Tropical

Evergreen Forests, reptiles (including king cobra),

and occasional barking deer, sambar, gaur, and

giant squirrels that frequent the periphery of the

township.

The town also has all the modem amenities

such as STD/I SD, Cable TV network, a well

maintained public park, half-hourly bus services

(greater part travelled through the National

Highway to Mangalore, c. 85 km, 2 hour drive) to

the nearest airport/railway station.

Inevitably, KIOCL will have to surrender

the mining lease and the land will have to be

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KUDREMUKH NATIONAL PARK

reverted and added to the National Park. All

the infrastructure that lies within the township

and the mining area needs to be dealt with

somehow.

For the KIOCL there are two choices:

a. Demolish all structures and establishments in

the township and mined area (and leave behind

a ghastly concrete/steel rubble of a ghost

town), or

b. Hand over the main structures to an

appropriate agency (National Park Autho-

rity) for possible future action/usage along

with substantial seed money to deal

with it.

Considering the fact that the area will

legally be reverted back to a National Park and as

such a full-scale independent township cannot

exist within the Park, private commercial activities

cannot be allowed, nor can there be any chance

of promoting a holiday resort or a hill station for

tourist activities. Perhaps the only pragmatic way

to overcome this dilemma is to utilise the existing

infrastructure to establish some kind of regional/

national research/convention centre. The

following possibilities can be seriously

considered after setting up an appropriate

taskforce with a mandate to devise ways and

means to achieve the desired and pragmatic

objectives.

Refer

Daniels, R.J.R. (1984): Bird Communities of Uttara

Kannada. Ph.D. Thesis. Submitted to Indian

Institute of Science, Bangalore.

Hussain, S.A., S.V. Krishnamurthy, N.R. Rao &

S. Nagraj (1999): An Overview of Some

Aspects of the Biodiversity of the Kudremukh

National Park, Karnataka. Pp. 181-201. In:

Biodiversity of the Western Ghats Complex

of Karnataka (Eds.: Hussain, S.A. & K.P.

Achar). Biodiversity Initiative Trust,

Recommended course of action

Constitute a taskforce with a definite

mandate and time frame to:

a. Conduct a preliminary feasibility study and

report,

b. Negotiate with KIOCL and all the likely

stakeholders,

c. Prepare an appropriate Action Plan,

d. Source out funding regimes (including income

generating schemes - e.g. facilities can be hired

out as an intemational/national environment,

scientific and conservation related conference

and convention centre).

The possible nodal agencies to be involved

are given in Appendix III.

Some immediate recommendations

1. Stop all “afforestation schemes” and avoid

plantations on either grasslands or deforested

areas. The tropical rainforest is capable of

regenerating itself provided it is given

adequate protection.

2. Monitor the growth of pioneer species in open

areas and control the spread of any alien weeds.

3. Carry on experiments in the broken up mining

areas to find the best way to encourage re-

colonisation of pioneering species such as

Buddlija asiatica, Blumea oxydonta and

Crotalaria pallida.

•NCES

Mangalore, India.

Kadambi, K. (1942a): The Evergreen Ghat Rain Forests

of the Tunga and Bhadra River Sources (Part-I).

Indian Forester 68(5): 233-240.

Kadambi, K. (1942b): The Evergreen Ghat Rain Forests

of the Tunga and Bhadra River Sources (Part-II).

Indian Forester 68(6): 305-132.

Rai, S.N. (1981): Productivity of Tropical Rain Forests

of Karnataka. Ph.D. Thesis. Submitted to Mumbai

University, Mumbai

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211

KUDREMUKH NATIONAL PARK

Appendix I: Existing facilities at KIOCL township

Permanent structures (other than mining area):

Data compiled from different sources.

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KUDREMUKH NATIONAL PARK

Appendix III: Possible agencies to be involved in deciding the future of KIOCL

Nodal agencies : Ministry of Environment & Forests, Government of India (MoEF/GOI)

Karnataka State Ministry of Environment & Forests.

Stakeholders: MoEF/GOI:

Zoological Survey of India

Botanical Survey of India

Forest Research Institute, Dehra Dun

Wildlife Institute of India, Dehra Dun

Department of Science & Technology (DST)

Indian Council for Agricultural Research (ICAR)

University Grants Commission (UGC)

Any other appropriate agency

Karnataka:

State Wildlife Department

State Forestry College

University of Agricultural Sciences (UAS), Bangalore

Centre for Ecological Studies, Indian Institute of Science (CES/IISc)

Indian Remote Sensing Agency (IRSA)

Indian Space Research Organisation (ISRO), Bangalore

Kuvempu University, Shankarghatta, Shimoga

Any other appropriate agency

NGO/lnstitutions:

Bombay Natural History Society (BNHS), Mumbai

Salim Ali Center for Ornithology & Natural History (SACON), Coimbatore.

Any other national agency

Funding sources:

National:

Government of India - MoEF

Ministry of Science, Technology & Space

Ministry of Agriculture

Government of Karnataka - MoEF

International:

United Nations Development Program (UNDP)

United Nations Environment Program (UNEP)

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213

DEVELOPING RESPONSIVE INDICATORS

FOR THE INDIAN BIOSPHERE RESERVE PROGRAMME

Anirban Ganguly1, Yogesh Gokhale2’3 and Madhav Gadgil2-4

Key words: Biosphere Reserves, India, indicators, monitoring and evaluation,

community participation

Biosphere Reserves (BRs) are intended to reconcile challenges relating to conservation and sustainable

use of biodiversity, social and economic development and maintenance of associated cultural values.

In keeping with these objectives, India has established a network of 1 1 Biosphere Reserves. The

study attempts to evaluate the performance of these BRs using a set of indicators relating to

community participation, legal and institutional mechanisms, management capacity and effectiveness.

The specific indicators relate to the Indian context and give special attention to issues like people-

wildlife conflict, understanding of programme objectives at various hierarchical levels, continuance

of traditional conservation practices and promotion of scientific research. Indian BRs have, by and

large, failed to resolve or even added to resource conflicts due to inter agency disputes or imposition

of an inappropriate model of development. Moreover, major management decisions seem to be

taken at higher bureaucratic levels without reference to livelihood concerns of local people and

traditional resource management systems followed in local areas. On the other hand, Indian BRs

have been successful in areas like supplementary income generation. The study also points to a

methodology of using ‘discriminatory’ performance indicators which would be adequately sensitive

to the proximate needs of ecologically handicapped communities.

Introduction

Biosphere Reserves are intended to

reconcile challenges relating to conservation and

sustainable use of biodiversity, social and

economic development and maintenance of

associated cultural values (UNESCO 1996). In

keeping with these objectives, India has

established a network of eleven Biosphere

Reserves till October 1999. Out of these eleven,

only three — Nilgiri, Gulf of Mannar and

Sundarban — are recognised under UNESCO’s

Man and Biosphere programme as of May 17,

2002 (http://www.unesco.org/mab/brlist.htm). The

present study attempts to evaluate the

performance of these Reserves, using a set of

indicators relating to community involvement and

participation, legal and regulatory mechanisms,

'Tata Energy Research Institute, Darbari Seth Block,

Habitat Place, Lodhi Road, New Delhi 110 003, India.

2Centre for Ecological Sciences, Indian Institute of

Science, Bangalore 560 012, Karnataka, India.

Email: [email protected]

management capacity and effectiveness, drawing

from a number of case studies.

The idea of Biosphere Reserves was mooted

by UNESCO in 1973-74 under its Man and

Biosphere (MAB) programme. Biosphere Reserve

(BR) is an international designation coined by

UNESCO for representative parts of natural and

cultural landscapes extending over terrestrial or

coastal/marine ecosystems. In India, BR is not a

legal conservation category, unlike Wildlife

Sanctuaries and National Parks. However, areas

earmarked as BRs often overlap with areas which

are legally protected, often resulting in conflicting

plans and programmes.

Material and Methods

Field experience

One of the authors (MG) has been associated

with the Indian Biosphere Reserve Programme

since its early beginnings in 1 980. At that time, he

was commissioned by the Government of India to

prepare the project document for the

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THE INDIAN BIOSPHERE RESERVE PROGRAMME

establishment of the Nilgiri BR. This subsequently

became the first biosphere reserve to be

established in 1986. The Centre for Ecological

Sciences established a Field Research Station in

the Nilgiri BR in that year and has been continually

involved in monitoring the functioning of that BR

since its inception. In 1992, the Ministry of

Environment and Forests (MoEF), Government

of India commissioned the Centre for Ecological

Sciences (CES), Indian Institute of Science,

Bangalore in 1 992 to undertake a mid-term review

of the Biosphere Reserve Programme. As a part

of this exercise, Madhav Gadgil and R. Prabhakar

undertook a specific review of the Nilgiri BR and

Niraj Joshi undertook field visits to the following

Biosphere reserves: Gulf of Mannar, Nandadevi,

Nilgiri, Nokrek, Manas and Sundarban. This mid-

term review was followed in 1993 by a training

programme for the managers of the biosphere

reserves in India at Mudumalai Wildlife Sanctuary

in Nilgiri BR, which provided an opportunity for

obtaining further inputs on the BR programme.

Hans Raj Negi of CES conducted extensive field

research in the Nandadevi BR during the years

1994 to 1997.

Subsequently, the Biosphere Reserve

Programme was once again reviewed by CES in

collaboration with RANWA, an environmental

group, in 1996-98 as a part of the Biodiversity

Conservation Prioritisation Programme (BCPP) of

WWF-India. Under this programme, collaborative

field visits were undertaken in the following

Biosphere Reserves:

The data collection has involved interviews

with forest and other government officials, local

people, researchers, and NGOs working in the area.

Several field visits were conducted for rapid

assessment of the various components of the BR

programmes at various Biosphere Reserves by

one of the authors or by the collaborators

mentioned earlier. Secondary information sources

in the form of reports, newspaper articles and

official documents were also referred to.

Evaluation methodology

Of late, there has been a proliferation of

literature on the evaluation of developmental

interventions. The World Bank, World Resources

Institute, IUCN and Biodiversity Support Program

of WWF, for example, provide detailed guidelines

on the kind of scientific and economic indicators

which may be used to evaluate the performance

of a wide range of projects. These range from

country-specific biodiversity indicators like

percentage of major forest, types covered in a

protected area network, as suggested by IUCN,

to broader region-oriented measures like change

in institutional or management structures leading

to change in resource utilisation practices. These

indicators have been extensively applied to assess

the efficacy of funds provided at the international

level, both by specialised funding mechanisms

like the Global Environment Facility (GEF) and by

bilateral donor organisations and aid agencies like

DANIDA and NORAD. Of late, there is also a

tendency to use indicators sensitive to equity

concerns, specifically from the angles of gender

and socially/economically disadvantaged

communities.

At the same time, there has also been a

major development in the economic and socio-

political theory in the area of cost-benefit analysis

and broader evaluation techniques. The major

landmark in this regard would be the development

of the Guidelines for Project Evaluation by

UNIDO, Vienna (Dasgupta et al. 1 972 ). These

guidelines spell out the economic rationale behind

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THE INDIAN BIOSPHERE RESERVE PROGRAMME

project choice, evaluation and monitoring using

models of uncertainty and social choice. From

then onwards, there has been a rapid growth of

understanding on the valuation of ecosystem

functions (Barbier 1 992, Dixon 1 992) and the social

consequences of disruptions of such functions

(Ruitenbeek 1992, Munasinghe 1990). Also, one

school of economists, pioneered by Amartya Sen,

have emphasised the use of discriminatory

analytical tools like weighted indices of human

development to assess programme level

interventions, since indices which are aggregative

or even relative at an inappropriate scale tend to

suppress major equity and (re)distribution

concerns. In his recent work development as

freedom, Sen (2000) advocates the use of

analytical tools that view development as a

process of expanding substantive freedoms,

reflected in socio-economic arrangements (e.g.

health and educational facilities) and political/civil

rights (e.g. the freedom to participate in public

discussion and scrutiny). Performance evaluation

of development interventions, when informed by

such a view, involves active consideration of the

ability of an intervention to achieve tangible

enhancements in freedoms of the target groups,

such as increased capacities of individuals and

groups to indulge in public debate on issues that

affect their livelihoods and lifestyles.

There is clearly a need to harmonise these

two streams of literature in order to develop a set

of responsive indicators for evaluation of

interventions at project and programme levels.

We make here a distinction between project and

programme levels because project interventions

very often follow from strategies developed at

the programme level. Thus, reasons for success

or failure of a specific project could lie both in

problems intrinsic to the project itself or be the

result of incorrect strategisation or poor learning

at a higher decision-making level which gets

reflected in project implementation phases. In this

paper, our focus is on the use of indicators to

make a programme level evaluation of the

Biosphere Reserve programme, as implemented

in India.

These indicators should, on the one hand,

test the efficiency of financial, technical and

management inputs provided in terms of a set of

carefully designed criteria, and on the other hand

be adaptive enough to ground realities so as not

to miss out relevant equity and social justice

concerns. These concerns are almost always

region and culture specific and, therefore, difficult

to aggregate over projects. For example, increase

in human-tiger conflicts in the Sundarban BR area

— and the resultant agony of local communities

— is now a part of local cultural ethos or even the

mindscape of local communities. It is impossible

to weigh this against reduced conflicts in other

areas or offset positive economic indicators

against the psychological cost of loss of a family

member or living a differently-abled life.

The currently adopted evaluation

methodology could draw from the rich literature

now available in a range of disciplines including

social choice, information and uncertainty theory,

while the theoretical literature can be enriched

through the use of appropriate case studies and

best practice lessons. International funding

mechanisms such as GEF do undertake programme

implementation reviews periodically; such

reviews can also inform and enrich the theoretical

literature in many of these fields. For instance,

case examples of how communities do or do not

exhibit group rationality when involved in a donor

driven programme can be a key input to social

choice literature. In arid areas of Rajasthan, water

harvesting initiatives supported by external

funders have been most successful when local

communities have been mobilised by local NGOs

to bear a significant component of the programme

cost; in other words, the ability of a community to

translate its group decision-making behaviour to

actual burden sharing (financial or otherwise) is a

requisite to the sustained success of a resource

‘wise-use’ programme (Rajender Singh, Tarun

Bharat Sangh, pers. comm.).

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An attempt at harmonisation of these two

streams of literature on evaluation methodology

and Biosphere Reserves is made in the following

paragraphs, drawing from available case studies

and personal experience of the authors with regard

to the Indian Biosphere Reserve Programme.

The Case of Indian Biosphere Reserves

Biosphere Reserves, as mentioned earlier, are

areas of terrestrial and coastal/marine ecosystems

or a combination thereof, earmarked for innovative

conservation and management (an alternative to

the National Park/Wildlife Sanctuary model) and

motivated by the framework of UNESCO’s Man

and Biosphere Programme. Each Biosphere

Reserve is intended to fulfil three complementary

functions: a conservation function — to preserve

genetic resources, species, ecosystems and

landscapes; a development function; and a

logistic support function — to support

demonstration projects, environmental education

and so on. Biosphere Reserves are not covered

by an international convention, but must meet a

set of criteria allowing them to fulfil properly their

three functions. A number of Biosphere Reserves

simultaneously encompass areas protected under

other systems (such as national parks or nature

reserves) and other internationally recognised

sites such as Ramsar wetland sites.

India launched its own Biosphere Reserve

Programme in 1979. Currently, 11 Biosphere

Reserves — Nilgiri, Nandadevi, Nokrek, Gulf of

Mannar, Manas , Great Nicobar, Sundarban, Dibru-

Saikhowa, Simlipal, Dehang Debang and

Pachmarhi operate in India. In addition, Namdapha,

Valley of Flowers, Thar Desert, Rann of Kutch,

Kanha and the North Islands of Andaman have

been identified by the Indian Man and Biosphere

(MAB) committee as potential sites for BRs.

The extent to which this network of BRs

has fulfilled its multiple objectives could be

evaluated using a carefully selected set of

indicators. We provide below a sample of such

indicators, using available information and

insights gained from personal fieldwork. These

are by no means exhaustive; rather they are

indicative of how a development intervention may

be evaluated in a responsive and adaptive way.

Extent of awareness about goals of the

programme at various hierarchical levels

Changes in levels of information about

programme objectives and implementation at

various hierarchical levels would be an important

indicator of management capacity and

effectiveness. In the context of Biosphere

Reserves, such an understanding is, to a great

extent, available at international and national

levels, but not at the State or local levels. Indeed,

the ability of local people to appreciate the

objectives of a development intervention is a

requisite for their involvement at various levels

of planning and implementation. Our experience

suggests that people living inside a Biosphere

Reserve area rarely have any idea of the objectives

of the programme, though people are more

conscious of other conservation categories, like

National Park, overlapping with a BR area.

Moreover, there is often a different kind of

understanding available at the local level that does

not get transferred upwards. For example, in the

Nandadevi BR, banning of seasonal grazing in

the alpine meadows, locally called bugyals , inside

the BR (core area) has affected traditional

livelihoods; at the same time, locals believe that

this has reduced the diversity of medicinal herbs,

replacing them with extensive growth of a few

species — a development not accounted for in

plans decided higher up (Hans Raj Negi, pers.

comm.) .

Incorporation of local management and cultural

practices

The degree to which a development

intervention incorporates existing management

and cultural practices could be an indicator of its

responsiveness. The costs imposed on specific

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THE INDIAN BIOSPHERE RESERVE PROGRAMME

communities as a result of a course of action taken

at a large physical (and hierarchical) distance are

typically undervalued. In this regard, the Indian

Biosphere Reserve Programme seems to have

performed poorly as there have been few

systematic studies in this area (an ongoing study-

on Bhotias, a nomadic tribe in Nandadevi BR

undertaken by the Wildlife Institute of India is a

notable exception). Consequently, development

plans have gone haywire; tribes like Shompens,

for example, have refused to accept a development

model based on improved access to amenities

unrelated to their cultural perceptions. Ecotourism

plans have similarly missed out the need to merge

local cultural practices (represented through

performing arts, for example) as a part of the

development strategy.

In many cases, locally evolved conservation

practices still continue in a limited way. The Bodo

tribals in the Manas BR area protect parts of the

forest as sacred groves — a practice which

harmonises well with the concepts of supply and

safety forests. The Bodos also maintain their

traditional varieties of crop species (whereas

Assamese, Bengali and Nepali settlers in the area

have taken to cultivating high yielding varieties),

while the Bodo practices still continue. Bhotias

of the Nandadevi BR area graze their livestock

and collect medicinal herbs from alpine meadows,

maintaining an optimum grazing level to ensure a

continued supply of herbs. The closure of the

core zone of Nandadevi BR to human activities

has deprived local communities of their traditional

health practices.

There is also significant local knowledge

available with fisherfoik all over India. The

fisherfolk on Moyar in the Nilgiri BR are able to

describe in great detail the time course of siltation

and shallowing of the river stream and the

consequent changes in fish fauna. However, very

little official documentation is available on the

aquatic ecosystems of most Indian BRs. Again,

herders in Nandadevi BR narrate the changes in

the alpine meadows following the grazing ban,

pointing to the disappearance of several medicinal

plant species. Such knowledge almost never plays

a meaningful role in management plans for Indian

BRs.

Enhancement of entitlements

Improved access to a basket of goods and

services consistent with livelihood needs

indicates the success of the development function

of Biosphere Reserves. However, this kind of

indicator is to be used with caution when the

alternative livelihood activity imposes non-

monetised costs on specific communities. Thus,

loss of family members due to dependence on a

hazardous activity needs to be suitably weighted

against additional income generation. In other

words, additional incomes, as Sen (2000) argues,

need to create an expansion of ‘human freedoms’

and elimination of ‘human unfreedoms’. The

‘unfreedoms’ are typically imposed by historical

disadvantages or ecological vulnerability. The

Indian Biosphere Reserves seem to have

performed well on the income generation criteria

alone but not so much when judged by the costs-

imposed criterion.

In income generation activities, the role

played by voluntary agencies deserves mention.

In the Sundarban BR (SBR) area, a number of

NGOs currently operate, most notably the

Ramkrishna Mission Lok Shiksha Parishad

(RMLSP) and the Tagore Society for Rural

Development (TSRD). Both these organisations

play a potent role in making people appreciate the

development objectives of Sundarban BR. TSRD,

for example, has set up handloom weaving and

honey processing units in the Sundarban BR area,

with a view to providing alternative employment

opportunities (alternative to uncontrolled

brackish water aquaculture and forest based

livelihoods). These units serve an important

development function, relieving pressures on

forest based occupations and reducing human

hazards due to human-wildlife tensions. Similarly,

RMLSP works in direct collaboration with the

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Sundarban BR directorate to involve local people

in the social forestry programme and other eco-

development activities. This is a step in favour of

merging the conservation and development

functions of BRs, though an overwhelming

majority of farmers are totally ignorant of such

efforts. This leads us to the issue of strengthening

extension functions of a BR programme; at the

programme level, inadequate attention given to

resource based extension activities reduces the

impact of the interventions being implemented

by the non-governmental sector.

While the Sundarban BR has successfully

created income opportunities through

agriculture, aquaculture and cottage industries,

it has failed to reduce man-tiger conflicts due to

continued dependence of the Tiger Reserve

fringe population on non timber forest produces

(NTFPs). In the Sundarbans, about a million

people living close to the Project Tiger area have

been heavily deprived of access to their traditional

resource bases inside the forest areas, due to

restrictions imposed by the project. This

suggests that income generation schemes have

not harmonised well with traditional liveli-

hoods or have been planned on an inadequate

scale.

Moreover, human development as

measured by access to a bundle of amenities like

healthcare, primary education and

telecommunication is not seen as an integral part

of the development component of the Biosphere

Reserve strategy, especially in remote areas of

Nokrek and Nandadevi BR.

People-nature conflicts

The degree to which conflicts between a

community and its resource catchment zone,

defined by livelihood imperatives, are reduced due

to a development intervention would be an

indicator of its sustainability and effectiveness.

On this count, Indian Biosphere Reserves have

clearly failed with such conflicts reported from six

of them.

Absence of employment options during

slack periods has, for example, led to frequent

people-tiger conflicts in the Sundarban BR.

Sundarbans, the only mangrove habitat with a

tiger population, has an interesting ecological

history. The British government had settled large

numbers of landless people from Midnapore

district and the neighbouring state of Orissa, in

the area, to maximise revenue earnings from the

land. When Project Tiger was launched in 1973

and access to the forest was curtailed, little

attention was paid to the biomass needs of the

local population and major livelihood activities

like NTFP collection were compromised. Prawn

fishing gradually became the major livelihood

activity but in the absence of genuine alternatives

in the slack season, people still entered the forest.

Man-tiger conflicts became frequent enough to

turn tigers into man-eaters. Man-crocodile

conflicts also increased due to over-dependence

of local people on fishing. The typical approach

to resolving these problems has been to try to

change the behaviour of tigers, so that they do

not attack humans. Efforts to check the aggression

of tigers have included schemes in which tigers

are habituated to drink fresh water (based on the

view that drinking of salt water is the cause of the

unusual aggression of Sundarban tigers). The root

solution to the problem lies in tailoring the

development functions of the BR programme to

create long-term livelihood options for the people

affected by these conflicts.

Manas BR also overlaps with a Tiger Reserve

area which does not have a buffer zone on its

southern periphery. Consequently, man-tiger

conflicts are very common, as human habitations

abut on restricted areas. Man-crocodile conflicts

pose a major problem in Great Nicobar BR as all

rivers in the region have significant crocodile

populations. Indeed, most Indian BRs have failed

to reduce such conflicts because of inadequate

integration of its development goals with the

conservation goals of the legally conserved areas

with which they overlap.

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Management Plans

Legal framework

The extent to which the legal framework in a

country harmonises with an intervention is an

indicator of its effectiveness. The Indian legal

system, for example, recognises National Parks

and Wildlife Sanctuaries, but not Biosphere

Reserves, as legal entities and thus imposes

purely conservation-oriented restrictions on

protected areas. Biosphere Reserves frequently

overlap with such protected areas and

consequently are governed by the relevant laws,

which may not be compatible with the BR

programme objectives. This leads, for example, to

total prohibition of economic activities like grazing

in Nandadevi BR, as mentioned earlier, depriving

a community of its livelihood and development

needs.

Simultaneously, zonation based on short-

term calculations has led to severe imbalances in

several areas. In the Manas BR, for example, paddy

fields and tea estates extend right up to the

southern boundary core zone of the designated

Tiger Reserve. Till about two decades ago, this

area was mostly covered by grasslands and

deciduous forests and was inhabited by a few

Bodo tribals. Today, refugees from neighbouring

Bangladesh have settled in these areas. An area

which could have been a natural buffer between

the densely populated Brahmaputra Valley and

the Tiger Reserve area is now deforested and

cultivated.

Inter-agency collaboration

The extent to which various government

departments and agencies coordinate among

themselves in the implementation of a programme

also indicates its effectiveness. The Indian

Biosphere Reserve programme seems to have

performed inadequately in this regard. Thus, there

is a lack of coordination between the Sundarban

Development Board (SDB) and the Forest

Department on the issue of mangrove plantation

in degraded forest patches around villages in the

Biosphere Reserve area. Similarly, in the Gulf of

Mannar BR, the key island of Krusadai in the BR

core zone is under the control of the Fisheries

Department, which is carrying a programme of

establishing aquaculture practices. Oyster

culturing is also carried out on a commercial scale

on the island. In the Manas Biosphere Reserve, a

seed farm is functioning in an area of 900 ha in

the core zone which used to be prime grassland

habitat for several endangered species.

However, there are also instances where

successful programmes conceived outside the BR

arrangement have been meshed into a BR area.

For example, the Joint Forest Management (JFM)

programme in West Bengal has been launched in

the Sundarban BR in collaboration with the

Sundarban Development Board and local NGOs.

Forest Protection Committees (FPCs) in the area

are now protecting mangrove patches in the

barren mudflats and barren intertidal spaces

between embankments.

Efficacy of development plans

The efficacy of development plans is an

important indicator of the BR programme. BRs,

by definition, need to fulfil integrated

conservation and development objectives.

Whether funds and equipment intended to fulfil

the development goals actually enhance

capabilities of target groups is the key question

here. In general, the funds made available by the

MoEF for the BR programme have rarely reached

target groups because of a general lack of

awareness about the objectives of funding both

among the implementors and the target groups.

In the Nandadevi BR, the Forest Department was

provided with vehicles to prevent illegal poaching,

but the vehicles could hardly reach the trouble

spots due to the mountainous terrain, and

poaching of musk deer and illegal export of musk

to neighbouring countries still continues from

Pithoragarh area. It is also reported that television

and VCR sets meant to screen documentary films

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THE INDIAN BIOSPHERE RESERVE PROGRAMME

for awareness creation in the area were actually

used by local politicians to show commercial films

to bag votes in the elections.

There are other examples of misplaced

priorities. Bhotias, the local shepherds in the

Nandadevi BR , were randomly provided with fuel

efficient chullahs which would consume less

wood and check deforestation. However, the local

people of the area found these of hardly any use;

since they needed big fires to tackle the severe

cold, which these chullahs could not provide. In

the Sundarban BR too, the experiment with fuel

efficient chullahs largely failed because the local

people were given practically no training on how

to use them and ultimately, the chullah chimneys

came to be used as farm implements.

Scientific research

Capacity building and demonstration

objectives of Biosphere Reserves may be

evaluated through the extent of ongoing scientific

research. The Indian Biosphere Reserves have

performed fairly well in this regard, when each

Reserve is taken separately. However, there has

been no attempt to create an integrated information

system for the entire network. Also, while major

biological or socio-economic studies are going

on in many of the BRs, multifaceted research

integrating ecological, social and management

imperatives seem to be lacking except in the Nilgiri

BR where the Kerala Forest Research Institute

(KFR1) and the Indian Institute of Science,

Bangalore have an active research presence.

Outer links — interaction with the larger

economy

BRs should aim at developing systems that

enhance positive interactions with the larger

economy. A significant area of such interaction is

the development of good information on

medicinal plant resources. However, Central

Institutes entrusted with this job (such as the

Central Institute of Medicinal and Aromatic Plants)

seem to have no interaction with BR authorities.

The Forest Departments too, normally have very

scanty information about these resources, and

the only source of information is possibly the

local collectors. With no mechanism to involve

these local people in managing these

bioresources, the present trends clearly support

the short-term profit making behaviour of

commercial interests. In Gulf of Mannar BR, the

fishing trawlers from outside countries exploit

more fishes in lesser time. Also several demand

driven activities like pearl fishing, and export of

white and black sea cucumbers as a food delicacy

to Southeast Asian countries continues. Similarly,

exploitation of seafood like crabs and lobsters

continues to supply the demand from countries

like Japan.

Negative environmental impacts of activities

dictated by the external economy need to be

tackled under the BR programme. In the Sundarban

BR, for instance, the local people collect prawn in

juvenile stages for aquaculture. In the process,

other small fish and crustaceans are incidentally

destroyed. Furthermore, ponds constructed to

store the prawn seeds breach the embankments

which are essential for keeping out saline water

from the cultivated land. No government agency

has so far paid attention to the issue, although a

few local NGOs have expressed concern. The

basis of BR management needs to be broadened

to address these issues.

A related issue is the extent to which BRs

create new conflicts or alleviate or accentuate

existing conflicts among resource users. In the

Manas BR, for instance, a state sponsored seed

farm exists in the core zone. This farm largely

employs urban people and caters to the demands

of urban centres in the Brahmaputra Valley. The

local Bodo tribals who live on the fringes are left

out of the process, resulting in insurgent

tendencies.

Similarly, in Nandadevi BR, excessive

tourist traffic around pilgrimage areas creates great

pressure on local resources like fuelwood and

generates large amounts of solid waste. However,

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a summary ban on trekking in the core area has

hampered the local economy. In this situation,

carefully planned ecotourism or ecotrekking could

be organised rather than going for a summary

ban on trekking, while allowing uncontrolled

pilgrim traffic.

Responsive Indicators

A robust programme level evaluation of the

Biosphere Reserve Strategy calls for more

information with regard to a much wider set of

scientific indicators. However, our snapshot view

suggests that the system of indicators should be

responsive enough to address specific concerns

of ecologically handicapped communities such

as those facing major conflicts with wildlife.

The following issues could dictate the

choice of indicator methodology:

a) Indicators need to be discriminatory at

stress and response level, alternatively in the

baseline and project scenarios. Ecosystems are

subject to varied amounts of stress based on a

range of social and ecological factors. For

instance, much of the Sundarban BR area was

historically subject to severe population

pressures due to settlements driven by a

revenue-maximising policy adopted by the British

government. An indicator of capabilities should

be sensitive (discriminatory) to stresses of this

kind. It should be able to assess whether the BR

strategy has been able to address this specific

local condition. As we have mentioned, judging

by this criterion, Sundarban BR has failed to

reduce man-wildlife conflicts in the area due to

lack of employment alternatives (in the slack

season) even though statistics of overall

employment creation for Sundarban BR give a

favourable picture.

b) Development interventions need to

improve access to an appropriate resource bundle

sustainably across communities. Thus, indicators

are to be community- (or user group-) specific

rather than aggregative when dealing with

resource access. Special attention should be paid

to ecologically or socially disadvantaged groups.

Local shepherds in the Nandadevi BR are a case

in point. These people traditionally depend on

their livestock for their livelihood. Collection of

medicinal herbs is a secondary activity. They live

in a relatively closed society with limited external

links. An intervention that aims at meeting a

broader conservation objective needs to carefully

address community needs such as alternative

livelihoods. Our indicator should necessarily

disaggregate these communities and their needs.

In this case, the intervention — banning of grazing

— failed not just in addressing the local livelihood

issue but also in taking into account local

ecological understanding. For instance, moderate

level of grazing maintains herb diversity and

contributes to local medicinal practices.

c) Participatory process documentation

built into a project implementation plan could

generate valuable information on local conflict

resolution, innovative cultural practices, etc.

Indicators could assess impact of interventions

on these issues. It is common for development

projects to be planned from above and evaluated

from above. Even indices for participation in the

project are calculated through limited appraisal

exercises planned at higher official levels. What

is needed is an in-built Monitoring and Evaluation

strategy where local people would be able to

continually evaluate the development

intervention through participatory processes.

Biodiversity Registers could be a possible tool

for this. Local people could be involved by teams

of local college/school teachers and students or

local NGOs in a participatory documentation

exercise regarding status of biodiversity,

management practices, development aspirations,

etc. This process could be repeated for a set of

indicators periodically to assess project

performance.

The major challenge becomes the use of

an adaptive evaluation methodology sensitive

to the proximate concerns of stakeholder groups.

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The typical logical framework adopted by the

World Bank and other donor agencies often

misses out an analysis of the appropriate

disaggregration level (See Table 1). This paper

suggests a modified frame incorporating this need,

which should lead to a more responsive

evaluation of funding efficacy.

Lessons Learnt

Biosphere Reserves are eminently suited

to fulfil our obligations under the Convention on

Biological Diversity which stresses on

conservation and sustainable use of biodiversity

and equitable sharing of the benefits flowing

therefrom. Biosphere Reserve, as a model for

conservation and development, is undoubtedly

a viable alternative to the protected area based

approach which stresses conservation at the

expense of livelihood options of people. However,

as our study shows, the BR model needs to

overcome several difficulties if it has to be

effectively applied. On the one hand, it needs to

be integrated with various regional planning

processes; on the other, the implementers need

to clearly understand its objectives and percolate

these down to target groups.

The emergent possibilities in the field of

biotechnology have made scientific research and

good information management all the more

important. As we mentioned earlier, while individual

BRs have taken up isolated research programmes,

there is no attempt to organise resource data at

higher levels. In Costa Rica, for example, the INBIO

institution is engaged in fulltime screening of

biodiversity of the country’s forests and Japan has

set up an institute for marine biodiversity

prospecting in Micronesia. India is rich in traditional

use-related knowledge of various medicinal plants

and similar activities ought to be organised as part

of the BR programme to take advantage of its

megadiversity status.

Promotion of the involvement of local

people in management of BRs ought to be

another thrust area. As we have mentioned, local

people often have a strong ecological perspective

on issues like grazing of livestock; this is not

accounted for in plans decided by higher officials.

Conflicts on resource use, as between Bodos and

urban people in Manas BR, need to be carefully

addressed in BR plans. For instance, the creation

of the Orinoco-Casiquiare BR in Venezuela,

covering 83,000 sq. km of rainforest area was a

response to concerns of Yanomani and Wekuana

tribals regarding development of the area by

outside interests. The presidential decree that

established this BR states measures to protect

the traditional livelihoods of the tribal

communities and acknowledges their land

ownership rights. The decree also prohibits any

colonisation of the area or any other interven-

tions that violate the rights of the communities

(http://nativenet.uthscsa.edU/archive/nl/9 lb/

0307.html).

Currently, 356 BRs are designated in 90

countries as part of the international network of

BRs. However, participation of a BR in the network

is voluntary. Only three out of India’s eleven BRs

are currently official members of the network. Such

networking would be crucial to fulfil the logistic

support role of BRs.

The Global Environment Facility with a

$2.75 billion replenishment in 1998 is currently

emerging as the major funding mechanism for

biodiversity. Funding from UNESCO’s regular

programme has progressively decreased over the

years, reaching a modest amount of $300,000 in

1995. The GEF, on the other hand, allocated $600

million to biodiversity during its first (post-pilot)

phase (1994-97). The GEF is progressively

stressing on funding integrated conservation-

development projects as opposed to those based

on protected areas. The present funding climate

thus favours a renewed interest in the BR model.

BR projects thus need to be streamlined to meet

the emerging funding criteria and the BR

philosophy needs to be embedded in country

priorities.

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG. -DEC. 2003

223

Table 1: Sample logframe for performance evaluation of Biosphere Reserves

THE INDIAN BIOSPHERE RESERVE PROGRAMME

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224

JOURNAL , BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG. -DEC. 2003

THE INDIAN BIOSPHERE RESERVE PROGRAMME

Acknowledgements Hans Raj Negi, and Silanjan Bhattacharyya. We

are also thankful to the Ministry of Environment

The paper draws from reports prepared by and Forests and WWF-India for funding

Shonil Bhagwat of RANWA, Pune, Niraj Joshi, support.

Bibliography

Anon. (1999): Guidelines for Monitoring and Evaluation

of Biodiversity Projects, World Bank.

Barbier, E. (1992): Valuing Environmental Functions.

London Environmental Economics Centre

Discussion paper.

Bhattacharyya, S. (unpubl.): A Report on the Human

Component of the Sundarban Biosphere

Reserve.

Dasgupta, P., A. Sen & S. Marglin (1972): Guidelines

for Project Evaluation, UNIDO.

Dixon, J. (1992): Analysis of economic-ecologic systems.

Oxford University Press, Oxford.

Gadgil, M. & RANWA (2000): Biosphere Reserves of

India. Pp. 680-690. In: Setting Biodiversity

Conservation Priorities for India. Vol II. World

Wide Fund for Nature - India,

Munasinghe, M. (1990): Managing water resources. World

Bank, UNO.

Negi, H.R. & M. Gadgil (unpubl.): Nandadevi Biosphere

Reserve: A status report.

Porter, G., R. Clemencon, W. Ofosu-Amaah & M. Philips

(undated): Study of GEF’s Overall Performance.

Global Environment Facility.

Ruitenbeek, H.J. (1992): Mangrove Management: An

Economic Analysis of Management of Options

with a Focus on Bintuni Bay, Irian Jaya. Report

for Environmental Management Development

in Indonesia Project, Halifax, Canada and Jakarta,

Indonesia.

Sen, A. (2000): Development as Freedom. Oxford

University Press, New Delhi.

UNESCO (1996): Biosphere Reserves: The Seville

Strategy and the Statutory Framework of the

World Network. UNESCO, Paris.

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG. -DEC. 2003

225

PERSPECTIVES ON THE USES OF BIOCIDES:

CONSERVATION STRATEGIES FOR THE NEXT CENTURY

Robert W. Risebrough1

Key words: India, biocides, DDT, wildlife

DDT has played a critically important role in anti-malarial programmes but future usefulness will

be limited by increasing resistance of the mosquito vectors to DDT and other insecticides. In areas

of past intensive use, DDT, in the form of its derivative DDE, has depressed the productivity of

sensitive species of birds, leading in some cases to their local extinction. In India, environmental

levels have appeared to remain below the threshold of chronic harm to wildlife, in part because of

lower levels of use and in part because of high rates of loss to the atmosphere. DDT use could

therefore continue in the anti-malarial programmes without environmental damage. The longer-term

threat to biodiversity and species abundance from the continuing use of biocides, including

insecticides, herbicides and rodenticides, in India, is expected to derive from harm to non-target

organisms in the form of poisoning and reductions of the biomass that constitutes the food source.

Although there have been tragedies such

as the loss of the cheetah (Acinonyx jubatus

venations ), and poaching is still a threat to the

remaining tiger ( Panthera tigris) populations,

India has maintained a higher density and

diversity of wildlife living in close proximity to

people than any other nation on the earth, a

consequence of the reverence for life inherent in

the majority cultures. In spite of increasing human

populations and their continuing encroachments

into parks and reserves, the biodiversity and

abundance of wildlife have remained relatively

high during the current period of rapid economic

growth. In this expanding economy, India has

become not only a major consumer of biocidal

chemicals in agriculture and in public health

programmes but also a major manufacturer and

exporter of these products. The Farm Chemicals

Handbook 2001 reports that India, with China, is

now the world’s largest and only source of DDT

and, with China, is now the world’s largest user of

DDT [Anonymous 200 1 ; DDT : 1,1,1 -trichloro-2,2-

bis (p-chlorophenyl) ethane, related compounds

and derivatives]. The American Bird Conservancy

has reported that India now accounts for about

43% of the global yearly use and a substantial

'The Bodega Bay Institute, 2711 Piedmont Ave.,

Berkeley CA 94705, USA.

Email: [email protected]

amount of the yearly production of 30,000 tonnes

of the insecticide Monocrotophos (also known

as Azodrin) (Anonymous 2002), one of the most

poisonous chemicals ever sold to the public

(Skripskv and Loosli 1994, Hudson et al. 1984);

its use has frequently caused massive mortalities

of birds (e.g. Mendelssohn and Paz 1 977, Goldstein

etal. 1999).

The decision by the Government of India

on March 5, 2002, to accede to the Stockholm

Convention on Persistent Organic Pollutants will

eliminate the use of DDT in agriculture and limit

its use in India to the control of malaria and other

vector-borne diseases. The Treaty will also end

the uses of 8 other chlorinated biocides which,

like DDT and its derivatives, move through

ecosystems, accumulate in food webs, and pose

various degrees of hazard to both humans and

wildlife. The era in which these chlorinated

hydrocarbon “pesticides” were to have eliminated

all vector-borne diseases and created a pest-free

agriculture has ended; the once-promising

technology had major inherent flaws. But the

production and use of large amounts of other

insecticides, herbicides and rodenticides, all

included here under the label of “biocides,”

continues, with the specific aims of reducing

populations of selected wildlife and plant species

that are considered to be injurious to humans.

226

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, I00(2&3), AUG. -DEC. 2003

PERSPECTIVES ON THE USES OF BIOCIDES

A Centenary offers an opportunity not only

to look back over the previous century but also to

anticipate the next century with the experiences of

the past as a guide. This essay will attempt to

achieve some perspective on the “pesticide” issue

with a focus on India, in part by addressing the

significance of the continuing use of DDT in the

malaria programmes. The word “pesticide”,

however, will be avoided; its use overlooks the

reality that not only “pests” are killed by

applications of these chemicals. Moreover,

“wildlife” is defined to include insects and other

invertebrates, so many of which have been the

subjects of manuscripts published in this Journal.

A Broader Perspective

The human species is hardly unique in

using chemicals to promote its survival and well-

being. Three of the most familiar plant species in

India, the tea ( Camellia sinensis) and coffee

(Coffea sp.) plants and the neem tree ( Azadirachta

indica ), derive their usefulness to man from a

complexity of chemicals that do not contribute to

the metabolic processes that keep these

organisms alive. Rather, they are the instruments

of a sophisticated form of chemical warfare in

which these and other plant species have engaged

throughout their evolutionary history to avoid

being eaten. Caffeine and a large number of other

chemicals have served this purpose well.

Strychnine, an alkaloid from the seeds of

Strychnos nux-vomica , and rotenone from the

roots of Lonchocarpus sp. and Derris sp. are

other “natural” chemicals that have become potent

biocides used by humans. An increasingly

important group of biocides, the pyrethroids, are

variations of the “natural” chemical pyrethrum

which is synthesised by Chrysanthemum

cinerariifolium and discourages its insect

predators. Such chemicals are therefore used by

both plants and humans for the same purpose.

In turn, biochemical processes have

appeared throughout the evolutionary history of

insects and other invertebrate plant consumers

that metabolised or otherwise inactivated the

chemical defences of the plants. The variety of

biocidal chemicals used by humans does not yet

match the arsenal used by other animals to eat or

to avoid being eaten. Within this broader

perspective, therefore, we might view the use of

biocidal chemicals by humans as a “natural” act,

but with critically important qualifications. The

use of “warfare” chemicals by other species is

highly selective and very rarely harms non-target

species. Selective pressures, against the needless

expenditure of energy, work to reduce the amounts

of these chemical warfare agents to no more

than what is necessary. Nature has therefore

provided guidelines that humans might profitably

follow.

Silent Spring

Not only did this book (Carson 1 962) launch

the environmental movement, its author has

become its Patron Saint. Is this acclaim justified?

The chemical industry vilified it for a number of

years but appears now to have accorded it a

measure of acceptance — it is hardly in the long-

term interest of the industry to kill organisms that

are beneficial or that maintain the biodiversity of

food webs. Nor is it in its longer-term interest to

produce products that threaten human health.

Moreover, production of biocides that are

substitutes for DDT and other chlorinated

hydrocarbons has been profitable. Many,

however, have criticised silent spring as

“unscientific” or as scientifically flawed (e.g. Ames

and Gold 1998). Advocates of DDT use continue

to assail it; Ross (2000) has described the

restrictions on DDT use as “the unfortunate

legacy of the self-styled environmentalists, whose

bible remains Carson’s silent spring.” If a

balanced and scientifically accurate historical

perspective on the merits of this book is to be

achieved, a task this essay can only begin, these

criticisms deserve to be fully and adequately

addressed.

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG. -DEC. 2003

227

PERSPECTIVES ON THE USES OF BIOCIDES

Rachel Carson was wrong in attributing the

mode of action of the biocides then in use to a

disruption of oxidative phosphorylation, the

universal process that creates usable energy

within the cell in the form of the ATP molecule.

Yet she had assembled a vast amount of

information and undertook what appears to have

been the best possible synthesis at that time, a

synthesis no one else was apparently even

attempting. Later research was to demonstrate a

broader and somewhat different diversity of

biocide effects. An imperfect and inaccurate

conclusion, which was nevertheless intellectually

honest, about the way in which these poisons

exert their effects, does not affect the validity of

her principal themes that biocides are poisonous

chemicals that were being applied in massive

amounts to the environment where they were

killing many non-target organisms, producing

unintended and unanticipated results such as

resistance, and posing potential dangers as yet

unsuspected.

She was also wrong on the cancer issue, in

assuming that the “natural cancer-causing

agents” are “few in number” and that most of the

modern cancers can be attributed to synthetic

chemicals produced by humans. Rather, only a

small percentage of the cancers not caused by

smoking can be attributed to synthetic chemicals

(Ames and Gold 1997, 1998). Apologists have

pointed out that completion of the book was a

race against her own impending death from breast

cancer. Here also, however, she was providing an

interpretation of the overall meaning of a

considerable body of information about chemicals

and cancer. The much larger amount of knowledge

about environmental cancers, that has

accumulated over the past 40 years, has not

supported her interpretation that was based on a

much smaller information base. Yet her attempts

to assess the significance and to predict longer-

term effects of the use and misuse of huge

quantities of poisonous chemicals can only be

considered as both intellectually honest and

socially responsible, values that later

environmentalists have not always maintained.

Interpretations inevitably change as more

information and more data become available.

Her explanation of the low productivity of

populations of bald eagles ( Haliaeetus

leucocephalus) that was observed in 1947 in

Florida (Broley 1958) and in a sanctuary

established for bald eagles on the Susquehanna

river [H.H. Beck, pers. comm, to R. Carson (Carson

1 962)] was confirmed only some years later: “There

is some occupancy of nests by adults, some

production of eggs, but few or no young birds. In

seeking an explanation, only one appears to fit all

the facts. That is that the reproductive capacity

of the birds has been so lowered by some

environmental agent that there are now no annual

additions of young to maintain the race.” This

explanation has since received full scientific

support. The “environmental agent” turned out

to be DDE, a derivative of p,p' -DDT, the active

biocidal component of technical DDT. Its principal

effect on sensitive bird species could never have

been predicted — alterations of the eggshell

structure such that the more contaminated eggs

would either break or fail to hatch. Like other

organochlorines, it moves readily throughout the

global ecosystem; its non-polar character results

in its accumulation in food webs, affecting those

species at the top of the food webs that are

sensitive to its effects. Carson’s explanation,

derived from the relatively small amount of

information then available, could not have been

more accurate — and scientifically sound.

Her documentation of the massive

mortalities of non-target species, the futility of

spreading large amounts of poisons over the

landscape, and the inevitable appearance of

resistance of the target species to biocides cannot

be challenged. Her position as Patron Saint of the

environmental movement is secure; imperfect

interpretations in the public interest, in response

to a genuine problem, have a value that is immune

to the attacks of either commercial interests or the

228

JOURNAL BOMBAY NATURAL HISTORY SOCIETY. I00(2&3), AUG. -DEC. 2003

PERSPECTIVES ON THE USES OF BIOCIDES

inflated egos of scientists defending an imperfect

technology that they, to their credit, have helped

to build.

How Bad is — or was — DDT ?

The deliberations leading up to the creation

of the Stockholm Convention prompted another

chapter in the continuing debates on the merits

of DDT (e.g. Roberts etal. 2000, Smith 2000), with

some environmental groups including World

Wildlife Fund (Anonymous 1998, 1999a) and the

Physicians for Social Responsibility (Anonymous

1999b) pushing for an ultimate global ban on the

use of this chemical, with organisations such as

the World Health Organization pointing out its

continuing value to public health programmes

(Kapp 2000), and with many individuals and

organisations proclaiming either that the use of

DDT has been one of the greatest achievements

of human civilization or that its hazards justified

an immediate global ban. At best, this essay will

be only a modest first attempt at a definitive

assessment of the pluses and minuses of a

chemical that will — whatever the final verdict —

have a prominent place in the history of

technology.

Certainly, there is no disagreement that its

use was spectacularly successful in the

eradication of malaria in many areas of the world.

By creating a temporary absence of mosquito

vectors while the disease took its course in

humans, the cycle of transmission was broken

and the Plasmodium species responsible for the

disease became locally extinct. But malaria has

returned with a vengeance to Mumbai (Kamat

2000) and the rest of India (Sharma 1 996). Would

the eradication campaign in India have been

successful, as it was in countries such as Taiwan

(Yip 2000), had not the extensive use of DDT in

agriculture hastened the inevitable appearance

of resistance? Would a more concerted effort of

spraying the walls of houses with DDT within a

relatively short time period have resulted in

success? These are questions for which no

adequate answers are available, although it is

likely that in both cases the answer would be

positive. In any case their relevance to current

policy is diminished; in India there is a high degree

of resistance to DDT, and to several other

insecticides, among the vectors of malaria (Sharma

1999); the current global emphasis in malaria

control programmes is on treatment rather than

the elimination of the vectors (Baird 2000).

Also, it cannot be disputed that DDT is still

being successfully used to control and to

eradicate malaria in some areas of the world

(Dapeng etal. 1996).

The first objections to DDT came after the

massive mortalities of birds, that followed its use,

mainly at higher doses and particularly in a

programme to “eradicate” the gypsy moth

Lymantria dispar on Long Island in New York

State, USA. The gypsy moth is an introduced

species that each year has defoliated oak ( Quercus

sp.) woodlands of the northeastern USA. A

lawsuit was brought in the 1950s against the US

Department of Agriculture by Robert Cushman

Murphy and Grace Murphy, to stop the

“eradication” programme on the grounds that

birds were worth more than the loss of foliage,

which usually did not kill the trees, and that the

programme was not working as the moths came

back year after year. Although lost on

technicalities, the lawsuit nevertheless exposed

the weaknesses in the “eradication” concept and

prompted the beginnings of change in public

perceptions of “pesticide” use. So entrenched

were these public perceptions that the Audubon

Society, the major US conservation organisation

at that time, did not support the lawsuit of the

Murphys, nor would it provide a pre-publication

endorsement of silent spring (R. Clement, pers.

comm.).

The next lawsuit was brought in 1966 by a

Long Island housewife, Carole Yannacone, who

objected to a fish kill in a local marsh. Ultimately

the court decided that it did not have jurisdiction,

JOURNAL , BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG. -DEC. 2003

229

PERSPECTIVES ON THE USES OF BIOCIDES

with no law upon which to base a decision, but the

public awareness generated by the case resulted in

the ending of DDT use in the local marshes.

In the autumn of 1968, American elm trees

Ulmus americana were to be sprayed with DDT

in the city of Milwaukee, Wisconsin, to combat

the Dutch elm disease, caused by the fungus

Ophiostoma ulmi which is carried from tree to

tree by either of two species of beetles, the

European elm bark beetle Scolytus multistriatus

or the native elm bark beetle Hylurgopinus rufipes.

The disease was then killing the elms over

northeastern North America, where it had long

been a cherished shade tree; in previous years

the programme had killed many hundreds,

probably thousands of birds. The Citizens Natural

Resources Association, Inc. of Wisconsin, joined

by the Izaak Walton League of America, requested

a hearing that would determine whether DDT

might be considered an environmental pollutant

under Wisconsin law, which defines

environmental pollution as “contaminating or

rendering unclean or impure the air, land or waters

of the state, or making the same injurious to public

health, harmful for commercial or recreational use,

or deleterious to fish, bird, animal or plant life.”

The hearing was undertaken in a courtroom

format, with vigorous and frequently hostile

questioning of witnesses. Critically important

financial support was provided by the National

Audubon Society, which was no longer neutral

on the DDT issue.

I was privileged to participate in this

hearing, initially to testify on the mobility of DDT,

and later on the effects of DDT on pelicans and

other fish-eating birds. Conflicts with industry

representatives were frequent, and bitter. The

species principally affected in Wisconsin were

the peregrine falcon Falco peregrinus that had

become locally extinct, the bald eagle that had

almost disappeared, the osprey Pandion

haliaetus that was producing thin-shelled

eggs, and the double-crested cormorant

Phalacrocorax auritus that was much reduced

in numbers. In the state of Wisconsin, the value

of these species and of the integrity of the

ecosystem was considered higher than the

economic benefits of a cheaper insecticide; the

Hearing Examiner, Maurice Van Susteren, ruled

that DDT met the criteria of an ‘environmental

pollutant’ as defined by the law and its use ended

in Wisconsin.

There were no public health issues to be

addressed in Wisconsin. Ideology did not intrude

on the relatively straightforward scientific

arguments that were relevant to application of

the law. On the industry side, however, ideological

considerations occasionally obscured the

defence of economic interests; the questioning

of “pesticide” use was considered unpatriotic,

seditious and perverse (e.g. McLean 1967).

Later, in 1970-72, the newly formed

Environmental Protection Agency conducted

hearings in Washington, D.C., to address the

question whether the use of DDT should be ended

on a national level. More than 100 witnesses

participated, representing almost all facets of DDT

use and documenting a wide range of

environmental effects. Here also, confrontations

between environmental scientists and the DDT

industry were frequent and bitter. The Hearing

Examiner, a retired lawyer with no scientific or

environmental credentials, ruled that DDT did not

constitute a danger to wildlife or people, but he

was overruled by the Administrator, who found

that continued DDT use posed an unacceptable

potential risk to human health (Ruckelshaus 1 972).

The wisdom of this decision continues to

be challenged (e.g. Lieberman and Kwon 2002).

The decision was supported largely by the

possibility that DDT or one of its derivatives might

be a human carcinogen, on the basis of rodent

studies undertaken by the International Agency

for Research on Cancer (IARC) using high

amounts of DDT (Anonymous 1991, and earlier

reports of the IARC). A vast amount of

subsequent research has failed to find a link

between DDT compounds and human cancers,

230

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003

PERSPECTIVES ON THE USES OF BIOCIDES

including breast cancer (e.g. Adami et al 1995,

Ahlborg et al 1 995, Wolff et al. 2000, Laden et al.

2001). Yet this decision by Ruckelshaus in 1972

not to permit a continuing build up of DDT

residues in the environment must be considered

the only responsible one that could be made at

that time in a country with no major problems of

vector-borne diseases. It was just too early to

make any definitive statement about the

carcinogenicity or other adverse effects of DDT

on humans; contamination of the global

environment could not be quickly reversed if such

an effect were discovered. The thinning by DDE

of eggshells of a number of bird species, and

depression of productivity in many of those

species, provided an example of the deleterious

effect of a synthetic chemical in the environment,

that most likely would never have been predicted

from the broadest range of toxicity tests in the

laboratory. Similarly, a longer-term subtle but

deleterious effect on humans might not be

detected until years later. The same argument was

used for ending the use of the polychlorinated

biphenyls (PCBs), which also were produced in

large quantities, and like DDT are mobile and

persistent, and are accumulated in food webs

(Risebrough and Brodine 1970). In the case of the

PCBs, the validity of the argument has been

abundantly vindicated; an entirely unexpected

effect of the PCBs is a depression of learning

capacity of human children exposed to PCBs in

the womb (e.g. Darvill et al. 2000). Any increase

above current levels of PCB contamination in the

environment would have disastrous effects on

human welfare; the release of large amounts of

persistent, mobile, toxic chemicals into the global

environment constitutes an experiment with

unpredictable results. A policy that would prevent

their accumulation is both sound and defensible.

Meanwhile, other persistent chemicals with

demonstrated biological activities and which

accumulate in food webs are now being detected

throughout the global environment; the

brominated diphenyl ethers, widely used as fire

retardants, are an example (Kuehl and Haebler

1 995, Noren and Meironyte 2000, She et al 2002).

Fluorinated surfactants, with unknown biological

properties, have recently been detected in marine

wildlife in concentrations that must be considered

“high” (Giesy and Kannan 2002).

The lessons from DDT and the PCBs about

the hazards of releasing large amounts of

persistent mobile chemicals into the global

environment have yet to be learned. Moreover, it

is still not possible to conclude that DDE has had

no harmful effects in humans. A report of an

association between DDE levels and human

premature births (Longnecker et al. 2001) might

be compared with an earlier study of premature

births of California sea lions Zalophus

calif ornianus. Concentrations of DDE in the

blubber of female sea lions giving birth

prematurely, in 1972 on the southern California

island of San Miguel, were eight times higher than

in females with full-term births. Yet, confounding

factors, such as the possible presence of a disease

factor, age, nutritional status, possible

contributions by other contaminants and the

absence of previous pregnancies, precluded any

definitive conclusion that DDE was a contributing

cause (Gilmartin et al. 1 976). The hypothesis that

DDE was at least in part responsible, however,

remains valid, as does the hypothesis of a

comparable DDE effect on humans. Although

confounding factors may eventually be shown to

be the responsible factors, an association between

DDE levels and the duration of lactation of nursing

human mothers (Gladen and Rogan 1 995) appears

to be another probable effect of DDE on humans.

Although solid evidence for an adverse

effect of DDT use on human health remains

elusive, the evidence for an adverse chronic

environmental effect in the form of a depression

in productivity of a number of bird species has

strong and convincing scientific support. The

Environmental Defense Fund, which undertook

the legal challenge to the use of DDT in both

Madison and Washington, other environmental

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003

231

PERSPECTIVES ON THE USES OF BIOCIDES

groups, and individual environmental scientists

had built their case against DDT on the basis of

the scientific evidence for adverse environmental

effects. DDE has been shown to produce eggshell

abnormalities in these species, which result in

either breakage of the eggs or their failure to hatch;

the biochemical mechanism, however, is yet to be

adequately described. Experimental studies have

produced the same effect with concentrations of

DDE that existed in the environment; other

environmental contaminants such as the PCBs

and methyl mercury produced no effects on the

eggshells (Risebrough 1986). Following the

national ban on DDT use, and the ending of the

discharge of waste DDT into the sea from a

manufacturing plant in Los Angeles, recovery of

populations of bald eagles in eastern Canada (Grier

1 982), ospreys of New England (Spitzer et al. 1 978),

brown pelicans Pelecanus occidentalis of

California (Anderson et al. 1975) and peregrine

falcons throughout North America (Cade et al.

1988) was rapid. Bald eagles are still unable to

reproduce in the contaminated marine

environment of southern California (Garcelon et

al. 1989, Jenkins et al. 1994), but elsewhere in

North America the environmental levels of DDE

are generally now below the threshold for harm.

By the late 1960s, both DDT and PCB

compounds were being detected in marine birds

at concentrations that were likely to be above or

near the thresholds of biological effects

(Risebrough 1969, 1 97 1 , Risebrough et al. 1968).

Recognition of the magnitude of this

contamination (e.g. Goldberg et al. 1971) was a

principal factor in changing perceptions about

the scale of human activities within a global

ecosystem of a finite size. DDT (or PCBs) used

anywhere in the world would ultimately move into

the global environment. However, in spite, of the

earlier interest in global marine pollution, no recent

data are apparently available that would indicate

the magnitude or direction of current trends.

In summary: DDT played a critically

important role in reducing the incidence of vector-

borne diseases. Locally it is still effective for this

purpose, but since resistance of the insect vectors

to DDT continues to increase, its longer-term

usefulness is low. Concerns about hazards to

human health cannot be dismissed, but these

hazards appear to have a lower risk of harm than a

vector-borne disease. Its derivative DDE has the

unique capacity to depress productivity of

sensitive bird species, but only above

environmental thresholds of harm. If low-volume

use of DDT does not increase environmental

levels of DDE to the threshold for the most

sensitive species, environmental damage would

not occur. Like other biocides, however, DDT may

kill many non-target organisms and disrupt local

food webs.

DDT Contamination

of the Indian Environment

Press and internet communications have

reported that the manufacturing capacity of

Hindustan Insecticides Limited, a government-

owned company that is the only producer of

technical DDT in India, is in the order of 9-10,000

tonnes/year, that domestic consumption has been

3.5-5 tonnes and that an equivalent amount has

been exported in recent years. It has not been

possible to verify these figures and they are

therefore considered to be estimates only. Data

from the US Tariff Commission indicates that the

production of DDT in the USA reached its peak

in the mid 1960s at about 73,000 tonnes/year. If

we assume that half of this was exported, then

excluding Alaska, domestic use at that time was

about 5 kg/sq. km/year. Indian domestic consump-

tion at a rate of 3,600 tonnes/year would therefore

be only 10% that of peak use in the USA, but

since the land area of India is 39% that of the

USA outside of Alaska, recent DDT use in India

has been about 1/4 of peak US use on an area

basis. Before considering other factors such as

differences in rates of evaporation and loss,

environmental effects in India would be expected

232

JOURNAL , BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003

PERSPECTIVES ON THE USES OF BIOCIDES

to be lower than in the USA by a factor of

about 4.

On the basis of information on population

declines of peregrine falcons from North America

and Europe (Hickey 1969) and available

information about the worldwide effects of DDE

on peregrine falcons (Peakall and Kiff 1979), it

can be expected that the black shaheen Falco

peregrinus peregrinator , the local peregrine race

in India, might be the Indian species that would

show the greatest depression of productivity and

perhaps also a decline in population numbers. As

a predator of other birds it occupies the top of the

food web. There are apparently no data on

productivity or on residue concentrations that

would confirm or negate this prediction. Yet

shaheens have continued to breed near Mumbai

and continue to be seen in the city during the

non-breeding season and elsewhere in India

(R. Naoroji and L. Pereira, pers. comm.). Although

inhabiting a country of relatively high DDT use,

this race of the peregrine appears to have escaped

the severe effects this species experienced

elsewhere in its global range.

The North American bald eagle, feeding

primarily on fish, occupies a somewhat lower

position in the food web than does the peregrine

falcon. Its productivity is, however, depressed at

concentrations of DDE 4-6 times lower than the

threshold that affects the productivity of

peregrine falcons (Nisbet and Risebroughl994,

Peakall and Kiff 1979). Although sensitivity to

DDE, measured as effects on productivity, varies

widely within the genus Falco (Fyfe et al. 1988),

the Pallas’s fish-eagle Haliaeetus leucoryphus ,

of the same genus as the bald eagle, is likely to

show depressed productivity in response to

accumulation of DDE. This species has not bred

in Keoladeo National Park in eastern Rajasthan

since the 1 980s (V. Prakash, pers. comm.), but there

are no data to suggest that DDE was a contributing

cause.

The relatively few studies of the

distribution of DDT compounds in selected bird

species (Muralidharan et al. 1992, Ramesh et al.

1 992, Tanabe et al. 1 998, Senthilkumar et al. 200 1 )

have shown that concentrations generally were

about an order of magnitude lower than those

associated with severe shell thinning and

reproductive failures of related species in North

America. Concentrations of DDT compounds in

three Ganges river dolphins Platanista gangetica

from the Ganges river at Patna (Kannan et al. 1 994)

were comparable to those in other species of

dolphins in other areas of the world, except for

the much higher levels that have been recorded

in dolphins from California coastal waters (Table 1 ).

Environmental levels of DDT compounds in India

appear to have remained below thresholds of

harm.

In part, this can be attributed to relatively

high mean temperatures. The rate of volatilisation

of DDT compounds into the atmosphere

increases sharply with increasing temperature

(Lloyd-Jones 1 97 1 , Risebrough 1990, Racke et al.

1997), enhancing their mobility through

ecosystems. DDT leaving India, however, enters

the oceans, thereby contributing to global

contamination of the oceans, and constitutes a

part of the DDT that has reached Antarctica.

Conservation Strategies and Policies

When I was on the witness stand at the

DDT hearings in Wisconsin in 1968, 1 was asked

if a small amount of DDT could not be used, in

Wisconsin, for any emergency purpose without

contributing to environmental harm. The answer

was “no.” Peregrine falcons had disappeared as a

breeding species, bald eagles had almost

disappeared, the eggs of an increasing number of

species were thin-shelled, and alarmingly high

concentrations were being measured in marine

wildlife. Environmental contamination levels were

above the threshold of harm; any additional DDT

would have increased the magnitude of damage.

Today, except for the marine environment of

southern California where bald eagles are still

JOURNAL , BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003

233

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PERSPECTIVES ON THE USES OF BIOCIDES

unable to reproduce, the response would have to

be a qualified “yes,” the qualification deriving

from an uncertainty about the longer-term trends

in the contamination of the global marine

environment. As long as environmental levels

remain below a threshold of harm to the most

sensitive species, any additional contribution of

DDT to the ecosystem could not be considered

as environmentally harmful. Particularly when in

India the use of DDT in agriculture is to end, under

the terms of the Stockholm treaty, opposition to

the continuing use of DDT for house spraying in

the anti-malarial programmes appears to derive

from ideological considerations rather than from

a pragmatic conservation policy.

The assumption that indoor use of DDT

does not result in environmental contamination

cannot, however, be supported. Studies in the

Brazilian Amazon have shown that local

contamination of soils and the food web of a

nearby river resulted from the use of DDT to spray

the interiors of houses in an anti-malarial

programme (Torres et al 2002). Volatilisation into

the atmosphere and hence into vegetation, soil,

water, etc., and back into the atmosphere,

processes that are highly dependent on

temperature (Risebrough 1 990), are the pathways

by which DDT compounds move from sites of

application or deposition. A dramatic example from

India is provided by the contamination of the soil

and of the human population in the vicinity of a

DDT manufacturing factory in Delhi (Saxena et

al. 1 987a, 1 987b). Yet, on the basis of the argument

presented above, DDT contamination resulting

from continuing use in anti-malaria programmes

in India will not be of environmental concern and

need not be a component of a conservation

strategy.

Among the other biocides to be banned by

the Stockholm treaty, aldrin and dieldrin have done

the greatest damage to wildlife (Nisbet 1988). The

poisoning of sarus cranes Grus antigone in the

vicinity of Keoladeo National Park (Muralidharan

1993) is an example from India that was

documented, among the probably countless

others that were not. These threats will diminish

as the uses of the chlorinated hydrocarbon

biocides end.

The major concerns in the past about the

use, misuse, and overuse of biocides have derived

from the effects these have had on the non-target

organisms that constitute much of the Indian

biodiversity, many of which have been described

and discussed in the pages of the Journal of the

Bombay Natural History Society over the past

one hundred years. It is likely that these will remain

the principal concerns throughout the foreseeable

future. Reductions of the magnitude of these

effects on wildlife and elimination of any adverse

effects on humans might therefore be considered

as the immediate goal; the longer-term goal must

be the maintenance of maximum biodiversity.

Whereas description and cataloguing of the

Indian biodiversity have been a principal activity

of the Journal over the past century, the

maintenance of the scientific basis for the

conservation and long-term survival of this

biodiversity will surely be a major focus of the

next century.

There are as yet very few hard data on the

impacts, on populations, of reductions in available

food supply following biocide applications.

Applications of insecticides generally do not kill

only those insects that are considered injurious

to humans, but other insects and invertebrates as

well, that are the food base for many birds and

other vertebrates. Applications of herbicides kill

plants that not only provide seed that supports

seed-eating birds but also are the food source of

many insects. Establishing monitoring

programmes that would detect changes over the

next century in the abundance of selected birds,

other vertebrates, invertebrates, and plants may

not initially appear to be an exciting scientific field

worthy of participation and support. Yet if the

problem is reframed to ask how changes in the

abundance of species can be detected and

measured, and how such changes can be

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG. -DEC. 2003

235

PERSPECTIVES ON THE USES OF BIOCIDES

adequately explained, it becomes a challenge to

the best of scientific minds and of all available

resources. It is such an approach that is needed,

to assess not only the impacts of future use of

biocides, but also of climatic change and all other

factors that might affect species abundance and

biodiversity. Moreover, without such

scientifically sound information, it would be

difficult to propose and to justify policies that

would protect the biodiversity.

The excessive influence of the “pesticide”

industry in California on the volume of chemicals

used and the ways in which they were applied

has been documented in a now-classic book by

van den Bosch (1978). In India, economic interests

of domestic manufacturers and the need for

foreign exchange will inevitably become factors

that influence biocide use patterns in both India

and the countries to which the biocides are

exported. An alert public that is well informed on

technical issues is critically important in

correcting balances towards the overall needs of

society. Available information provided at the

present time by the environmental groups,

however, is woefully lacking in accuracy. If this

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The research in India was supported by the

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PERCEPTIONS OF THE DEVELOPMENT OF WILDLIFE CONSERVATION

IN INDIA, HIGHLIGHTING THE PAST QUARTER CENTURY,

AND THE INPUT OF THE U.S. FISH AND WILDLIFE SERVICE

THROUGH THE BOMBAY NATURAL HISTORY SOCIETY

AND THE WILDLIFE INSTITUTE OF INDIA

David A. Ferguson1

Key words: Conservation, partnerships, cooperation, Service, assistance, research, wildlife

The Indian subcontinent has a rich natural heritage that has been largely maintained historically

because of the traditional respect for all life forms by the local peoples. Increasing pressures from

a rapidly growing human population have brought about deterioration of habitats and elimination of

species. These pressures and their consequences have gradually gained more attention over the

years, but in the last quarter century, a sea change in the actions and concerns of not only the

Government of India but also a number of organisations, have attracted widespread attention. The

Bombay Natural History Society (BNHS), a major non-governmental organisation, and the Wildlife

Institute of India (WII), a major Government of India institution, are making a difference in this

battle and have benefited from the input of international partners such as the U.S. Fish and Wildlife

Service (USFWS). Details of the two and a half decade relationship of the Service’s contribution to

India’s conservation movement through the BNHS and WII are provided, reflecting the benefits of

long-term commitment and working together for India’s wildlife heritage.

Introduction

What we know as the present-day Indian

subcontinent is a diverse landmass with a rich

natural fauna and flora where some of the world’s

earliest known human civilisations flourished over

5,000 years ago. There are signs that these early

civilisations not only recognised and used the

rich, natural biological diversity around them, but

developed views and policies governing use of

these resources mirrored in their sacred writings,

the Vedas and Upanishads. These teachings on

reality and morality linked to the five elements of

nature, “earth, water, fire, air and ether,” laid down

over three millennia ago, are recognisable today

and parallel modern-day thinking on ecology and

ecological relationships.

The legacy of recognition of, traditional

respect for, and protection of, all life forms by the

peoples of India, has resulted in the maintenance

'Division of International Conservation,

U.S. Fish and Wildlife Service, 4401 North Fairfax Drive,

Arlington, Virginia 22203, USA.

Email: [email protected]

of a rich natural heritage. This legacy was also

bolstered by the actions of royalty such as

Emperor Ashoka, who created reserves of forests

and issued edicts on the values of forests and

wildlife, describing acceptable behaviour, and

prohibited activities which could damage

renewable natural resources. While the forest

resources were often subsequently used by the

kings and their guests for hunting and were off

limits to others, their value as protected areas has

survived to the present, as many of the former

royal preserves now form the bulk of the national

parks and conservation areas network.

Since the last part of the 19th century and

early part of the 20th, many of the remaining

patches of forest and rangeland came under the

administrative hand of State Forest Departments

established under the British Raj for the primary

purpose of managing them for forest produce,

particularly timber. Despite the difficult transition

during the post-independence period and the

desperate need for resources from a rapidly

growing human population, the major wild land

resources represented in the forest preserves were

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able to survive because of the Forest Department

establishment.

Concurrent with the Government’s actions

to more formally protect and manage India’s

forests, the emergence of non-governmental

interests, in the late 1 800s, to focus attention on

other components of the biological realm was an

interesting phenomenon. Among the first of these

groups was the Bombay Natural History Society

(BNHS) which championed the study of natural

history in all its forms. Added to the more scientific

studies conducted by the BNHS, the writings of

colonials, wildlife observers, shikaris (hunters),

and foresters began to reveal the marvels of

India’s wildlife.

The achievements of the BNHS over its first

century have been documented elsewhere.

Likewise, the importance of foresters, the

evolution and management of wild lands, the

institutional description and linkage to wildlife

and wildlife management has been widely

chronicled. Despite these advances, gaps

between information on wildlife and their habitats

and management responses by the Forest

Department remained great.

In the late 1950s and early 1960s, India’s

wildlife scene started to pique the interest of the

international community. One of the earliest

ventures was what became a decades-long

relationship between the BNHS and the United

States’ Smithsonian Institution, focusing on the

study of birds. Dr. George Schaller of the New

York Zoological Society ushered in a new era with

the pioneering study of chital and tiger at Kanha

National Park, which employed some recently

evolving techniques, merging scientific method

with the behavioural study of animals and their

habitats.

The 1969 IUCN General Assembly, held in

New Delhi, could perhaps be viewed as a

watershed for the subsequent events that

triggered multiple actions affecting the wildlife

conservation movement in India — a chain of

events perhaps unparalleled anywhere else in the

world. This meeting brought together

international conservation leaders and

conservationists in India, including a small group

of influential Indian businessmen, academicians,

foresters and politicians (chief among whom was

the then Prime Minister, Indira Gandhi). Problems

were discussed, the plight of the tiger was

highlighted, and recommendations were made.

The key to the future development and

implementation of recommendations from the 1 969

meeting was the dedication and determination of

Mrs. Gandhi. In 1970, a 6 month duration course

in wildlife management was initiated in the Indian

Forest Research Institute (FRI) for in-service

Forest Officers, the first such course devoted

specifically towards wildlife. With the full support

of Mrs. Gandhi, the Wildlife (Protection) Act was

enacted in 1972. Under this landmark legislation,

wild animals and birds were given protection

through the creation and establishment of wildlife

advisory boards, hunting regulations, sanctuaries,

national parks and protected areas, wildlife officer

position, policies on trade and commerce in wild

animals and animal products, and a list of

protected species. In 1973, Project Tiger was

launched, and eight reserves were designated

specifically as tiger reserves.

At the request of the Government of India

(GOI), the Food and Agriculture Organization

(FAO) of the United Nations Development

Programme (UNDP) sent a consultant to India in

1974 to assess the crocodile situation and

recommend a programme for crocodile farming.

The relationship with FAO led to a number of

visits, creation of a captive breeding facility at

Hyderabad, a long-term restoration programme

for gharial and other crocodilian species, and a

major contribution to what later became the

Wildlife Institute of India (WII).

In 1976, India’s Constitution was amended

to allow the states to legislate forestry and wildlife

matters within their jurisdictions.

In the mid 1970s, the scene was set for the

entrance of another, perhaps unlikely, international

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player, the U.S. Fish and Wildlife Service (FWS).

Over the next quarter century up to the present,

the FWS has been engaged in a major wildlife

conservation programme with the GOI. This

programme, facilitated through local institutions

such as the BNHS and the WII, has influenced

and, hopefully, productively altered the

organisation and application of wildlife research

and management in India and the surrounding

region. It is recognised that major milestones were

occurring due to actions taken at the highest levels

of the GOI. A variety of other activities were also

going on, some as a result of the major events,

some as catalysts for these events, and some that

came about synergistically because of FWS input.

U.S. Fish and Wildlife Service and its Mission

As the principal U.S. Federal agency

responsible for conservation of wildlife, the FWS

has a mission to promote fish and wildlife

conservation and species enhancement, both

inside and outside U.S. boundaries. In 1973, a

special directive was given to the FWS with the

passing of the Endangered Species Act (Public

Law 93-205). The Act not only charges the FWS

to determine the status of threatened and

endangered species worldwide, but also allows

U.S. assistance to be offered to foreign nations

for the development and management of

programmes necessary or useful for threatened

or endangered species conservation. While the

drive behind FWS’s international programme is

conservation of endangered species and their

habitats, it was recognised that the best way to

proceed would be to engage and help existing

local institutions in other countries strengthen

their abilities and skills. An approach was

developed that sought out conservation-oriented

projects, focusing on research, management,

education and training, which could be addressed

by local institutions. By linking with various U.S.

organisations that could provide the technical

expertise not available with FWS and/or additional

financial or other administrative or technical

support, the FWS was able to marshal finances,

technical expertise, and logistical support that

have made permanent contributions to a multitude

of issues.

FWS AND THE BNHS

Fortuitously, early FWS engagement with

the GOI in the late 1970s was directed towards

BNHS, not only because of its pre-eminent

position and long history in facilitating bird

studies in the Subcontinent, but because it was

the only major organisation in India at the time,

with technical personnel experienced in natural

history research. Spirited by the leadership of

Dr. Salim Ali, the BNHS developed several multi-

year project proposals for Indo-US cooperation.

The first of these major projects to be endorsed

by the GOI was a 5-year effort starting in 1 979 to

monitor movements and habitats of birds. This

was quickly followed by two other 5 year projects

in 1980, the first an ecological study at what is

now Keoladeo National Park and the second on

the ecology of the Asian elephant and great Indian

bustard.

To date, the FWS has supported 16 major

multi-year conservation-oriented research projects

with the BNHS, covering 68 project years and

including studies on Bengal and lesser floricans,

birds of prey, wolves and blackbuck, hill stream

ecology, wetland ecology, grassland ecology and

many others. Each project is conservation-

oriented, implemented by the BNHS with local

personnel, has training as an important

component, and is done in conjunction with the

local state government (Forest Department). The

focus is to arrive at practical management options

in order to address resource issues and make some

permanent contribution to the knowledge base of

the country.

Besides the hundreds of peer-reviewed

scientific publications produced by these projects,

nearly 70 young and emerging Indian scientists

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have got their start or added to their experience

through participation in various FWS supported

projects. A total of 27 researchers of the BNHS

obtained their doctoral degrees, with many more

obtaining M.Sc. degrees, while working in FWS

funded projects. Several of these wildlife scientists

now play crucial roles in the wildlife research and

conservation scenario in India.

A number of projects, while fulfilling their

original objectives, often created new questions

to be asked, and were either extended or reformed

into new proposals. The original 5 year avifauna

project was extended to 7 years and spawned two

subsequent projects, one 5 year project on bird

migration and a 3 year bird banding project. The

original 5 year project on the hydrobiology of

Keoladeo National Park was extended to 10 years

and became one of the most comprehensive

ecological studies of an ecosystem in Asia. The

Birds of Prey project gave rise to 3 additional

studies, and so on. This is how science works.

Questions are posed as a hypothesis, then tested

to see if the hypothesis works. Along the way all

kinds of new questions and information arise.

The full impact of these projects with the

BNHS, as well as other projects supported by the

FWS with other organisations, is seen in the

ecological approach they brought to all studies

of species and their habitats. Species were no

longer viewed as inanimate objects to be studied

without context, but as parts of dynamic natural

systems that function or do not function in an

interdependent manner. If one part of the system

is affected, the effects are generally felt in the rest

of the system. Indian wildlife scientists were

becoming recognised internationally for their

ground-breaking work.

A more complete treatment of the BNHS/

FWS cooperative partnership in these long-term

projects can be found in Anon. (2000) and

Ferguson (2002), including lists of publications,

theses, project reports, presentations, and a list

of the biologists in their present positions who

worked in the BNHS/FWS projects.

Besides the FWS support to the many long-

term BNHS conservation research studies, the

partnership was further strengthened through a

variety of other activities which have fed back

into India’s conservation movement. FWS has

facilitated the participation of BNHS personnel in

international scientific meetings to present

findings from project related research. This not

only provides them with the opportunity to

disseminate information on India’s natural

systems to the international community, but also

provides them with experience in representing

India, builds self-confidence, and helps them make

contacts with potential cooperators and

collaborators.

The FWS has also provided opportunities

for BNHS scientists to visit American and other

foreign institutions and field sites for training,

interaction with scientific peers, making contacts

and confidence building. These opportunities

invariably serve to open minds and present

alternative approaches to problem solving and

ways to do things.

Top U.S. and European wildlife biologists,

managers and administrators have been sent to

India by FWS for interaction and information

exchanges with BNHS colleagues. Many of these

relationships have blossomed and endured for

lifetimes, creating new opportunities and

exchanges between personnel and institutions.

Believing in the need for a good library and

reference collection of scientific information, the

FWS has attempted to maintain a flow of scientific

literature to the BNHS’ main reference collection

in Mumbai, as well as to individual scientists at

their field sites upon request. A rough estimate of

the number of such books, publications and

environmental education materials would number

in the thousands.

Joint efforts with the BNHS on other

activities such as environmental education and

public awareness have also created multiple

effects. The 1983 publication of a pictorial guide

TO THE BIRDS OF THE INDIAN SUBCONTINENT arose OUt

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of the recognition that good illustrations of birds

were fundamental for identification and field

observation and no single-volume publication

existed that depicted all 1,200+ species of the

Indian subcontinent. Beginning with an effort in

1979, the FWS and BNHS persevered in this first

attempt to include 1,241 species in a one-volume

bird guide that could be taken to the field. John

Henry Dick, the well-known American bird painter,

provided the 106 illustrations free of charge, as

did the authors of the text, Drs. Salim Ali and

S. Dillon Ripley, as a tribute to the seminal work of

the BNHS in raising public awareness in natural

history and nature conservation.

With financial assistance from the FWS, this

publication was offered to the public on a non-

profit basis, in order to assure its widest possible

distribution and diffusion of knowledge

concerning India’s rich ornithological heritage

among low and middle income groups, among

government and private scientific, cultural and

educational institutions. The guide has become a

mainstay in the BNHS publication offerings, with

repeated reprinting to meet public demand, and

enabling well-informed citizenry to participate in

the appreciation and conservation of wildlife and

its habitats, and hence, in the preservation of a

healthy natural environment. Proceeds from sales

of the book also help support the production of

other BNHS publications aimed at creating

awareness of our natural world.

1983 was highlighted as the centennial year

of the BNHS. A year-long series of events was

scheduled to suitably mark the occasion of this

venerable institution’s 100th birthday,

culminating in an international seminar in Mumbai.

The FWS supported the participation of 12

representatives from U.S. scientific institutions

and universities, with prior experience in India, to

attend the seminar and present results of their

work, to add to the asset of growing data on the

country’s biodiversity. The FWS also invited

representatives from the BNHS on a U.S. tour to

promote the centennial celebration, disseminate

information on BNHS activities and objectives,

and attract a wider membership network. The

BNHS group was led by its President and

charismatic leader, 87 year old Dr. Salim Ali, and

included Mr. J.C. Daniel, Director and Curator, and

Mrs. Dilnavaz Variava, a member of the

BNHS Executive Committee. Making appearances

in Washington, DC, Chicago and Baraboo,

Wisconsin, the group made presentations, passed

out literature, signed up new members, were feted

at various functions, and received funding

donations.

One unusual, but significant action which

occurred during his time, may have extended

Dr. Salim Ali’s productive years. For many of his

later years, the Old Man (as he was affectionately

called) suffered from a hearing problem, which

became so severe that it had become embarrassing

for him to appear in public fora as he had difficulty

hearing questions, and carrying on meaningful

conversations. Though equipped with hearing

aids, they often seemed to be more bothersome

than helpful and BNHS requests for new batteries

became a frequent call to the FWS. Concerned

that this hearing impairment would detract from

the effectiveness of the U.S. visit, Mr. Daniel

confidentially asked the FWS to take the Old Man

to a hearing specialist soon after arrival. The Old

Man was tested and custom fitted with top-of-

the-line hearing aids despite his truculence and

unwillingness for this bother. Secretly paid for by

donations from a number of U.S. admirers, the

hearing aids — which the Old Man decided he

liked when they allowed him to hear the calls of

birds — remained a mystery to him as to how

they came about. The rest of the U.S. tour was a

success with the Old Man avidly interacting with

his host, returning to India and actively pursuing

his work for another four years until his death in

June 1987. Prior to his death, he was honoured

with an appointment to the Rajya Sabha

(Parliament) by the Prime Minister in 1985, and

used this platform to effectively raise the cause

of conservation to the highest levels of the

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USFWS AND WILDLIFE CONSERVATION IN INDIA

government. It is conceivable that his improved

hearing helped serve this mission more effectively.

Not to be overlooked, is the effect the

BNHS/FWS long-term conservation projects had,

on the growth and staff training of Indian wildlife

institutions, particularly the BNHS and its

offshoot, the Salim Ali Centre for Ornithology and

Natural History (SACON). Early on in the BNHS-

FWS relationship, concern was expressed by the

BNHS directorship that the projects were serving

as excellent training vehicles for young biologists,

but due to the “soft money” nature of their

support, the BNHS had little or no ability to retain

the best of these personnel once the projects came

to an end. Several proposals addressing this

problem were put forward to the GOI for

consideration. The main objective was to create a

mechanism within the BNHS structure to provide

positions and work for the qualified project

biologists. Unfortunately, the GOI never saw the

same view as that proposed by the BNHS. The

concept changed and energies were directed into

a somewhat different channel that eventually led

to the creation of SACON. Basically, the core staff

from the BNHS ecological study at Keoladeo

National Park moved to Coimbatore and formed

the new research centre at Anaikatti. SACON

continues to be led by a former BNHS researcher

from the Keoladeo ecology study and employs a

number of former BNHS staff and ‘graduates’

from the cooperative projects.

The FWS support for the BNHS continues

to the present and BNHS continues to hold its

pre-eminent position in studying, reporting on,

and generating conservation actions for India’s

natural heritage.

FWS AND THE WII

As noted earlier, it was not until 1970 that

formal instruction in wildlife management was

offered through the Forest Research Institute

(FR1) at Dehra Dun. Five years later, following

some initial visits by an FAO consultant, FWS

was requested by FAO to consider a request to

train several Indian wildlife personnel in the U.S.

Lack of funding prevented any actions at that

time and it was not until 1980 that the FWS had

much direct interaction with the programme that

would one day become the WII. Perhaps the event

that set the stage for future FWS involvement in

India, from a formal standpoint, was the adoption

of wildlife as a “High Priority” area of cooperation

at an India-U.S. Joint Sub-commission on Science

and Technology meeting in Washington, DC, in

June 1977. The sub-commissions were high-level

mechanisms to facilitate bilateral cooperation

between the two countries. Subsequent bilateral

fora continued to endorse this cooperation and

foster a programme involving a broad array of

interests. This mechanism has proved highly

valuable in providing a forum where FWS, as a

U.S. governmental agency, had access and a

working relationship with the appropriate Indian

governmental agency, in this case the Ministry of

Agriculture. Discussions and training for serving

wildlife conservation needs became a two-way

street.

In an attempt to encourage India to become

more networked into the global conservation

community, the FWS sent a CITES (Convention

on International Trade in Threatened and

Endangered Species of Fauna and Flora) team to

India in 1979 to discuss CITES implementation

requirements and to review the agenda of the

upcoming Second CITES Conference of Parties

(COP) in Costa Rica with GOI officials. The GO!

Joint Secretary, Ministry of Agriculture (Forests

and Wildlife) attended the Costa Rica COP. The

FWS provided support for the GOI Joint Secretary

to attend the CITES Technical Meeting in Bonn

in 1 980, whereupon India officially joined CITES.

In the same year, 1980, at the GOI request,

FWS sent two specialists in wildlife capture,

immobilisation and radio tracking techniques to

India to conduct training for Project Tiger

personnel, representatives from WWF-India, and

students from the wildlife classes at Dehra Dun.

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USFWS AND WILDLIFE CONSERVATION IN INDIA

Accompanying the U.S. technical team was a

cinematographer who recorded the capture and

immobilisation techniques on video. Copies of

the videos were then made available to a number

of Forest Department and Park offices as well as

at the FRI, Dehra Dun. They have served as

important training aids for many years. This

activity was followed by the visit of a team of two

FWS wildlife veterinarians to the FRI and the IVI

facility at Izatnagar to present classes on wildlife

diseases.

In 1981, India hosted the Third CITES COP

in New Delhi. The FWS provided financial support

for 15 participants from other countries and

assisted with other administrative costs. With this

meeting, India began several years of leadership

in Asia on CITES issues. Also in 1981, India

promulgated the Forestry Conservation Act which

required permission of the Central Government

before any forest land could be diverted to any

non-forest use. India adopted the World

Conservation Strategy, and joined a number of

other International Conventions calling for global

preservation of endangered species and their

habitats — Wetland (RAMSAR) and Whaling in

1981, Migratory Species (BONN) in 1 982, and the

Indo-USSR Treaty on Protection of Migratory

Birds in 1984.

During this period. Dr. Salim Ali was

pleading the case for more trained Indian

biologists in all aspects of wildlife biology and

management. This call translated into a month-

long workshop on wildlife management

techniques at Kanha National Park in January 1 982.

Sponsored by the FWS, the GOI, and the Madhya

Pradesh Forest Department, the goal was to

transfer wildlife ecology, management theory and

techniques to 60 forest officers and biologists

from Central and State Governments, universities

and non-governmental wildlife organisations in

India. Also present were representatives from

Bhutan, Nepal, Pakistan, Sikkim and Sri Lanka.

They gathered with 25 senior U.S. and FAO wildlife

biologists and the same number of Indian

foresters. The instructional material was converted

into book form, distributed widely to all

participants, and made available to a wider

audience at a nominal cost. Some of the present

WII curricular content resembles the syllabus of

the Kanha workshop and many of the participants

established important personal and professional

links with their colleagues in India and abroad.

The concept of establishing a training

school focusing on wildlife at the national level

had been building up for some time.

Recommendations and preliminary plans were

formulated by the UNDP/FAO specialists and a

draft plan was circulated at the Kanha Workshop.

The Kanha Workshop proved to be a major

catalytic action to the movement to create such

an institute and helped decide on its eventual

location.

Concurrent with the creation of the WII,

there was a push to separate the administration

of wildlife management from its existing position

under the Forestry Department in the Ministry of

Agriculture to a more independent position.

Advocates strongly felt the need for a separate

Ministry if wildlife was ever to get the attention it

deserved. A high-level committee was

commissioned in 1980 to study the matter and

make recommendations to the Prime Minister. This

culminated in a complete re-organisation of the

Central Government’s administration of natural

resources, resulting in a new Department of the

Environment which formally became the Ministry

of Environment and Forests in 1985. In the

process, wildlife was also separated from forestry,

with the Forest Wing being headed by the

Inspector General of Forests, and the Wildlife

Wing headed by the Additional Inspector General

of Forests (Wildlife). The newly created WII

answers to the Additional Inspector General of

Forests (Wildlife).

The charge of the new WII was to operate

training courses for foresters in wildlife

management and conduct field research to help

formulate priorities and guidelines for wildlife

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USFWS AND WILDLIFE CONSERVATION IN INDIA

conservation. Despite challenges such as the lack

of trained personnel for faculty positions, no

organised curricula for its training programmes,

no structural facilities for housing, and no

precedent for breaking out of the insularity of the

traditional education system, the organisers and

planners moved steadily forward. A portion of

the FRI at Dehra Dun was set aside for the WII

programme. On May 22, 1982, the WII formally

came into being as an autonomous body situated

at Dehra Dun. With the UNDP and the GOI sharing

the costs of the project, Mr. V.B. Saharia (a Forest

Service officer in charge of the Wildlife Diploma

Course at the FRI) was designated Acting Director

for the new Institute with Dr. John Sale, formerly

assigned to the UNDP/FAO crocodile project at

Hyderabad, as the FAO Chief Technical Advisor.

In the next five years, a permanent Director,

Mr. Hemendra Panwar, was selected. The

positions and criteria for faculty were developed

and staff selected, the Government of Uttar

Pradesh donated land for a new campus, and plans

for a new campus were designed and construction

initiated. State foresters began receiving more

specialised training in wildlife subjects.

In October 1988, the FWS and WII entered

into a 5 year cooperative arrangement to train WII

faculty in modem research techniques and provide

equipment for use in wildlife training and research,

including advanced computer hardware and

software. Endorsed by the Governments of India

and the U.S. under the Indo-U.S. Sub-Commission

on Science and Technology, this innovative

project has served as an excellent example of what

a small bilateral technical development programme

can achieve.

There were 12 subject areas originally

decided upon, in wildlife ecology and

management, but only 1 0 were dealt with in detail.

The remaining two were not developed as full-

fledged independent programmes. The

programmes initiated included:

Wildlife in Managed Forests

Field Research Methods

Interpretation and Conservation Education

Geographic Information Systems

Systems Analysis

High Altitude Ecology

Wetland Ecology and Management

Wildlife Health

Animal Damage Control

Library and Documentation

Represented in the group of U.S. scientists,

eventually totalling 33, who participated in this

programme to transfer technology, was expertise

from the FWS, U.S. Forest Service, National Park

Service, a university and the private sector. Each

expert stayed in India from 1 to 3 months and

then hosted their counterpart for varying lengths

of time during.a study tour in the U.S. Under the

programme, eighteen WII faculty members,

including the Director and project nodal officers,

visited scientific institutions, protected areas

and other field locations in the U.S. to get

acquainted with U.S. training programmes and

management techniques. A final component of

the exchange was a ‘show and tell’ workshop in

India by the Indian and U.S. counterparts to

demonstrate to WII faculty and other organisa-

tions and universities what was learned in the

transfer.

Besides the wildlife courses for foresters

developed at WII which included a 3 month

Certificate, 10 month Diploma, and a number of

1 week specialised Capsule Courses, its premier

programme is a 2 year M.Sc. course to foster

original wildlife research and enquiry, often

independent of the needs of the government. As

a further reflection of the intellectual expansion,

WII hosts a number of post-graduate Research

Scholars from cooperating universities to carry

out original field studies on subjects of mutual

interest. The consequences of this movement

away from the didactic, resource exploitative,

hierarchical approach to a questioning,

academically rigorous, independent and dynamic

approach to wildlife research, cannot be

overstated.

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247

USFWS AND WILDLIFE CONSERVATION IN INDIA

With the output of about 20 Diploma, 14

Certificate, and 75 Capsule Course participants

each year, besides the 7 M.Sc. and 25 post-graduate

Research Scholars at various stages of their

studies, the WII had a tremendous impact in

providing a direction and mass to the wildlife

movement in India in the early 1990s. The WII/

FWS project has been immensely successful in

achieving its objective of facilitating the

development of WII and its faculty in its formative

stages. An important by-product has been the

establishment of formal as well as informal

scientific information exchange relationships

between the WII faculty and a large number of

research, training and management institutions

and agencies in the U.S.

When the Indo-U.S. project between the

WII and FWS was completed in June 1994, a

second phase was approved, in 1995, to run for

another 5 years. The second phase was envisaged

to consolidate the gains obtained thus far and

expand the application of modem wildlife research

and management techniques. Phase II was made

up of 7 specific projects on management oriented

biodiversity research and on developing

laboratory and field technology and curriculum,

conducted by Indian scientists with the support

of U.S. scientists.

The 7 projects under Phase II were as

follows:

1) Identification of potential areas for

conserving biodiversity in the Indian

Himalaya.

2) Evaluation of Panna National Park in

Madhya Pradesh with special reference to

the ecology of sloth bear.

3) The relationships among large herbivores,

habitat and humans in Rajaji-Corbett

National Parks.

4) Impact of fragmentation on the biological

diversity of rainforest small mammals and

herpetofauna of the Western Ghat

mountains, south India.

5) Establishment of an Indian Cooperative

Wildlife Health Programme.

6) Establishment of a wildlife forensic capacity

at the WII.

7) Planning and development of interpretive

facilities in selected protected areas in India.

The WII and the FWS remained the main

collaborating agencies under this programme.

Several of the projects were extended for an

additional year. All are essentially completed as

of date, although the overall programme is not

yet formally completed. Several U.S. and Indian

organisations were also involved in the

completion of the programme, including the Indira

Gandhi National Forest Academy (IGNFA;

formerly FRI), Dehra Dun, Salim Ali Centre for

Ornithology and Natural History (SACON),

Coimbatore, and 4 veterinary colleges.

The results from this collaborative phase

are not yet all in. But it is clear that a long-term

professional relationship has been established

between WII and FWS. The programme has

fostered similar relationships with other major

conservation agencies such as the USDA Forest

Service, the U.S. National Park Service, the U.S.

Geological Survey’s Biological Resources Division

as well as several universities. The programme

has enabled WII faculty to develop professional

links with a host of scientists and institutions

around the world.

Although the collaboration with FWS was

designed to provide the most useful modern

technologies relevant to Indian wildlife work, the

more lasting legacy may be the inculcation of

problem solving capabilities employing a scientific

method, and the opening up of the WII to take up

an important role in the global conservation scene.

WII is now recognised not only in India, but also

internationally, as a major centre for training and

research in wildlife conservation in Asia.

1 would like to conclude by saying that the

FWS is proud of its role in helping further the

cause of wildlife conservation in India.

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USFWS AND WILDLIFE CONSERVATION IN INDIA

References

Anon. (2000): The U.S. Fish and Wildlife Service - BNHS

Partnership for Wildlife Research and Conservation

in India. Buceros 5(3): i-vi, 1-69.

Ferguson, D.A. (2002): U.S, Fish and Wildlife Service

and Bird Related Projects in India. Pp. 17-23. In:

Birds of Wetlands and Grasslands: Proceedings of

the Salim Ali Centenary Seminar on Conservation

of Avifauna of Wetlands and Grasslands, Feb. 12-

15, 1996 (Eds.: Rahmani, A.R. & G. Ugra).

Bombay Natural History Society, Mumbai.

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003

249

CHRONOBIOLOGY, ECOLOGY AND BEHAVIOUR

OF SOME INSECTIVOROUS BATS OF SOUTHERN INDIA

( With twenty-three text-figures and one plate)

M.K. Chandrashekaran1

Key words: Insectivorous bats, chronobiology, ecology, ethology, echolocation, breeding cycles

This review describes the experiments and results of our work with six species of insectivorous

bats, inside natural caves, in open spaces and in the laboratory, performed by my students and me,

in the Department of Animal Behaviour and Physiology, School of Biological Sciences, Madurai

Kamaraj University in the two decades from 1 975-1995. We had worked out in detail the behavioural

expressions of biological clocks in four species of locally occurring insectivorous bats: Taphozous

melanopogon, T. kacchensis, Hipposideros speoris, and Rhinopoma hardwickei and the activity/

roosting patterns of colonies of these bats in their habitat. Working at a depth of 40 m in a natural

cave, we discovered that there is mutual social synchronisation of the circadian rhythm underlying

the exodus flight of a colony of c. 600 Hipposideros speoris bats around the local sunset time. The

circadian rhythm in the flight/rest activity of a solitary bat in a solitary cave without other

conspecifics (‘social informers’) free-ran. We also report here results of experiments with H. speoris

showing that daylight dimmer than starlight (0.0001 to 0.0006 lux) streaming into a cave for c. 90

mins could entrain their circadian rhythm. H. speoris is very sensitive to light, and light flashes

lasting only 0.0625 msecs could shift the phases of the circadian rhythm. The spectral sensitivity

of the photoreceptors in H. speoris indicates that they have colour vision. The Indian false vampire

Megaderma lyra echolocates prey (frogs) by ‘listening’ to prey-generated noise on land (passive

mode) and by active echolocation of the frogs in water (active mode). The eyes of most insectivorous

bats are very small and are unlikely to participate in vision in the darkness of the night. It is

suggested that eyes may participate in detecting dawn, sunrise, dusk and sunset, thus measuring

daylength over the seasons of the year. There is annual breeding periodicity in Hipposideros

speoris. There are 4 distinct breeding cycles in Pipistrellus mimus , the female giving birth to twins

each time, making it the most prolific breeder among Chiroptera. Many of the observations reported

here were first reports of their kind when they were made and published. The ecology of the day

roosting sites, place fidelity and social interactions of insectivorous bats are fascinating facets that

deserve to be investigated further.

Introduction

Bats, like rodents, are the largest group of

mammals in the world with 1,001 species

(Mickleburgh et al. 2002) and wide

zoogeographical distribution. They can fly

thousands of kilometres to inhabit remote islands

in the Pacific and the Indian Oceans, live and

flourish in all habitable regions of the earth. The

Latin name for bats is Chiroptera which means

hand-winged, as the forelimbs are modified into

'Evolutionary and Organismal Biology Unit,

Jawaharlal Nehru Centre for Advanced Scientific

Research, RO. Box No. 6436, Jakkur RO.,

Bangalore 560 064, Karnataka, India.

Email: [email protected]

simple wings. There are two sub-orders:

Microchiroptera (insectivorous bats) and

Megachiroptera (fruit-eating bats).

Microchiroptera are found in every continent

except the Arctic and the Antarctic, whereas

Megachiroptera (about 175 species, all belonging

to one family Pteropodidae) are confined to Africa,

Asia and Australasia. The fruit-eating bats have

very large eyes and excellent eyesight, which helps

them to find their way and their food in the dark;

in contrast the insectivorous bats which rely

almost exclusively on echolocation for navigation

and foraging, usually have small eyes and very

poor eyesight.

The world around us in the tropics is a

fascinating place after sunset. There is swarming

250

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CHRONO BIOLOG Y, ECOLOGY AND BEHAVIOUR OF INSECTIVOROUS BATS

of myriad insects (mostly Lepidoptera, Diptera,

Coleoptera and Orthoptera), the air has the

fragrance of the flowers which open only at night

to their pollinating agents, there is the alluring

smell of ripening fruit too. This nightly world is

the arena of bats, metaphorically called ‘the birds

of the night’. The insectivorous bats, without

competition from birds, all of which (with a very

few exceptions like nightjars and owls) are day

active, crop the rich fare from this sea of insects

on the wing. The power of flight and ability to

echolocate prey are contributory factors for the

evolutionary success of insectivorous bats.

G. Neuweiler (Zoologisches Institut der

LMU, University of Munich) and his students

and the dedicated group of bat researchers at the

Department of Animal Behaviour & Physiology,

School of Biological Sciences, Madurai Kamaraj

University (MKU), Tamil Nadu, undertook a series

of experiments related to field ethology,

neurophysiology and chronobiology of nine

species of echolocating bats of the Madurai

region (9°58'N, 78° 10' E). Neuweiler (1984) has

summarised the findings of his team and reported

that the insectivorous bats of Madurai foraged in

three modes: 1) Surface gleaning, 2) Foraging

within foliage, and 3) Open air foraging. Neuweiler

elegantly demonstrated that the constant

frequency/ frequency modulation (CF/FM)

features and the bursts or train of 150 to 250

ultrasonic pulses per second emitted by the bat

species while hunting, adaptively varied

according to the mode of foraging and

topographic features of the feeding habitat

(Neuweiler 1984, 1990, Linked/. 1986, Marimuthu

and Neuweiler 1987).

One of the most spectacular sights

coinciding with the sunset in many parts of the

world is the exodus flight of huge colonies of

bats from their caves. In Bracken Caves in Central

Texas, people assemble at sunset to see millions

of free-tailed bats Tadarida brasiliensis mexicana

stream upwards out of the cave for minutes on

end. I noticed the same phenomenon when I

accompanied my students to cave sites in Madurai

when colonies of Taphozous melanopogon,

T. kachhensis, Hipposideros speoris, Tadarida

aegyptiaca, Megaderma lyra and Rhinopoma

hardwickei flew out of their caves, evening after

evening, within minutes of sunset, like

cloudbursts. This ‘emergence by coup’ obviously

is a collective behavioural expression of the perfect

synchrony of the biological clocks of members of

the bat colonies. This was the genesis of my

interest in investigations on physiology of timing

in various activities, chronobiology, and ecology

of the insectivorous bats of Madurai. A study of

the literature on the subject revealed that a few

people had indeed reported a daily periodicity or

circadian rhythm in insectivorous bats of Europe

(DeCoursey and DeCoursey 1964, Erkert 1970,

1976, 1978, Laufens 1969, 1973, Vout eetal. 1974)

and the U S A (Menaker 1961, Rawson 1 960), the

earliest of them being Griffin and Welsh (1937).

Griffin had discovered the phenomenon of

acoustic tracking of prey by insectivorous bats,

and famously coined the term ‘echolocation’

(Griffin 1958). Most research effort in India on

bats has been on taxonomy and embryology,

contributions to the latter made mostly by

M.A. Moghe, A. Gopalakrishna and colleagues

(Gopalakrishna 1949, 1950, 1955; Moghe 1952,

1958). Therefore my students and I decided to

concentrate on experimental studies of circadian

rhythms, and field ethology observations on

breeding cycles, ecology and the foraging

strategies that these bats employ. There was no

scientific information at all on biological rhythms

of tropical bats in literature, except the

observations of G. Neuweiler on the onset of

activity in a colony of flying foxes in Madras

(=Chennai) and dates of parturition (Neuweiler

1969). Brosset’s papers (Brosset 1962a, b, c, 1963),

first brought to my attention by Mr. J.C. Daniel,

served as an excellent introduction to the bats of

India.

The present review describes the

experiments and results of our work, in the

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CHRONOBIOLOG Y, ECOLOGY AND BEHAVIOUR OF INSECTIVOROUS BATS

laboratory, inside caves and in the open, carried

out by my students and me from 1975 to 1995. We

worked on the behavioural expressions of

biological clocks in four species of locally

occurring insectivorous bats: Taphozous

melanopogon , T. kachhensis , Hipposideros

speoris and Rhinopoma hardwickei , and have

accumulated, as a result, the largest database on

biological clocks of bats anywhere. This report

includes data obtained in the laboratory on free-

running flight activity rhythms in these bats and

their responses to light pulses and monochromatic

light pulses.

Terminology and Abbreviations

In the interests of readability, much of the

jargon of ‘chronobiology’ has been left out of

this essay. Some essential abbreviations and

symbols, standardised by Aschoff et al. (1965),

and since then much in use are given below.

LD: light/dark cycle

LL: continuous light

DD: continuous darkness

Circadian rhythm: daily rhythm with period

close to 24 hours (Latin circa - about; dies =

day).

Period: t (tau): natural period of the

circadian rhythm. In practice, time of onset of

activity in LL or DD averaged over several days.

Entrainment: When environmental factors

modify a circadian rhythm such that it has an exact

period of 24 hours as it happens in nature.

Zeitgebers: Environmental cues (LD cycles

for example) which entrain biological rhythms.

Free-run: State of the rhythm in LL or DD

and constant temperature.

Phase: Any point along the circadian

rhythm. Often expressed as CT (circadian time)

denoting that it is not local time. (e.g. Sunrise =

CT 0 hrs; Sunset = CT 12 hrs).

PRC: Phase response curve. Plot of the

responses of a circadian rhythm, in terms of phase

shifts, to perturbations (discrete displacements

of the rhythm on time of day — X-axis) as a

function of phase. (0° phase = CT 0 hrs; 180°

phase = CT 12 hrs). Perturbations can be light,

temperature, chemicals, social stimuli, etc. In this

review, only light pulse PRCs are described.

Housing of Bats and Experimental Methods

Freshly captured bats that had been kept

in the laboratory for close to a week, in a normally

lit room, were used in the experiments. Our first

problem was the capturing of the bats, but we

soon became experts at catching them in fine mesh

nylon mist nets. Taphozous spp. were the most

difficult to capture.

Taphozous melanopogon : Body weight

20-24 g, best hearing frequency 26-28 kHz.

Field observations: This is a very common

species and hunts high above the canopy in the

open air. The field observations were made in the

rock complex of the Jain Hills (Samanar Malai) in

the vicinity of the Keela Kuyil Kudi village some

8 km southeast of the Madurai Kamaraj University

campus. A colony of 150-180 animals of both

sexes inhabited the vertical cracks and deep

crevices of the rocks facing north. The bats

crouched on all fours in clusters of 6-8 in the inner

recesses of the crevices, some of them leading to

dark, dank caves which progressively became

narrow and inaccessible to observers. Older males

and females often roosted in regions closer to the

entrance, and were visible for observations during

the daytime, often bathed in sunlight. Early

observations lacked details about cluster size,

sex composition, age distribution, hierarchical

order, and exact numbers, for we had not yet

acquired infrared viewers. Later observations by

R. Subbaraj (unpubl.) confirmed the observations

of Brosset (1 962a) that the species was extremely

polymorphic, variable in size and colour. Males

were bearded. Later observations were made from

comfortable perches, on a colony that lived in the

outer yard of a temple 6 km west of the MKU

campus on the banks of a bend in the Vaigai river.

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CHRONOBIOLOG Y, ECOLOGY AND BEHAVIOUR OF INSECTIVOROUS BATS

Information on the various species of

insectivorous bats of the Madurai region, location

of cave sites, observing techniques of evening

exodus flight of bat colonies, foraging areas and

the bat lore could be obtained orally from the

villagers living around the MKU campus. Bat

counts were made during the onset of colony

activity following sunset, the observer lying on a

rock surface at the cave mouth and looking upward

against the dimly lit dusk sky and counting the

swift outward flight of the bats. In later studies,

we used an infrared sniper-scope (FJW Industries,

USA). Light intensity was measured during

evening out-flight at the cave mouth using an

AEG luxmeter and an optometer (United Detector

Technology, USA). The lowest level of light

intensity that could be directly and reliably

measured with the luxmeter was 0.25 lux, but using

the optometer on the energy scala scale we have

measured 5% of starlight intensities (c. 0.000 1 lux).

Counting the bats when they returned to the cave

was a more weary exercise, for each bat returned

at its own time.

Captive bats were kept in the laboratory in

lmxlmxlm wooden frame cages, wrapped

around with nylon mosquito net, with a sleeve to

put a hand in to feed the animals, change the

drinking water, etc. Our German colleagues

working in their laboratories, procured meal worm,

commercially available in Germany, in abundance

to feed their bats. Our meal worm was of very

small size, so we resorted to feeding our bats on

de-gutted cockroaches with elytra, wings, outer

cuticle and legs removed. First the bats refused

this unnatural fare but learnt to eat it after a

sufficient build-up of hunger. They were always

fed at night, and in later experiments during day

into activity time, when we had offered the bats

inverted light/dark (LD) cycles of 12: 12 hours, with

dark prevailing from 0800 hrs to 2000 hrs. Water

with a few drops of Vitamin B- 12 was available ad

libitum.

Taphozous kachhensis : Body weight 48-

54 g, best hearing frequency 24-26 kHz.

This is the largest and most sturdy species

among the commonly occurring echolocating bat

species in Madurai. It is as fast a flier as Tadarida ,

foraging at high speed (10-15 m/s) at heights of

17-30 m above the ground in unobstructed areas.

A colony of these bats lived in the deep cracks

and crevices of a rock complex called Pannian

Malai, 5 km west of the MKU campus, not far

from the road leading to Theni and Thekkady.

The eyes of the bats were seen glistening during

the day, when one peered into the cracks and

crevices. The temperature in the inner recesses of

the crevices was interestingly much cooler, at 27

to 28 °C, than outside where the temperatures

could rise to 42 °C in the summer months. Both

Taphozous melanopogon and T. kachhensis

clung to the rock face on all fours. Unlike

T. melanopogon , T. kachhensis was never seen

roosting in temples or other human artifacts.

For experimental recording of the locomotor

activity of both T. melanopogon and

T. kachhensis , which did not fly within limited

spaces but moved swiftly and laterally on all fours,

we devised sturdy wooden tilting cages of 50 cm

x 30 cm x 20 cm with a sliding door at one end and

mesh net at the other. The cages were poised on

knife edges and would tilt laterally, picking up the

slightest ambulatory movements of the bats. The

tilts activated the writing stylets of 20-channel A

620 X Esterline Angus Event Recorders (Esterline

Angus Electronics Co., Indianapolis, USA).

Experiments were performed in photographic

darkrooms (chronocubicles) in desired DD, LL or

LD conditions. Red light >630 nm was used to

represent darkness (‘safe light’). The temperature

in the chronocubicles was constant at 28 ±1 °C

and the relative humidity was artificially raised to

60 to 65% for experiments with Taphozous

melanopogon and T. kachhensis.

A small colony of 25 Taphozous kachhensis

bats comprising both sexes was maintained in an

outdoor bat enclosure of 7.5 m x 3.0 m x 3.75 m

with a fishpond of dimensions 4. 1 5 m x 2.39 m x

0.64 m filled with water. Frogs in good numbers

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were introduced into this pond. The water in the

fishpond helped to increase the relative humidity

inside the enclosure. The sides of the enclosure

were limited on all four sides, by walls 2 m high

and the enclosure was roofed over by steel rods

forming a grating, which permitted the air

movement but was too restrictive for the bats to

fly or squeeze out. The enclosure was in the midst

of a mango grove in the Botanical Garden. A row

of Polyalthia longifolia trees on the east side of

the enclosure, and mango trees on all sides,

provided shade in the early morning hours. At

the east end of the pond was a dark wooden

enclosure in which bats roosted during daytime.

The humidity varied from 35 to 65% and the

temperature from 22 to 30 °C. This outdoor bat

enclosure was our own idea and the outfit came

in handy for the ‘Prey capture by the false Indian

vampire bat’ experiments (Habersetzer and

Marimuthu 1986, MarimuthuandNeuweiler 1987)

and for video recording flight patterns of our bats.

The onset of flight activity following the sunset

and end of activity of the group of captive

T. kachhensis bats were monitored for a whole

year from January 1979 to January 1980. These

bats flew around and fed on insects attracted to a

mercury lamp (125 W) mounted within the

enclosure. The bats drank the water from the pond.

Hipposideros speoris. Body weight 6.5-

7.0 g, best hearing frequency 137 kHz.

This bat typically forages close to the

canopy, around bushes and trees and very close

to obstacles. A colony of 550-600 bats inhabited

a true cave, which was 40-45 m deep in some of its

pockets, in the Jain Hills. Actual observations of

emergence flight of bats and their return were

made at this site (Marimuthu 1984). In a nearby

cave, which was more a hollow in a rock-front,

lived a much smaller colony of c. 50 bats of

Hipposideros bicolor fulvus. The mortality after

capture of these bats was so high that it was

decided that G. Marimuthu and Dilip Joshi would

work with the sturdier H. speoris. As a result, we

have accumulated a wealth of information about

the biology, behaviour and circadian rhythms in

this species. These bats also lived for long periods

of time in captivity in good health.

Freshly caught bats were brought to the

laboratory and placed in a 1 m x 1 m x 1 m nylon

mosquito mesh cage for a week to allow them to

acclimatise to the laboratory conditions. These

bats spaced themselves out without ever

clustering. In fact, each male bat had his ‘personal’

space, inside the cave as well as in the laboratory,

which he would defend from intruders. The

average space between two neighbouring male

bats was 18 ±3 cm (n=12 observations on 46

animals) (Selvanayagam and Marimuthu 1984).

Females seem to choose their roosting position

in relation to males. Males, in addition to urine-

marking their personal space, also adhere to a

strict , hierarchy in their roosting

(Chandrashekaran, unpubl.).

For the experiments, the bats were brought

into the chronocubicles and housed in flight

activity cages of dimensions 30 x 30 x 30 cm, one

bat in one cage. The cages had light aluminium

frames and were covered with mosquito mesh

netting and suspended from firm arms of tripod

stands with strips of spring. H. speoris bats

resorted to brief bursts of sustained flight and

such activity jiggled the cages. The vertical

oscillations and displacement of the cages

depressed microswitches activating the stylets

of Esterline Angus Event Recorders.

In the cave experiments performed by my

students, G. Marimuthu (from 1978 to 1983) and

Dilip Joshi (from 1980 to 1985) the same actograph

(activity recording device), as described above,

was used, employing a hand-wound Lambrecht-

KG-Gottingen thermohygrograph instead of the

electricity-run Esterline Angus Event Recorders.

The thermohygrograph completing one

revolution in 24 hours was adopted for tracing

activity/rest patterns by writing ink stylets fixed

on the flank of activity cages. The sturdy set-up,

shown in Plate 1 , Fig. 1 , was placed 40 m deep into

the cave. The activity/rest patterns of three bats

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Chandrashekaran, M.K.: Thermohygrographs

Plate 1

Fig. 1: The experimental set-up of suspended flight activity cages

and hand-wound thermohygrographs

Fig. 2: Thermohygrographs placed in different parts

of the Rhinopoma hardwickei cave

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could be simultaneously traced, the longest

experiment lasting up to 60 days. The ambient

temperature in the chronocubicles was 28 ±1 CC

and relative humidity was maintained high at

85 ±5% (since the relative humidity in their cave

is 90% or higher).

Rhinopoma hardwickei’. Body weight 14-

15 g, best hearing frequency 35 kHz.

Rhinopoma hunts flying insects at medium

heights of up to 10 m above the ground and

always keeps away from dense background. A

stable colony of c. 1 ,500 bats of both sexes of the

mouse tailed bat Rhinopoma hardwickei roosted

in a narrow cave with high ceiling in the western

flank of the Nagamalai Ridge, which forms the

backdrop to the MKU campus. The population

was intensively studied (Usman et al. 1990) for

emergence and return flight patterns from August

1978 to August 1979. Additional observations

were made during the period January to December

1980. The temperature and relative humidity in

the various pockets of this irregular cave were

measured using thermohygrographs. Wind

conditions were recorded with a portable wind

meter (Lambrecht KG Woelfle type) erected on a

tripod stand in the study area. Sunrise and sunset

times for all our field experiments were obtained

from the Indian Ephemeris Nautical Almanac

published by the Director of Observatories,

Kolkata and were adjusted for latitude, longitude

and Indian Standard Time. No laboratory

experiments were performed either with

Taphozous kachhensis or Rhinopoma

hardwickei.

Results of Field Ethology

and Laboratory Studies

Activity and roosting patterns of a colony

of Taphozous melanopogon : Voute et al. (1974)

and DeCoursey and DeCoursey (1964) had

observed that members of their bat colonies of

Myotis spp. crowded at the entrance to their roost

before flying out in the evenings to forage. In

Germany, Rudolf Rubsamen and Michael Eckrich,

who had each spent a few weeks and a whole

year respectively at MKU, took me to the

countryside churches in Upper Bavaria to see

huge colonies of Myotis 'myotis roosting there.

At sunset, these bats came to ledges from where

they appeared to be ‘sampling light’ for close to

40 minutes in the long twilight of the higher

latitude, before taking off. Such Tight sampling’

behaviour could not be established for

T. melanopogon. On the other hand, these bats

appear to be exposed to the ambient light all

through the day and could perceive nightfall,

which came in a matter of 6 to 8 minutes in Madurai,

directly. The time of the first flier could be

generalised as the exact time of onset of foraging

flight activity of the colony, for hordes of bats

followed within seconds in clusters of 6 to 8.

These observations were made at intervals of 10

days during January 1975 to January 1976. The

difference between the longest and shortest days

in Madurai is well below 2 hours. The entire colony

empties out within 1 8 to 2 1 minutes, indicating

great inter-individual synchrony in onset of

activity but not in its termination, as shown in

Fig. 1 . Even though the time of sunset during the

course of the year varied over a range of 41

minutes, the bats showed astonishing rigidity in

the time of their emergence which was restricted

to a narrow ‘gate’ of 16 mins, where the first bat

emerged between 1 825 hrs and 1 841 hrs. The onset

of foraging flight of the first ‘sentinel’ bat under

field conditions, plotted against the months of

the year, is illustrated in Fig. 2. Each dot of the 2

sets of 36 dots represents single observations on

the ‘onset’ and ‘end’ of the activity of the bat

colony made at 1 0-day intervals from January 1 975

to January 1976.

The precise timing of emergence of the bats

is well illustrated by the following incident.

Eberhard Gwinner (Director MPI for Behavioural

Physiology, Andechs, Germany) was visiting me

in 1977 and wanted to see for himself this great

precision in the onset of the outward foraging

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1 50— s

w

ro

JD

0

XI

E

3

z

100'

50-

H - Time taken for first bat to last bat to emerge (mere 22 minutes)

1ZZZZ3 = Time taken for return flight (spread over 6 hours and 14 minutes).

Fig. 1 : Pattern of ‘onset’ and ‘end’ of the nightly foraging flight of members of a colony

of Taphozous melanopogon (After Chandrashekaran et al. 1983). Actual numbers flying out and returning do

not tally since the bats took different routes, especially during the onset of foraging flight.

1975 JAN FMAMJ JA SON DEC

MONTH OF YEAR

O = Onset of activity, E = End of activity.

SS = time of sunset, SR = time of sunrise.

Fig. 2: Field data of flight activity of Taphozous

melanopogon bats (After Subbaraj and

Chandrashekaran 1977)

activity of the colony. We took him to the site.

Gwinner lay on a rock facing the crevices and was

impatiently looking at his watch and called out at

1826 hrs, “Where are your bats?”. The exodus

started within seconds of the question and

Gwinner was greatly impressed.

The timing of end of activity was imprecise

in all seasons. The narrow gate in the timing of

emergence flight implies that there is a seasonally

changing threshold in sensitivity to twilight. Thus

bats began flying when it was very dark on short

days (0. 1 lux) and flew out in sunlight even as the

sun was going below the horizon (50 lux) on long

days. It is unusual for a nocturnal mammal not to

have a reasonably definable and fixed lower light

intensity as a trigger to nightly activity, and

therefore our findings need to be further

investigated. Our findings are also at variance

with the observation of Brosset (1962a) on

Taphozous melanopogon , “The nocturnal

outings of the colony take place 25 to 30 minutes

after sunset (Kanheri and Mandu)”. But Brosset,

of course, as he himself has pointed out, did not

make a systematic year-round study of these

phenomena.

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Werner Siefer, a project student of

G. Neuweiler, who was visiting us in 1990, declared

one day that he had seen bats ‘soaring’ at heights

of c. 200 m soon after sunset. My response was

that it was not possible for bats to soar. Textbooks

tell us that bat wings are designed for flapping

flight. “To stay airborne, a bat requires 8-15

wingbeats” (Neuweiler 2000). R. Subbaraj was

requested to look into the matter. He accompanied

Siefer with a pair of powerful binoculars. It was a

warm full moon evening and it had drizzled. The

bats were indeed soaring 1 50-200 m above the

Jain Hills rock formation (Siefer and Kriner 1991).

R. Subbaraj identified the species as being

probably Taphozous kachhensis. The bats were

apparently soaring in the thermal layer arising from

the rocks below and cropping a rich fare of insects.

Unfortunately, this interesting phenomenon was

not further investigated, since both Werner Siefer

and Eva Kriner left for Munich soon after making

the discovery. It makes sense to assume that these

bats had switched off their echolocation system

while soaring, and the insects would have been

plentiful anyway. They just had to soar with their

mouth open and close it with a mouthful of insects.

Activity and roosting patterns of a colony

of Hipposideros speoris : The observations on

onset and end of activity were made at fortnightly

intervals from December 1977 to January 1979. It

was possible to count bats flying out in the

twilight, against the still blue sky. The time of

emergence of the first bat was recorded, and

thereafter the number of bats emerging every

minute was counted for as long as the prevailing

twilight permitted. Light intensity was measured

during the evening out-flight at the cave mouth,

using the optometer with the photosensor

pointing to the zenith.

Prior to emergence, approximately an hour

before sunset, bats become restless and exhibit

circular scanning head and ear movements, wing

flapping, elaborate autogrooming, rocking

forward and backward, and brief stretches of

flights. The ear and head movement, noticed

through the noctovision apparatus, may just

mean that these bats were echolocating our

presence in the total darkness of their roosting

site. The commotion inside the cave gradually

builds up, but since adults of H. speoris are “silent”

and only produce ultrasonic pulses of 134 kHz, in

this case there is none of the noisiness associated

with the onset of colony activity in bats of other

species. Flying around of the bats also intensifies

the smell of bat guano inside the cave. When the

light intensity steeply decreases during sunset,

bats fly farther and farther toward the entrance of

the cave. Bats of both sexes live in the innermost

recesses of the cave. The darkness in the deeper

parts is absolute and 1000 seconds exposure of

the photoelement of the optometer on the energy

scala scale did not register any light. About 1 0 to

20 minutes after sunset, in all seasons, a solitary

bat invariably darts out of the cave to return

immediately. We have never been able to verify if

it was the same ‘alpha’ bat that emerged every

evening and banding the bat did not help. A little

before exodus, the entire colony of bats remain

milling around very close to the cave mouth

‘sampling light’ (Twente 1955), then groups of 1 0

to 15 bats break off and fly into the night sky.

However, on any given night, about 3 to 5% of

the bats remained inside the cave. It was not

possible to determine the returning time with any

degree of accuracy, since bats flew in and out

throughout the night. Further, bat mothers

returned frequently to check if their pups were

safe, when they did not carry them to the foraging

sites. Evening departure of bats usually occurred

at low twilight intensities that ranged over the

seasons from 4.5 to 40 lux. Fig. 3 summarises the

findings on H. speoris (Marimuthu 1 984). It is clear

from the figure that the onset and end of colony

activity systematically changed and remained close

to the timings of sunset and sunrise, respectively.

Fig. 4 illustrates the pattern of emergence

flight of the colony of H. speoris made on four

nights, in December 1977 (short daylength), March

(neutral daylength), June (long daylength) and

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October 1978. The bat counts were made every

minute and shown here added up for five minutes.

The pattern of scattered emergence on the night

of June 23, 1978 might have resulted from the

frequent outward and inward flight of suckling

bat mothers. Fig. 5 is a representative example of

- 13

- 12™

-11

C Z3

O O

z

>.

40 'w

(5

20 c J

0 §

100

80 —

60 I

40 I

1977 1978

1979

Month of year

Fig. 3: Summary of the nocturnal activity pattern

of a colony of Hipposideros speoris

for the period of a year (After Marimuthu 1984)

Key to panels:

A: Temperature variations at time of onset of activity.

Upper (solid) circles = temperature maxima,

lower (open) circles = temperature minima,

broken straight line = remarkably constant temperature

of 27 °C inside the cave.

B: Maxima and minima of humidity.

C: Sunset (dotted line) and onset of colony foraging

flight (solid circles).

D: Light intensity.

E: Sunrise (dotted line) and end of colony activity (solid

circles).

F: Relationship between daylength (dotted line) and

duration of activity (solid circles).

end of activity of the colony where numbers build

up with approaching sunrise and terminate

abruptly. The precision in the timing of onset and

end of activity over various seasons in the

colonies of insectivorous bats has been reported

by many authors (DeCoursey and DeCoursey

1964, Erkert 1976, 1978, Griffin and Welsh 1937).

The actual onset of exodus in Hipposideros

speoris is preceded by arousal, which is

obviously accomplished by the endogenous

circadian rhythm acting like a wake-up timer. The

light sampling is actually employed to fine-tune

the circadian clock. A bat that tries to fly out before

its time is exposed to bright light, which causes

the clock to delay its functioning. Similarly, a bat

that does not return to its cave early enough

would be exposed to the bright light of the rising

sun. A light pulse shock, at that phase in the

animal’s clock, ‘advances’ the phase. These

discrete phase delays effected by light during

sunset, and discrete phase advances effected by

sunrise, explain how the bats do not fly about at

unusual hours in nature. This tuning of the

biological clocks by light is called entrainment

(Aschoff 1 960, Pittendrigh 1 960).

Activity time. Marimuthu (1984) has made

a fine structure analysis of the seasonal changes

in the precision of the onset and end of the

H. speoris colony activity under natural

photoperiodic conditions. The duration of activity

of the colony was measured as the time elapsed

between the flying out of the first bat and the

return of the last bat. The duration of activity

followed the seasonal changes in the

photoperiod, being longer over shorter

daylengths and shorter over longer daylengths,

indicating a positive correlation between the

duration of colony activity and the length of night

(Fig. 6). Fig. 7 represents the regression analyses

of the timings of sunset and emergence of the

first bat, and Fig. 8 represents the regression

analyses of the timings of sunrise and return of

the last bat, over the seasons. The lower light

intensities triggering the onset of colony activity

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Fig. 4: Representative examples of the pattern of emergence of members of the colony

of Hipposideros speoris (After Marimuthu 1 984)

Fig. 5: Representative pattern of return of members of the Hipposideros speoris colony over the course of an

entire night (After Marimuthu 1 984)

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Fig. 6: Correlation between activity period

of Hipposideros speoris and duration of night

(After Marimuthu 1984)

varied marginally over the seasons, indicating that

there is no fixed lower threshold. The rate of

change of light intensities during the brief dusk

and dawn appears to be the primary and reliable

environmental cue for these bats to modulate the

onset and end of activity. It was not possible to

precisely determine the beginning of return flight,

because the bats indulged in both inward and

outward flights sporadically during most of the

night. By around 0300 hrs, the flight was

essentially back to the cave and the build up in

numbers was impressive an hour before sunrise.

At this time the bats did not dash into the cave

but undertook circling flights near the mouth of

the cave at an altitude of one or two metres. The

precise function of these manoeuvres is not clear,

but it has been understood by us that the bat was

daily fortifying its place memory. The bats then

ceased to emit the trains of ultrasonic pulses and

literally dived into the cave. These findings on

activity duration, onset and end, relative to the

Fig. 7: Linear regression of time of emergence

of the first H. speoris bat of the colony in relation

to the time of sunset over the seasons

(After Marimuthu 1984)

Time of sunrise (hrs)

Fig. 8: Linear regression of the time of return of the last

H. speoris bat of the colony in relation to the time of

sunrise over the seasons (After Marimuthu 1984)

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daylength (photoperiod) are in accordance with

the rules formulated by Aschoff (1960, 1969) for

diurnal and nocturnal animals, known as

Aschoff’s Rule (Pittendrigh 1960). Marimuthu’s

(1988) studies of these parameters in H. speoris

were the first of their kind on a tropical bat species.

Activity and roosting patterns of a colony

of Rhinopoma hardwickei : A number of caves,

caverns and crevices that accommodate bat

colonies exist in the Nagamalai Ridge. There, we

decided to study a colony of Rhinopoma

hardwickei which in many ways was different

from Hippos ideros speoris. The first feature I

noticed was that there was an unmistakable smell,

and that the members of the colony were found in

all parts of the cave and roosted in different parts

of the cave at different hours of the day. There

certainly was nothing like ‘personal space’ or a

marked hierarchy. These bats have two kinds of

roosting positions, either resting on all fours like

Taphozons spp. or hanging free like Hipposideros

spp. The cave was more like a crevice and its

topology is shown in Fig. 9. During field

observations 1 noticed that most bats of the colony

were close to the spacious high-roofed (c. 12 m)

cave mouth which faced the west, in the cooler

hours of the morning. The bats frenetically waved

their prehensile and longish tails by means of

which they detected and crept into narrow cracks

in the wall of the cave. Plate 1, Fig. 2 shows the

thermohygrographs placed at different positions

inside this cave. With the help of special,

elongated thermal probes, we could measure the

temperatures prevailing in nooks and comers. The

bats roosted in big clusters of 30-40 animals and

the clusters moved progressively inwards in the

cave, which was very dark and very cramped —

in places, less than 1 metre wide — for the observer

to position himself. By evening they had all

crouched in inaccessible recesses and cracks in

the inner walls of the cave. Careful measurement

of the ambient temperature at the sites in which

they roosted indicated that the bats were moving

to zones of constant temperature of 27-29 °C. A

rough sketch about how a cluster of bats moved

on a hot May day is shown in Fig. 10. In

R. hardwickei , constancy of temperature of

c. 27 °C was possibly being sought by the bats

rather than absence of light. This might also be

the situation with Taphozous melanopogon and

T. kachhensis, with none of the three species

having well-marked ‘personal space’.

FRONT VIEW

Fig. 9: Topology and dimensions of the cave inhabited by the colony of Rhinopoma hardwickei

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The Rhinopoma colony was also the

biggest in terms of numbers. Prior to exodus flight,

the members of the colony indulged in intense

audible and shrill vocalisation which was literally

amplified at the cave mouth. Fig. 11 summarises

patterns of onset of colony activity accompanying

sunset and return of the bats after foraging (Usman

et al. 1990). The exodus flight is impressive, like a

cloudburst, with the members scattering to fly to

their foraging sites. Emergence of the colony kept

pace with the seasonally varying time of sunset

and the ambient light at the cave mouth, which

varied between 1 .0 lux during shorter days to 60

lux during the longer days of summer.

One striking feature, which is common to

the nightly emergence of the colonies of

T. melanopogon, H. speoris and R. hardwickei,

is that the members of the colonies pour out

synchronously. However, the return flight is

spread over the night and not predictable. Based

on such observations, we postulated

(Chandrashekaran et al. 1983) that the arousal

and onset of activity in these animals might be

under the control of the circadian clock, whereas

the end of activity may be determined by external

environmental factors such as wind direction and

speed, heavy precipitation and satiation. The most

spectacular ‘emergence by coup’ of bats I had

witnessed was on January 12, 2002, in Thailand. I

was driven from Bangkok, to a place called Phu

Phaman in Khon Kean province, some 450 km

northeast, to watch a colony of bats emerge for

their evening foraging. My informants themselves

had not seen the phenomenon and therefore

could not prepare me for the sight awaiting us.

We arrived at the impressive cave site situated in

a massive rocky hill with a yawning mouth c. 50 m

high and 30 m across. There were eight tourist

buses and hundreds of tourists who were all

gazing at the cave. At 1 745 hrs, the crowd gasped

and we saw a thick black cloud ooze out of the

cave and stream along the left flank of the rock at

a height of 1 50 to 1 70 m. The bottom ledge of the

cave mouth was at an elevation of c. 80 to 100 m.

i

v-

z>

O

HI

§

o

Encircled numbers indicate positions of roosting at given time: 1. 0600-0900 hrs, 2. 0900-1200 hrs,

3. 1200-1500 hrs, 4. 1500-1800 hrs

Fig. 10: Rough illustration of daytime movements of a cluster of Rhinopoma bats

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300-1

200’

w

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Z

100'

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- 200

- 100

L. o

Outward flight of colony lasted only 26 minutes (1846 to 1912 hrs) on the evening of counting but return lasted

virtually the rest of the night.

Fig. 1 1 : Pattern of ‘onset’ and ‘end’ of the nightly foraging activity of the colony of R. hardwickei

(After Chandrashekaran et al. 1 983)

They looked, at that distance, like a swarm of bees,

but too numerous to count or even guess the

numbers. The sun had not set and several kites

were gliding aloft. The outflow of thousands upon

thousands of the bats continued steadily and

lasted seventeen minutes when I saw the last of

the millions of bats fly out at a great height, but

this time headed in our direction. We were

fortunate that a juvenile bat, possibly on his first

evening out, fell to the ground close to me. I picked

it up, examined it and placed it on a nearby rock.

Given its resemblance to Tadarida aegyptiaca in

Madurai, I presumed that the Thailand bat was

Tadarida brasiliensis. The fur was dark grey, the

bat was wrinkle-lipped, and had a snout like a

mastiff. If the species was indeed Tadarida

brasiliensis , then they were all flying out at great

speed of 27 m/s, the fastest bat flight known

(Neuweiler 2000). The temperature was 26 °C and

darkness descended very soon. Dr. Manjunatha

Rao, who took me to the Phu Phaman caves, writes

that there are several other caves nearby. I was

thrilled to see lovely, stylised statues in bronze,

of the locally available bat species, inscribed with

the Latin names.

Social synchronisation of circadian rhythms

in Hipposideros speoris: An interesting feature that

we noticed in the exodus flight of the H. speoris

colony was that bats inhabiting the innermost

recesses of the cave, mostly females and subadult

males, were the earliest to fly out. A group of Dutch

scientists (Voute et al. 1 974) had noticed a similar

phenomenon in the evening exodus flight of a colony

of My otis dasyncheme. A major question was: how

do bats inhabiting the depths of our natural cave of

perpetual darkness, invariant temperature and

relative humidity — an environment virtually devoid

of time cues — know the time of sunset in Madurai?

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We made the first observations inside the cave

(Marimuthu et al. 1978) with the help of infrared

noctovision binoculars. The individuals of the

colony were spaced out and hanging by their hind

feet from the ceiling. If we approached them, they

turned their heads in all directions with quivering

ears and flew away if we went too close. But when

Marimuthu and I sat still in the afternoon hours, the

bats became still and the colony appeared to be in a

state of deep rest (sleep?) until c. 1700 hrs. Soon

after, the bats appeared to show signs of arousal

and individual bats stretched their wings, began

preening themselves, yawned and began flying

about, one by one. Deep in this cave, where we

made our observations, the gurgle of an unseen

stream of water could be heard. Happenings

subsequent to ‘arousal’ of the colony, such as bats

flying to the light sampling chamber, and eventual

exodus flight of the members of the colony soon

after sunset have been described under ‘Activity/

roosting patterns of a colony of Hipposideros

speoris .’

The next question to be tackled was

whether each bat had to ‘see’ for itself the

darkening sky outside. If some bats did not come

to the cave mouth to sample light, would they still

know that the sun has set? Do the volunteer bats

relay the information to bats deep inside the cave?

We performed the first experiment in which we

kept three male bats captive at a depth of c. 40 m

in flight activity cages (described earlier) for an

extended period of 50 days and the bouts of their

flight and rest were continuously recorded on the

charts of hand-wound thermohygrograph drums

described earlier. It soon became apparent that

the captive bats began their flight activity at the

precise time at which the free-flying conspecifics

began their evening exodus flight. The actogram

describing the flight activity/rest patterns of the

three captive male bats for 40 days in one case,

and 50 days in the other two cases, is presented

in Fig. 12 (Marimuthu et al. 1981). The 24-hour

activity/rest strips were pasted on bristol board

one below the other chronologically. Activity

bouts are indicated by the vertical patches and

the horizontal traces indicate rest.

The captive bats were less active when the

cave emptied out, but they responded to stray

returning bats and to the flock of bats returning

in the small hours of the morning. Our excitement

was great, for we had confirmed that the free flying

bats were telling captive bats the time. Our first

communication (Marimuthu et al. 1 978) was a very

short one of less than 400 words based on activity/

rest patterns of one captive bat recorded for a

mere eleven days inside the cave. Confirmatory

evidence of social synchronisation (that is the

scientific term) of the circadian rhythms in

H. speoris came with a later paper (Marimuthu et

al. 1 98 1 ). We had also stated that it was not clear

to us how the phenomenon took place

and suggested involvement of: 1) pheromones,

2) wing flapping noise generated when

conspecifics flew out, and 3) acoustic

transmission of message.

Even though we were reasonably sure that

there was social information of sunset and time of

day, we wanted to demonstrate the opposite

situation also, i.e. a solitary bat in a solitary cave

without ‘social informers’ cannot synchronise its

circadian rhythm to the 24-hour periodicity of bat

colony activity. A practical problem was finding a

cave good enough to be habitable for bats but

still not colonised. When, finally, we did find a

solitary cave without hipposiderid bats, we

performed an experiment by placing a solitary male

H. speoris in a flight activity cage and recorded

its flight activity/rest patterns inside the cave.

The solitary bat was indeed helpless in the strict

24-hour periodicity of the onset and end of its

activity, as is shown in Fig. 13. The circadian

activity rhythm in the flight activity free-ran with

a period < 24 hours in the continuous darkness of

the cave (Marimuthu et al. 1981). The bat began

its nightly flight c. 20 mins earlier each subjective

evening. We terminated our experiment with a

touch of bravura. After 50 days in captivity and

free-run of its circadian rhythm, our bat began its

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1200 1800 0000 0600 1200 1200 1800 0000 0600 1200 1200 1800 0000 0600 1200

Time of day (hrs)

Fig. 12: Actogram showing flight activity patterns of three captive male Hipposideros speoris bats

(After Marimuthu et al. 1981)

nightly activity just when other free flying bats

were returning to the roost in the early hours of

the civil morning.

Please note that, as shown in Fig. 13, the

bat started its nightly activity close to 1 900 hrs

which is near civil sunset time. On a daytime

inspection on day 10, it turned out that the cave

was not impervious to bird calls (crows and mynas

at a nearby watering hole), at night resident

crickets stridulated. It looked as if the solitary bat

was taking time cues from these acoustic inputs,

but I was sceptical of such interspecific acoustic

entrainment and we continued the recordings. To

our surprise and excitement, the circadian rhythm

slipped into a state of free-run (as explained

above) from day 1 1 onwards (as seen in Fig. 13).

The obvious interpretation is that birds and

crickets cannot entrain the circadian rhythm of

this captive bat. These observations led us to

another major question. Do these bats need to be

told the time by other bats of the same species?

In other words, is social synchronisation in

Hipposideros speoris species-specific?

There have been interesting reports of

species-specific entrainment of perch hopping

rhythmicity of the common sparrow Passer

domesticus. Male courtship vocalisations were

played back to female sparrows held in continuous

dim light for four and half hours in a 24-hour cycle,

and the perch hopping, free-running circadian

rhythmicity entrained to song/silence cycles

(Gwinner 1966). Similar results came in for two

other species of birds the same year and these

results generated much excitement for their

ecological and behavioural implications. However,

a year later it was found (Lohmann and Enright

1967), rather unromantically, that cycles of

mechanical noise administered by a loud buzzer

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1200 2000 0400 1200 2000 0400 1200

Time of day (hrs)

Fig. 13: A double plot of activity/rest patterns of a

solitary male Hipposideros speoris bat recorded in an

empty cave without any conspecifics over a period

of 50 days (After Marimuthu et al. 1981)

for a few hours in a 24-hour cycle, also entrained

the activity rhythms of three species of passerine

birds. This is a demonstration of the phenomenon

of non-species-specific entrainment of circadian

rhythms at its best.

To return to bats, in order to investigate the

possibility of species-specificity in social

synchronisation of circadian rhythms in bats, we

performed an ingenious experiment. We

introduced a rank outsider bat, an alien species, a

male Taphozous kachhensis into the

Hipposideros speoris cave in the Jain Hills, and

recorded its flight/rest pattern while captive. The

circadian rhythm of flight/rest activity of the alien

could not be socially (or otherwise) entrained by

the 550-600 hipposiderid residents (Marimuthu

and Chandrashekaran 1983a). The circadian

rhythm of the emballonurid T. kacchensis

impressively free-runs as shown in Fig. 14. We

interpreted the results as indicating a kind of

communication gap between the hipposiderids

and the alien emballonurid. We can back up claims

from the results of neurophysiological experiments

in which we placed electrodes in the lower

colliculus of a T. kachhensis bat in the operation

theatre and played back the colony vocalisations

(including ultrasonic acoustics) of H. speoris

(using a homemade Lennartz tape recorder of 2-

200 kHz range). The emballonurid bat (under mild

nembutal narcosis) showed no spike (action

potential) responses. The message was not even

being heard. These results strongly suggest that

social synchronisation of circadian rhythms in

microchiropteran bats may indeed be species-

1200 0000 1200 0000 1200

Time of day (hrs)

Hatched area at bottom describes hours of

exodus activity during early night and returning

of the resident, free-flying Hipposideros speoris

bats.

Fig. 14: Double-plotted actogram illustrating the free-

running of the circadian rhythm in the flight activity

pattern of an emballonurid bat Taphozous kachhensis

confined to a hipposiderid (H. speoris ) cave for 60

days (After Marimuthu and Chandrashekaran 1983a)

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specific. The interesting field observation is that,

in some places, H. speoris and T. kachhensis are

known to share roosting sites (pers. observ., also

Brosset 1 962).

We have established for the first time that:

1) the circadian rhythm of a captive bat held in

isolation in perpetual darkness free-runs, and

2) in the presence of free flying conspecifics which

undertake foraging flights out of the cave and

back, the circadian rhythms of captive bats entrain

to the 24-hour periodicity of flight/rest of the

colony. Thanks to the light/darkness of the natural

environment (ubiquitous entraining agents) there

are no free-running rhythms out there in nature,

except in deep-sea organisms and those living in

caves (Koilraj et al. 2000). Circadian rhythms are

also known to free-run in organisms in the Arctic

winter (of perpetual darkness) and summer (of

continuous light) (Bunning 1973). We were

curious to know how bats dwelling in the darkness

of caves and foraging in further darkness of nights

would respond to exposure to continuous light

— an admittedly artificial condition that these

tropical bats never face. Artificial light was created

with car batteries and an incandescent bulb inside

the Hipposideros speoris cave, and the flight

activity/rest patterns of three male captive bats,

in activity cages placed in the vicinity of the light

bulb, were measured. The ambient light intensity

at the level of the cages was between 5 to 1 5 lux.

It can be seen from Fig. 15 that the circadian

rhythms in the activity/rest patterns in all three

bats free-ran (Marimuthu and Chandrashekaran

1983b) with a period > 24 hours. Continuous light

lengthens period and perpetual darkness shortens

it, in these dark active animals, effects that have

been codified in ‘Aschoff’s Rule’ (Aschoff 1960,

Pittendrigh 1960). A careful examination of Fig. 15

will reveal two ‘conflicting’ components. All three

bats do stir about briefly during the colony’s

exodus flight, but lapse into sleep and begin and

end activity according to their endogenous

circadian free-running schedule with a period >24

hours. The brief arousal during colony exodus

coinciding with sunset is called a positive masking

effect (Aschoff et al. 1982), and the free-run, the

true expression of the circadian clock, which is

apparently uncoupled from the 24-hour social

synchronising inputs, by the artificially created

continuous light. In spite of close to ten years of

working on the social synchronisation of circadian

rhythms in H. speoris we are still, quite literally, in

1200 0000 1200 0000 1200 1200 0000 1200 0000 1200 1200 0000 1200 0000 1200

Time of day (hrs)

Fig. 15: The flight activity patterns of three captive male H. speoris bats for durations of 55 days (a and c)

and 39 days (b), registered in LL of 10 to 20 lux (After Marimuthu and Chandrashekaran 1983b)

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the dark about the exact nature of the social cues

(ultrasonic acoustics? pheromones? wing-

flapping noise of free flying bats?) behind this

kind of entrainment.

One of the earliest reports to impute social

synchronisation among conspecifics was for mice

of the genus Peromyscus (Halberg et al. 1954).

Similar effects have been subsequently claimed

for blinded mice, male chevrotain antelopes, wolf-

coyote hybrids, beaver colonies of Castor

canadensis , macaques and sexual cyclicity of

female mammals (for an early review, see

Chandrashekaran 1982). Social synchronisation

of circadian rhythms deserves to be better studied,

preferably using social insects such as

honeybees and ants (Frisch and Aschoff 1987).

A daylight PRC for the circadian rhythm

of Hipposideros speoris

The ‘phase response curve’ (PRC) is a plot

of the responses of a circadian rhythm in terms of

phase shifts to perturbations (of light,

temperature, and chemicals) as a function of phase.

A vast body of scientific literature has

accumulated on this subject. A PRC informs the

state of sensitivity of the basic oscillator

(biological clock) at any given phase to the

zeitberger stimuli (perturbations). Such

information cannot be had, for instance, when we

record discrete events such as locomotion and

rest, which start and end abruptly. There is no

clue to the state of the clock during hours of rest

when nothing happens outwardly. Further, all

external events, both observable and measurable,

may be of the nature of the ‘hands of the clock’.

Pittendrigh ( 1 960) stated that the Drosophila PRC

reflected the time course and waveform of the

basic oscillator. All experimental studies to

construct PRCs have been made with fluorescent

and incandescent light (Chandrashekaran 1998).

We therefore constructed a daylight PRC for the

circadian rhythm of the cave dwelling bat

Hipposideros speoris, employing brief pulses of

daylight for perturbations and performing the

arduous experiments inside a natural cave. The

dimensions of this cave were less cramped (12x8

x 2.5 m), with uneven walls, ceiling and floor. The

5 original inhabitant bats ( H . speoris) were evicted,

and the cave was fitted with blinds and lightproof

doors. Temperature was constant at 28 ±1°C and

relative humidity of 85 ±5% prevailed. Employing

methods described earlier, the flight activity/rest

patterns were recorded on Lambrecht-KG-

Gottingen thermohygrograph drums. The bats

were brought to the cave mouth (close to the

hinged door) at different phases of the circadian

rhythm and exposed to diffuse daylight of c. 1 000

lux for 1 5 minutes. Fig. 1 6 illustrates atypical delay

phase shift and a typical advance phase shift of

the flight activity, and Fig. 17 illustrates the PRC

obtained on many bats. The daylight PRC we

obtained resembles those made for other

organisms using fluorescent and incandescent

light (Joshi and Chandrashekaran 1 983).

Daylight dimmer than starlight entrains

Hipposideros rhythm: I report an accidental

finding which we later investigated at some

length. This happened in the cave inside which

Dilip Joshi had worked out the PRC. Another

student, S. Raj an brought in a chart depicting the

flight activity of a solitary male H. speoris for a

period of 47 days under absolute darkness,

constant temperature and relative humidity. The

activity started at around 1 900 hrs, evening after

evening, and stopped a little before sunrise. The

calculated period length was exactly 24 hours,

unheard of in the literature on circadian rhythms.

This made me write an eccentric paper, ‘An

unusual circadian rhythm with a precise 24-hour

period’ (Chandrashekaran 1981). I philosophised

“The law of parsimony dictates that we consider

our 24-hour bat as an isolated instance of a

circadian system quite accidentally possessing a

very uncircadian circadian rhythm!”

Then Dilip Joshi brought in 4 or 5 other

cases of bats with precise 24-hour periods. To

make a long story short, light was apparently

leaking in. Careful scrutiny revealed a crack in the

uneven ceiling of the cave through which very

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a Time of day (hrs)

1200 0000 -|200

a. Illustrates a delay phase shift of 63 minutes at 90°

phase of light exposure.

b. Illustrates an advance phase shift of 42 minutes at

225° phase of light exposure.

V = light exposure

Fig. 16: Phase shifts caused in the flight activity

rhythm of two male Hipposideros speoris bats

by 15 mins exposure of the animals held in DD

to diffuse daylight of c. 1000 lux

(After Joshi and Chandrashekaran 1983)

dim light in the range of 0.0001 to 0.0006 lux

streamed in. Due to the inclination of the crack

the light came in for just c. 90 to 1 00 minutes during

midday hours. Light intensity was measured (with

a UDT optometer on the log-scale, i.e. time x

intensity) where it shone the brightest on the floor

of the cave.

Two experiments were performed with 4 and

3 bats. All these bats entrained. Fig. 18 shows the

pattern of entrainment of two bats used in the

first experiment over periods of 39 days (Fig. 1 8a)

and 35 days (Fig. 18b) respectively, in response

to very dim light of 0.0001 to 0.0006 lux illumining

the bats for c. 1.5 hours. Onset of activity

coincided with the local sunset time. Results of

Open circles = Mean; Vertical lines = SD. n = 4 or 5

Fig. 17: PRC obtained for the circadian rhythm in the

flight activity of Hipposideros speoris in DD inside a

cave on several bats, and for a solitary bat (PRC in

inset), over a protracted period of 156 days with 15

minutes daylight of c. 1 000 lux

(After Joshi and Chandrashekaran 1983)

the second experiment confirm that the

entrainment was indeed effected only by the dim

light. Fig. 19 shows entrainment of the flight

activity in 2 bats during the first 18 days. Then

the light leak was plugged. In one bat the free-run

of the rhythm set in immediately (Fig. 1 9a) with a

period < 24 hours, and the other bat continued in

the entrained rhythm for 2 weeks before its rhythm

ffee-ran with a period >24 hours (Fig. 1 9b) (Joshi

and Chandrashekaran 1982). An interesting

feature of entrainment in this case was that even

though the bats experienced exposure to the dim

light at midday, the onset of activity coincided

with local sunset time 6 to 7 hours later. Obviously,

the light pulse in addition to entraining also

influences the phase angle. In other words, the

bats recognised the dim light to be midday light.

We had speculated that this phenomenon might

have an adaptive value.

The lowest intensity of light reported in

literature as entraining a circadian rhythm (in the

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Time of day ( h r s )

1200

r

0000

1200

v % ♦ % n ♦. I

• • * m * I

«* * * ♦ * — k 1

- ? f

1 ♦

k ** J. « •* .1

I • i •*. # .

3vv-#v

♦ «

<• .

• #♦

A •> «.

Us «4Pi sAllll ||

— ■ k » a iw*arr>n»

41 . *

- w *

i v

^anfcatWiMMTrtrw^mri.iim'fe i

b

Fig. 18: Entrainment of the flight activity rhythms of two male Hipposideros speoris bats inside a natural cave

for 39 days (a) and 35 days (b) in response to c. 90 mins of dim light of 0.0001 to 0.0006 lux streaming in

at midday hours (box at top) (After Joshi and Chandrashekaran 1982)

sensitivity of the median eye of the scorpion

Androctonus australis L.) is 0.00025 lux shone

continuously for 16 hours (Fleissner 1977). It must

be pointed out that the actual light intensity at

the level of the flight activity cages, positioned

approximately 2 to 3 m away from the site of light

measurement, was beyond the sensitivity of our

optometer. The intensities of daylight that entered

the cave were about 5% to 30% of starlight

intensity. These are to date the lowest intensities

of light implicated in entrainment of circadian

rhythms. We have also constructed PRCs for

ultra-short light pulses of 0.0625 milliseconds, for

the circadian rhythm in the flight activity of

Hipposideros speoris , which are the shortest light

pulses in literature, demonstrated to shift phase

(Joshi and Chandrashekaran 1985a).

Spectral sensitivity of the photoreceptors

in H. speoris : Do the bats have colour vision?

We performed a series of experiments to study

the spectral sensitivity of the photoreceptors

responsible for phase shifting the circadian

rhythm of flight activity in the bat H. speoris. In

nature, these bats exist in a strange paradoxical

light-darkness regimen experiencing absolute

darkness during the daytime inside natural caves

where they roost, and some amount of light

(starlight, moonlight) during the night when they

forage in the open. They are thus exposed to

skeletal pulses of dim twilight of 4-40 lux in

intensity, during dawn and again during dusk on

the same day, that constitute the major phase

resetting stimuli responsible for entrainment by

natural light.

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Time of day (hrs)

1200 0000 1200 1200 0000 1200

Fig. 19: Pattern of flight activity rhythms of two captive male Hipposideros speoris bats inside the DD of a

cave for 60 days. Entrainment ensues for 1 8 days (a) and 33 days (b) in response to very dim light streaming

through a crack in the ceiling. The leak was fixed on day 1 8 (indicated by arrow)

(After Joshi and Chandrashekaran 1982)

We have monitored the phase shifts evoked

when the circadian rhythms, free-running in

constant darkness, were exposed to 15 minute

and 2.77 hour pulses of monochromatic light (1 00

pW/cm2) at various phases. Fig. 20 illustrates the

flight activity record of a bat in DD for 275 days,

exposed at various phases to monochromatic light

pulses. The marathon experiment lasted 275 days

and the actogram shows long lasting changes in

period that followed some phase shifts. For these

experiments, four phases were chosen to

investigate the wavelength dependent phase

shifts, and we reported that light pulses of 430

and 520 run unequivocally delay (at CT 1 8 hrs)

and advance (at CT 4 hrs) the phases, respectively.

It was postulated that there might exist two classes

of photoreceptors in the retina of H. speoris. The

S photoreceptors (short wavelength sensitive)

having a maximum sensitivity at 430 nm and the

M photoreceptors (middle wavelength sensitive)

having maximum sensitivity at 520 nm, that

mediate delay and advance phase shifts (Joshi

and Chandrashekaran 1985b). This is illustrated

in Fig. 21. The data presented in the figure was

derived from 156 phase responses for 36 bats;

for each point at least four measurements were

taken.

Prey capture by the false Indian vampire

bat: Megaderma lyra (Body weight 3 1 -35 g, best

hearing frequencies 1 1-65 kHz)

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i 1 — I J I

1200 0000 1200 0000 1200

Time of day (hrs)

Fig. 20: Flight activity record of a male IT. speoris

bat held in DD exposed to monochromatic light

pulses of higher energies of 100 pW/cm2for

durations of 2.77 hours given at different phases

(pulses 2 to 1 6). Horizontal arrows to the right

indicate the phases at which the light pulses were

given (After Joshi and Chandrashekaran 1985b)

The biggest colony of the false Indian

vampire bat Megaderma lyra lived in a very well

camouflaged cave, on the southern slope of

Nagamalai Ridge to the west of MKU, which was

difficult for humans to approach. This bat is

Advance phase shifts (CT 4 hrs) = solid line, maximum

at 520 nm. Delay phase shifts (CT 18 hrs) = broken

line, maximum at 430 nm.

Fig. 21: The spectral sensitivity curves obtained with

1 5 minute monochromatic light pulses that evoked

approximately 50% of the maximal advance and

delay phase shifts evoked by white light pulses at

CT 4 hrs and CT 1 8 hrs phases, respectively

(After Joshi and Chandrashekaran 1985b)

unique among Microchiroptera because its diet

is eclectic, including small vertebrates, birds,

reptiles and mammals. Our German colleagues

maintain their colony of M. lyra , captured in and

around Madurai, in Munich, offering each bat one

live mouse a day. In our laboratory, we had at

least twelve M. lyra at any given time, let loose in

the outdoor bat enclosure with a well stocked

frog pond (dimensions given under ‘ Taphozous

kachhensis\ These bats, which are the most

ferocious when handled, did their own foraging,

feeding on the frogs that jumped out to the edge

of the pond, or in water. G. Marimuthu has

performed elegant experiments and reported the

unusual modes of prey capture in M. lyra , on land

and in water (Link et al. 1986, Marimuthu and

Neuweiler 1987).

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On land: Six to ten bats were used in the

experiment. The bats generally hung from the steel

rafters of the outdoor enclosure, visible during

night, and inside the darkened wooden enclosure,

during day. These bats need ‘personal’ space but

unlike H. speoris , M. lyra huddle when they are

disturbed in their natural haunts or are feeling

threatened (Chandrashekaran and Marimuthu

1987). Also unlike H. speoris , the M. lyra stirred

out of the darkened wooden enclosure into broad

sunlight and hung from the steel rafters of the

enclosure. Marimuthu released medium sized

frogs on to the sandy floor of the outdoor cage

and made the interesting discovery that only

jumping frogs attracted the attention of the bats.

If the frogs sat still, the bats did not take any

notice of them. The moment a frog jumped, the

bats, in threes and fours, swooped down to catch

it, but only one of them succeeded in getting hold

of the frog by its scruff. The victorious bat stayed

for 2-3 seconds on the ground and used its wings

to properly position the frog in its jaws. Then it

flew up and off, back to its roosting position, to

ingest in the next 3-5 minutes the entire frog head

first, leaving only the hind legs, in interesting

contrast to the tastes of humans in frog

consumption (Marimuthu andNeuweiler 1987).

Marimuthu also demonstrated that dead

frogs tied by twine and dragged over a sandy

surface are promptly captured, but not dead frogs

that are noiselessly dragged over a smooth glass

pane over a thin sheet of water. It is clear that this

gleaner was not detecting the motion of the prey

but was listening to the “zic” noise of 10-15 kHz

that jumping frogs made. This mode of prey

capture by gleaners listening to the rustle of

scampering prey on the ground, without resorting

to echolocation, has been called ‘passive acoustic

localisation’.

In water: There are some species of bats

that preferably hunt over rivers, lakes, and marine

coastlines and take prey from the water surface.

Such species are Noctilio leporinus, N. labialis,

Megaderma lyra and Myotis spp. On field visits

at night time, in areas rich in ponds and small

waterbodies in south Madurai, one often noticed

Megaderma lyra silently and swiftly fly past at a

mere height of c. 1 m, obviously listening for

scampering prey or jumping frogs. Some species

of bats that hunt over ponds, rivers and lakes

have enlarged feet with which they scoop up small

fishes and arthropods from the water surface. An

ideally smooth water surface would act as an

acoustic mirror, making it difficult for echolocating

bats to receive an echo. Water ripples and objects

protruding from the surface create a kind of texture.

Hunting M. lyra seem to detect the protruding

snouts of stationary frogs in water, in this manner.

Ultrasonic recordings indicate that the false

vampire actively echolocate snouts of frogs, most

of which remained strangely motionless

(Marimuthu et al. 1995). While the bat hovers

over the water, its flapping wings fan the air, which

creates ripples in the water all around the snout

of the frog below. Megaderma lyra presumably

detects, using ultrasonics and echolocation, the

outward progression of ripples vis-a-vis the

motionless central protrusion of the snout of the

frog. As soon as the hunting bat splashes into

the water, the other frogs dive deeper into the

pond and swim to safety.

The role of eyes in echolocating bats: The

advantages of echolocation and the glamour

surrounding the subject is such that extensive

work has been carried out on this aspect

(Neuweiler 1990). But surprisingly little is known

about the precise use of the bats’ eyes in vision

and prey capture. A book devoted entirely to bats

(Altringham 1996) makes only one reference (Bell

1985) to the function of vision in prey capture. It

is common knowledge that hearing in micro-

chiropteran bats is much more efficient than

seeing. Morphologically, the auditory regions of

the brain of insectivorous bats are dispro-

portionately large compared to the optic regions.

The auditory regions of the brain are specialised

to receive, process, store and retrieve information

about the environment on the basis of soft echoes.

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We have not conducted specific

experiments to investigate the role of the eyes of

Madurai bats in landing or prey capture. The

Madurai bats forage as efficiently on new moon

nights as they do during a lunar eclipse (Usman

et al. 1980). We, however, have evidence that

Hipposideros speoris and Rhinopoma

hardwickei did forage on insects in the pitch-

black darkness of their caves. This often happened

when there was heavy rainfall outside, coinciding

with sunset and the bats could not fly out. Hordes

of insects were driven into the caves by the gusts

of wind. Link et al. (1986) have reported that

Rhinolophus rouxi, Hipposideros speoris and

H. bicolor approached and seized dead

cockroaches held by forceps, when these were

artificially vibrated. This indicates that any

oscillating movements and not specific aspects

of wing beating were the key stimulators for

catching-behaviour in all three species. Once,

Marimuthu was feeding bats held captive, in

activity cages, 40 m deep into the Jain Hills cave.

He was holding the live cockroaches with the aid

of forceps. The elytra, wings, legs and innards of

the insect had been removed. Even after this

operation the cockroach usually wriggled

between the tips of the forceps. On one occasion,

a free flying H. speoris wrenched a wriggling

cockroach off the forceps. On scanning the ceiling

of the cave with a torch emitting ‘safe’ light of

>630 nm, a male bat was found hanging from the

ceiling, chewing the cockroach. This prey capture

in absolute darkness was obviously accomplished

solely by means of echolocation, in which

H. speoris employs CF/FM signals of 5-10

milliseconds of pure tone of 132 kHz terminated

by a brief FM sweep (Neuweiler et al. 1988). On

the basis of these findings I conclude that the

tiny eyes of echo locating bats such as H. speoris

and H. bicolor are not efficient in prey capture,

and may act as photoreceptors for ‘sampling light’

recurrently every 24 hours, thus entraining their

circadian clocks. Similar views have been

expressed by other bat researchers (Kunz 1982).

Mother-infant relations in Hipposideros

speoris : Bats are altricial, with newborn pups being

highly dependent on the mother. Females lavish a

lot of attention on their young. H. speoris mothers

either left their young behind in a creche in the

cave or carried them to the foraging areas. At first

it appeared that there was no correlation between

the age of the young and the frequency with which

it got carried around by the mother (Marimuthu

1988). A more detailed subsequent study in our

laboratory and statistical analysis (Kolmogrov

Smirnov test,/? = 0.05) on the patterns of mothers:

i) carrying their young and ii) leaving them behind

as a function of the size/age of the young,

revealed that the two distributions (Fig. 22) differ

significantly and that more bat pups are left behind

as they become larger (Radhamani et al. 1 990).

The young that were left behind in the cave

clung on all fours to the ceiling and scarcely

moved. The mothers often returned before

midnight and retrieved them. During retrieval, the

mother moved towards her infant, gently touched

it with her forearm and presented her ventral

surface, especially the pubic region, moving in

such a way that the young could hold on to the

dugs with its teeth. Then the infant bat let go of

its perch on the rock ceiling, nestled under the

wing membrane of the mother, and oriented

towards the mammary glands to suckle. When

satiated, the young fluttered its wings and often

hung from the neck of the mother. During the day

the volant young bats routinely hung from the

necks of the mothers and as the observer moved

closer, they briefly flew away, to reunite with the

mother and assume the same posture in 1 0 to 15

minutes (Marimuthu 1988). In this posture, the

young have been seen to stretch their wings and

simulate flight. I am not sure if any other species

of Microchiropteran bat infants, in the tropics or

temperate climes, use their mother’s neck as a

perch.

Marimuthu (1988) has also reported, from

experiments performed under natural conditions

and under semi-natural conditions in the outdoor

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Fig. 22: The pattern of mother bats carrying their

young (hollow circles) to the foraging areas, or

leaving them behind in the cave (filled circles) as a

function of the size (which corresponds to age) of

the infants (After Radhamani et al. 1 990)

bat enclosure, that female Hipposideros speoris

identify their own pups. In an experiment

performed close to the mouth of the Jain Hills

cave, he placed six pre-volant pups on a cool rock.

Mother bats returned in the pre-dawn hours and

exhibited hovering and circling flight and possibly

emitted trains of ultrasonic pulses of 134 kHz

directed at the pups. The pups became agitated

as though they were responding to the hovering

mother bat, and raised their heads and emitted

audible multi-harmonic squeaks (Habersetzer and

Marimuthu 1986). Infant vocalisations attracted

the mothers, who flew towards them and alighted

close to them. The pups tried to cling to any bat

mother but the mother bats snuggled closer and

retrieved their own young. Mothers located their

infants even when the juveniles were displaced

from where the mothers left them. Behavioural

experiments under both natural and captive

conditions showed that the sound emission of

the pups attract the mothers, but may not contain

sufficient clues for them to correctly identify their

own from among a group of squeaking young

ones. The nuzzling of the mother among the pups

is indicative that olfactory means also mediate

final identification and retrieval (Gustin and

McCracken 1987, Habersetzer and Marimuthu

1 986, Marimuthu 1 988). On many occasions I had

observed infants of Rhinopoma hardwickei and

H. speoris fall to the ground and emit faint but

audible vocalisations, to be retrieved by obviously

their own mothers. So there can be no doubt that

vocalisations are the primary cue for infant

recognition. Isaac and Marimuthu (1993b) have

reported their accidental discovery of how a

mother pygmy bat Pipistrellus mimus responded

to the vocalisations of a one day old infant. A

maternity colony of Pipistrellus mimus roosted

for several years in a sleeve-like tunnel 1 .3 m wide,

0.85 m high and 24 m long with a right-angled bend

and just one entrance on the western side of the

Department of Animal Behaviour and Physiology

building. A small colony of 18 Rhinopoma

hardwickei had also chosen this tunnel as a

daytime roost. Suthaakar Isaac separated a one

day old infant bat from its pipistrelle mother after

tagging her, and took the infant to his work bench

nearly 70 m away from where the mother bat was

roosting on all fours. Pipistrellus spp. never carry

their young to foraging grounds. The infant

separated from its mother was emitting ultrasonic

distress and isolation calls continuously, in the

range of 30 to 80 kHz measured using a Mini-2 bat

detector (Ultrasound Advice, UK). The time of

day was 1 800 hrs to 1 900 hrs when the sun sets

and bat colonies stir out. The mother was

apparently flying around, heard the isolation calls

of her infant and in response flew into the room

through an open window and alighted on the

infant, covered it with her wings, lifted it up and

flew away. The entire rescue operation apparently

lasted only a few seconds (Isaac and Marimuthu

1 993b). Other researchers have also reported that

bat mothers respond to calls of their own young

(Balcombe 1990).

We commented in one of our papers

(Radhamani et al. 1 990), “The present study deals

with mother-young relations in a cave-dwelling

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insectivorous bat Hipposideros speoris and

explains how the mother bats carry even their

volant young to their foraging areas in order to

acquaint them with the topography and foraging

strategies”, and later in the same report, “. . .more

young ones are left inside the cave as they become

larger”. These two statements make, in fact,

contradictory claims. Though volant subadult

H. speoris can produce and hear CF/FM sounds

( 1 27- 1 3 8 kHz) (Habersetzer and Marimuthu 1 986)

in the adult range, they face opposite the direction

of the flight path of the mother while being carried.

Subadult Hipposideros may use ultrasonics to

form an acoustic picture of the interior of the cave

and the immediate environment (Marimuthu 1 988).

In the case of Megaderma lyra , the mothers even

carry infants which are nearly their own size.

These bats are known to have secondary night

time roosts. In Madurai, many of the M. lyra bat

mothers used a cowshed as a night time roost

and left their infants on the ceiling.

Breeding Periodicity of Insectivorous Bats

Breeding pattern of Hipposideros speoris :

Brosset ( 1 962a) writes of the breeding pattern of

H. speoris of the Poona/Khandala region “The

periodicity is absolutely strict for this species,

and all females deliver together in May. The first

to do so were observed around 5th May, and the

last around the 25th of the same month”. In

Madurai, there are two peaks of breeding in

H. speoris , the first and less pronounced peak in

May and the second in November, which is how

Radhamani (Radhamani et al. 1990) could make

her observations on Hipposideros mothers

carrying infants from August to December 1989.

G. Marimuthu’s detailed work on mother-infant

interactions in this hipposiderid bat are based on

observations made from December 1977 to

February 1979 and November 1980 to February

1981 (Marimuthu 1988). Since a single young is

the rule for most species of bats, we can only

conclude that there are female H. speoris that bear

young ones around May and other females that

bear young around September/October. Almost

all species of bats studied by Brosset (1962a, b, c,

1 963) had a very narrow period of parturition. This

is also the case in Madurai in the breeding pattern

of Hipposideros bicolor , which often shared the

same cave as H. speoris. All young of

Hipposideros bicolor were bom in May. The May

peak is the typical breeding pattern to the north

of the equator, and the November peak to south

of the equator. Professor Aschoff (1913-1 998) in

personal conversations had often told me that he

believed that the “biological equator” might be

around eight degrees north of the geographical

equator. Madurai interestingly is 9° 58' north of

the equator.

Breeding pattern of Pipistrellus mimus :

The vespertilionid bat, Pipistrellus mimus

Wroughton, is a commonly occurring and widely

distributed bat in India, except in mountainous

regions. It is physically the smallest species of

bats of India. It is eclectic in the choice of roost

sites and lives inside caves, even in small

depressions in rocks, cracks, crevices, buildings,

inside thatches, tree holes and even in

letterboxes. Adults weigh 3.9 ±0.4 gm (n=19) and

the forearm measures 27.6 ±1.1 mm (n=26). A

group of 20 members of P. mimus colonised the

tunnel in our departmental building described

earlier. Adult females and infants roost in groups,

and adult males roost singly throughout the

length of the tunnel. The study was carried out

over one whole year, from May 1 990 to May 1991.

Observations were made during the day as well

as the night, using a torch light with a red filter

(>610 nm). The reproductive condition of every

female was noted at each observation. In pregnant

females, the embryo was detected by palpation

and lactating females were easily identified by

the presence of well-developed mammary glands.

The young were also tagged. Infants with fresh

umbilical cords were noted as being one day old.

The time interval between the first parturition in

the batch until the last parturition denotes a cycle

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of breeding, somewhat confusingly called

breeding “season” in our publication (Isaac et

al. 1994).

Gopalakrishna et al. (1975) reported that

Pipistrellus mimus was a “continuous breeder”.

Suthaakar Isaac, a Ph.D. student of G. Marimuthu,

studied the breeding patterns of this vespertilionid

as a part of his research programme. It soon

became apparent that parturition in this bat occurs

in four distinct and discrete cycles in a year as

shown in Fig. 23. The first cycle of breeding lasted

from July 5-14, 1990 (n=6); the second cycle lasted

from September 14 to October 30, 1990 (n=8); the

third cycle lasted from December 21, 1990 to

February 23, 1991 (n=8); the fourth cycle lasted

from March 23 to April 1 1 , 1 99 1 (n=3) (Isaac et al.

1994). P. mimus , unlike other microchiropteran

bats of southern India, gave birth to twins. Females

gave first birth to twins at the minimal age of 1 03

days. Possibly facing high mortality rates, this

bat species would have to increase the rate of

reproduction. Dead infants were often seen on

the floor below the daytime roosts of this bat.

Since the mother bat does not carry her young,

the rate of predation at the roost may also be

higher than that in other bats. It is also a

behavioural oddity that P. mimus bats are the

earliest to fly about at dusk, and the last to return

to the roost at dawn, thus exposing themselves

twice in 24 hours to visual detection by predators

(Isaac and Marimuthu 1993a) such as the black-

shouldered kite El anus caeruleus. Their flight

activity pattern is crepuscular. We have, however,

never witnessed any birds of prey capture a bat.

We have confirmed that the same female bats

underwent successive pregnancies and

parturitions. Our results imply that P. mimus is

the most prolific breeder among the

microchiroptera. It would be interesting if similar

studies as ours (Isaac et al. 1994) are undertaken

for other organisms that are often claimed to be

“breeding continuously in the tropics.”

Fig. 23: The four breeding seasons of Pipistrellus mimus.

The upper panel represents number of infants from newborn to pre-volant stages, found in the tunnel.

The striped areas in the lower panel indicate durations of parturition (After Isaac et al. 1994)

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General Remarks

Personally for me, there remain many

unsolved puzzles in the bats 1 studied. Inside the

cave of Jain Hills I noticed, at around 1400 hrs to

1600 hrs, how still the colony of Hipposideros

speoris became. No bats flew around, even the

restless circular movements of the heads of the

bats had substantially abated, and it looked as

though the bats were indeed in deep sleep. At

such times if my movements happened to cause

the slightest displacement of a smooth pebble

under my feet, the ensuing muffled noise made a

hundred bats jerk up in alarm. This ‘alertness in

sleep’ is perhaps an adaptation against predators

like the smoothly gliding snakes. Dilip Joshi often

brought to the laboratory the sloughed skin of

big snakes, which he found inside the cave. The

cave was also visited by bandicoots, and we had

lost at least one experimental male H. speoris to

these nightly predators. There were also bam owls

and owlets at the cave mouth. I had on one

occasion, during our field ethology work on

foraging, noticed what might have been a barn

owl, capturing a bat. Interestingly, the same

H. speoris bats, ultra-sensitive to small noises

within the cave, flew later in the night around

2000 hrs in the central bus stop of Madurai or in

the noisy railway station. The hissing of the

engines and the throngs of passengers did not

appear to bother the hunting hipposiderid bats.

This suggests that there is a heightened threshold

sensitivity to audible sound when bats are on the

wing. This may also explain why the ‘best hearing

frequencies’ of insectivorous bats have such a

restricted band, often excluding sounds audible

to humans. Similarly these shy creatures, which

avoid the brightness of the sky on full moon

nights and fly under canopy cover, forage on

swarms of insects drawn to bright sodium-vapour

streetlights.

All the species of insect bats that we

examined showed roost fidelity, with the exception

of Tadarida aegyptiaca (body weight 22 to 24 g).

This bat is a strong flier that preferably forages at

considerable heights of 15 to 20 m above the

ground, well over the canopy and ponds. A colony

of c. 2,000 bats lived in crevices and cracks of a

vertical rock on the northern slope of the Jain

Hills. They were very noisy, squeaking in chorus,

a little before the onset of the evening foraging

flight and were difficult to observe. We had

planned to monitor activity/rest patterns of this

bat also, but were surprised, when we arrived at

the site of the colony one evening not a single

bat was to be seen. No other bat colony in the

Madurai region had vacated its roosting site as

T. aegyptiaca did. A few weeks later we were

informed that some bats had returned. In view of

the inconvenience involved in studying the

behaviour of this bat roosting in the narrow

crevices of the rock at a height of 50 to 70 m, we

decided not to continue with our observations

on T. aegyptiaca.

I have left out much of our labora-

tory experimental findings on light-induced

PRCs made for the circadian rhythms of

Taphozous melanopogon, and T. kachhensis and

H. speoris to avoid unnecessary technical details.

We had reported that the circadian rhythm in

activity-rest cycles free-runs in dim light of 5 lux

and responds to dark breaks of 2 hours and 4

hours, with advance and delay phase shifts as a

function of phase experiencing the ‘blackout’.

Similarly, phase shifts are also caused by light

pulses of 1 5 minutes and 1 000 lux given at different

phases. We reported for the first time for any model

system (Subbaraj and Chandrashekaran 1978),

that the time course and waveform of phase

response curves obtained from experiments using

pulsed light and pulsed darkness are mirror images

of each other. The idea of reviewing our studies

was to impart a flavour of the kind of work my

students and I had been doing on insectivorous

bats, among other objects, at the MKU for two

decades. Many of the observations reported here

were also first reports of their kind, when they

were made. The ecology of roosting sites, site

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fidelity and social interactions of some of these

bats were the most fascinating facets for me.

Acknowledgements

The kind of research described here could

not have been carried out without the enthusiastic

participation of venturesome students like

R. Subbaraj, G. Marimuthu, K. Sripathi, Dilip Joshi

and K. Usman. I thank Dhanashree Paranjpe for

helping me with the figures. I thank Professor

Hubert Markl for encouragement and the gift of a

jeep, when he was Vice-President of the Deutsche

Forschungsgemeinschaft, which enabled more

comfortable travel and transport of some

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( 1 994): Fecundity in the Indian pigmy bat Pipistrellus

mimus. J. Zool. (London) 234: 665-668.

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dimmer than starlight entrains the circadian rhythm

of a bat. Naturwissenschaften 69: 1 92.

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phase response curve obtained for the circadian

rhythm of a bat inside a cave. Indian J. Exp. Biol.

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bat. J. Exp. Zool. 233: 187-192.

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dwelling millipede Glyphiulus cavernicolus.

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TAIL LENGTH IN ENIGMATIC NORTHEAST INDIAN MACAQUES

AND PROBABLE RELATIVES

( With four text-figures)

Jack Fooden1

✓

Key words: Arunachal Pradesh, Macaca assamensis , Macaca thibetana , range

extension, subspecific contact zone, tail length variation

Recent sightings in western Arunachal Pradesh, Northeast India, of macaques with tail proportions

similar to those in Macaca assamensis assamensis apparently extend the known range of this

subspecies c. 400 km westward along the Lesser Himalaya, across the great bend of the Brahmaputra

river. The contact zone between M.a. assamensis and M.a. pelops apparently is at c. 2,000 m along

the slopes of the Lesser Himalaya west of the Brahmaputra river, in Arunachal Pradesh and

possibly also in adjacent Bhutan.

Introduction

Macaca assamensis and M. thibetana are

closely related species that inhabit contiguous

areas in southern and eastern Asia (Fig. 1 ; Fooden

1 988, p. 1 9); despite their close relationship, these

two species differ strikingly in tail length.

Recently, Choudhury (1998, p. 7; 2000, p. 14)

conducted field work in a previously unsurveyed

area in Northeast India and observed monkeys

that obviously belonged to the M. assamensis -

M. thibetana group, but in which tail length

differed from what previously would have been

expected in that area. The purpose of the present

paper is to consider Choudhury's new information

in the general context of tail length variation in M.

assamensis and M. thibetana (see Appendix).

Geographic Distribution and Tail Length

Variation in M. assamensis and M. thibetana

Macaca assamensis includes two

recognised subspecies, M.a. pelops and M.a.

assamensis, which differ in tail length (Fig. 1 ). The

known geographic distribution of M.a. pelops is

narrowly restricted to the Outer and Lesser

Himalaya from central Nepal (83° 36’ E) eastward

'Division of Mammals, Field Museum of Natural

History, 1400 South Lake Shore Drive, Chicago,

Illinois 60605-2496, USA.

Email: [email protected]

to northeastern Assam (94° 25' E) (Fooden 1 982,

p. 4; Choudhury 1997, p. 37). The known

geographic distribution of M.a. assamensis is

much more extensive, encompassing lower and

middle mountain ranges in continental Southeast

Asia, from c. 30° N in Xizang (earlier Tibet), China,

to c. 1 5° N in Thailand and Laos (Fooden 1 982, p.

5). Prior to Choudhury’s (1998, p. 7; 2000, p. 14)

recent research, the gap between the easternmost

Himalayan record of M.a. pelops (94° 25' E) and

the westernmost known Himalayan record of

M.a. assamensis (95° 45' E) was c. 175 km wide.

Macaca thibetana , in which no subspecies are

recognised, is broadly distributed in eastern China,

from c. 25° to 33° N and from c. 102° to 120° E

(Fooden 1988, p. 2).

The following summary of tail length

variation in M.a. pelops , M.a. assamensis , and

M. thibetana focuses on the ratio of tail length to

hind foot length (T/HF) in adult males, which is

the only relevant information available concerning

the newly observed monkeys in Northeast India;

however, tail length variation in adult females and

immatures in M. assamensis and M. thibetana is

generally similar to that in adult males (Fooden

1988, pp. 9, 10). T/HF in adult males clearly declines

progressively in M.a. pelops , M.a. assamensis ,

and M. thibetana (Fig. 2). In M.a. pelops adult

males, T/HF values (mean ±SD, extremes) are 1 .86

±0.109, 1.73-2.07, n = 8; in M. a. assamensis adult

males, these values are 1.26 ±0.127, 1.11-1.56,

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Fig.l: External characters in adult males and geographic distribution of Macaca assamensis pelops, Mcicaca

assamensis assamensis and Macaca thibetana\ slightly modified from Fooden (1988, p. 4; cf. Fooden et al.,

1994, p.623)

n = 21; and in M. thibetana adult males,

these values are 0.4 1 ±0.077, 0.33-0.54, n = 7. Within

each of these three taxa, no consistent pattern of

geographic variation in T/HF is evident (Fig. 3).

Tail Length in Newly Observed Northeast

Indian Macaques

Choudhury (1998, p. 7; 2000, pp. 6, 14)

observed macaques with unexpected tail

proportions in November 1997 at Piri La (27° 07' N,

92° 26' E), western Arunachal Pradesh, Northeast

India. At this locality, three troops of monkeys

were encountered in temperate broadleaf forest

at 2,000-2,700 m elevation; troop size, determined

for two troops, was > 1 5 and 21. In one of these

troops, Choudhury was able to sketch the

dominant adult male and to sketch and

photograph one of two subordinate adult males

(Fig. 4). Although circumstances did not permit

external measurements to be taken, Choudhury’s

sketches provide a basis for provisional estimation

of T/HF in these two adult males; tail length is

also clear in Choudhury’s ( 1 998, inside front cover)

colour photograph of the subordinate adult male,

but, unfortunately, the monkey’s hind feet are

obscured by foliage in this photograph, which

therefore precludes its use for calculation of a

second estimate of T/HF in this monkey. From

the available (albeit suboptimal) evidence of

Choudhury’s sketches, T/HF in the dominant male

is roughly estimated to be 1 .2, and T/HF in the

subordinate male is roughly estimated to be 0.8

(see Appendix).

Sightings of macaques said to have tail

proportions similar to those of the Piri La

286

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Tail length/Hind foot length

TAIL LENGTH IN ENIGMATIC NORTHEAST INDIAN MACAQUES

2.1

2.0

1.9

1.8

1.7

1.6

1.5

1.4

1.3

1.2

1.1

1.0

0.9

0.8

0.7

0.6

0.5

0.4

0.3

Fig. 2: Tail length/hind foot length ratios in adult male Piri La macaques

compared with ratios in adult male Macaca assamensis pelops, Macaca assamensis assamensis

and Macaca thibetana ; for documentation see Appendix

M. a. Piri La M. a. M.

pelops macaques assamensis thibetana

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TAIL LENGTH IN ENIGMATIC NORTHEAST INDIAN MACAQUES

288

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95° 100° 105° 110° 115°

Fig. 3: Geographic variation in tail length/hind foot length ratio in adult male Piri La macaques,

Macaca assamensis pel ops, Macaca assamensis assamensis and Macaca t hi be tana', for documentation see Appendix

TAIL LENGTH IN ENIGMATIC NORTHEAST INDIAN MACAQUES

Fig. 4: Field sketches of Piri La macaques: (a) dominant adult male, (b) subordinate adult male;

reproduced from Choudhury (2000, p. 13), by permission

macaques also have been reported by local

inhabitants at three localities 20-60 km west and

north of Piri La (Choudhury 2000, pp. 8, 15); no

quantitative estimates of T/HF are available from

these reports. All of these sightings of Piri La-like

macaques apparently are at upper elevations,

2,000-3, 100 m (Choudhury 2000, pp. 15,20). At

lower elevations, 1 00-2,000 m, longer-tailed M. a.

pelops (T/HF « 1 .85; Fig. 2) is reportedly common

in this part of western Arunachal Pradesh

(Choudhury 2000, pp. 15,20).

Choudhury (2000, pp. 4, 6, 19) further

suggests that the Piri La macaques may differ from

M. assamensis in slightly darker dorsal pelage

coloration, slightly more prominent side whiskers,

and in an unspecified slightly different

vocalisation. Information provided concerning

these subtle differences between Piri La macaques

and M. assamensis is insufficient to permit

evaluation of the postulated differences at this

time; within M. assamensis , variation in dorsal

pelage coloration and prominence of side whiskers

has been reported previously (Fooden 1982, p. 6).

Discussion

Judging from available information, T/HF in

the Piri La adult males clearly differs from T/HF in

M.a. pelops and M. thibetana adult males (Fig. 2)

(cf. Choudhury 1998, p. 7; Groves 2001, p. 235).

T/HF in the Piri La dominant adult male (c. 1 .2) is

near the mean value for T/HF in M.a. assamensis

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TAIL LENGTH IN ENIGMATIC NORTHEAST INDIAN MACAQUES

( 1 .26); T/HF in the Piri La subordinate adult male

( c . 0.8) is less than the minimum value known in

M. a. assamensis (1.11). For the present, it seems

reasonable to allocate the Piri La macaques to

M.a. assamensis , as tentatively suggested by

Choudhury (2000, p. 20). In this case, the low

T/HF value of the Piri La subordinate male may be

interpreted alternatively as an individual

abnormality, as an expansion of the range of

previously known normal variation of this

character in M.a. assamensis , or as an artifact of

the method of estimation. The uncertainty

concerning T/HF in the subordinate male can be

resolved, as indicated by Choudhury (2000,

p. 20), by future quantitative study of T/HF at Piri

La and other upper elevation localities from where

Piri La-like macaques have been reported.

Assuming that the Piri La macaques

are correctly allocated to the subspecies

M.a. assamensis , the known geographic range of

this subspecies is thereby extended c. 400 km

westward along the Lesser Himalaya, west of the

great bend of the Brahmaputra river (Fig. 3); a

Refer

Choudhury, A. (1997): Checklist of the Mammals of

Assam. Gibbon Books and Assam Science

Technology & Environment Council, Guwahati,

India. 103 pp.

Choudhury, A. (1998): Pere David’s macaque discovered

in India. Rhino Found. Nat. NE India, Newsl. 2(3):

inside front cover (photo), 7.

Choudhury, A. (2000): Survey of Pere David's macaque

in western Arunachal Pradesh: Final report. WWF-

lndia. North East Regional Office, Guwahati, India.

23 pp.

Dao Van Tien ( 1 978): Sur une collection de mammiferes

du plateau de Moc chau (Province de So'n-la, Nord-

Vietnam). Mitt. Zool. Mus. Berlin 54: 377-391.

previous designation of the Brahmaputra river

as the intersubspecific boundary between the

ranges of M. a. pelops and M. a. assamensis was

an extrapolation based on locality records then

available (Fooden 1982, pp. 32, 34). Choudhury’s

(2000, p. 20) new information indicates that the

contact zone between M. a. pelops ( 1 00-2,000 m)

and M.a. assamensis (2,000-3,100 m) is located at

c. 2,000 m along the slopes of the Lesser Himalaya,

west of the Brahmaputra river, in Arunachal Pradesh

and possibly also in adjacent Bhutan.

Acknowledgements

I thank the curators and staff of the

museums listed in the Appendix, for permission

to study specimens in their custody; I also

thank A. Choudhury for generous permission to

reproduce his sketches in Fig. 4. Valuable

comments on a previous version of this

manuscript were provided by L. Austin (Chicago

State University) and L.R. Heaney (Field Museum

ofNatural History).

ENCES

Fooden, J. ( 1 982): Taxonomy and evolution of the sinica

group of macaques: 3. Species and subspecies accounts

of Macaca assamensis. Fieldiana: Zoology (n.s.)

10: vii + 52 pp.

Fooden, J. (1988): Taxonomy and evolution of the sinica

group of macaques: 6. Interspecific comparisons

and synthesis. Fieldiana: Zoology (n.s.) 45: vi +

44 pp.

Fooden, J., Quan Guoqiang, Zhang Yongzu, Wu

Mingchuan & Liang Monyuan (1994): Southward

extension of the range of Macaca thibetana. Int.

J. Primal 15: 623-627.

Groves, C.P. (2001): Primate taxonomy. Smithsonian

Institution Press, Washington, DC. viii + 350 pp.

290

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TAIL LENGTH IN ENIGMATIC NORTHEAST INDIAN MACAQUES

Appendix

Documentation of collectors' flesh measurements of tail length and hind foot length

in Macaca assamensis and Macaca thibetana adult males (cf. Figs. 2 and 3)

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Appendix ( contd)

Documentation of collectors' flesh measurements of tail length and hind foot length

in Macaca assamensis and Macaca thibetana adult males (cf. Figs. 2 and 3)

1Key to abbreviations: AMNH = American Museum of Natural History, New York. BM(NH) = British Museum

(Natural History), London. BNHS = Bombay Natural History Society, Mumbai. FMNH = Field Museum of Natural

History, Chicago. IZCAS = Institute of Zoology, Chinese Academy of Sciences, Beijing. KIZ = Kunming Institute of

Zoology, Kunming. MCZ = Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts.

SCIEA = South China Institute of Endangered Animals, Guangzhou.

2Data from Choudhury (2000, pp. 13, 15); in Choudhury’s drawings (p. 13), tail length and hind foot length of the

dominant male are 17 mm and 14 mm, respectively, and tail length and hind foot length of the subordinate male are

11 mm and 13.5 mm, respectively.

3Data from Dao (1978, pp. 378, 382).

*Measurements taken on skeleton, recorded in collector’s field notebook.

292

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STATUS, ECOLOGY AND CONSERVATION OF THE INDIAN WOLF

CAMS LUPUS PALLIPES SYKES

( With two text-figures and one plate)

Yadvendradev Jhala1

Key words: Food habits, human - wolf conflict, threats, distribution

Two wolf subspecies were believed to inhabit the Indian subcontinent, the Tibetan wolf ( Canis

lupus chanco) whose range extends from the trans-Himalaya into Tibet and China, and the Indian

wolf ( Canis lupus pallipes) that ranges over much of peninsular India. Recent genetic data shows

three extant wolf lineages in the Indian subcontinent. Two of these, the Himalayan and peninsular

lineages, are ancient and unique to the Subcontinent, while wolves from Kashmir belong to the

widespread wolf-dog clade. The Indian wolf (peninsular clade) inhabits semi-arid agro-pastoral

landscapes and scrub forests. It is considered endangered, with its numbers ranging between 2,000-

3,000 individuals. Undisturbed habitat patches of 5-1 5 sq. km that offer good cover and water are

critical for successful breeding in established packs. Wolf territories range between 150 and

300 sq. km and are a function of prey and denning habitat availability. Indian wolves whelp in

December-January in an underground den excavated by the alpha pair. The mean litter size is 4.8

pups. Juvenile wolves can disperse from their natal pack after the age of 8 months. The majority of

wolf populations in India survive outside protected areas, and subsist primarily on livestock. In

some areas, wolves have been reported to attack children. The common factors in such areas are

high human population, poverty, too few or heavily guarded livestock, and poor wild prey availability.

Human-wolf conflicts are of serious magnitude over much of the wolf’s range in India. Wolves are

persecuted by smoking pups in their dens, sometimes by shooting and recently by poisoning.

Major threats to the continued survival of wolves in India are persecution by poisoning, and loss of

denning habitat to intensive agriculture, development, and industry.

Introduction

Canis lupus once had the largest natural

range of any land mammal, besides Homo sapiens

(Sheldon 1992), and had successfully colonised

much of the Northern hemisphere (Mech 1970).

Out of the 32 odd subspecies of wolves that are

currently recognised (Mech 1974), two are

believed to occur in the Indian subcontinent.

Canis lupus chanco or the Tibetan wolf is found

in the trans-Himalayan region and its range

extends into Tibet, China, Manchuria and

Mongolia. The Indian wolf C.l. pallipes ranges

over much of peninsular India and the same

subspecies is believed to occur in Iran and Israel

(Mendelssohn 1982, Shahi 1982). C.l pallipes is

'Wildlife Institute of India, RO. Box No. 18,

Chandrabani, Dehradun, Uttaranchal 248 001, India.

Email: [email protected]

much smaller in comparison to other subspecies

of wolves, except C.l. arabs that is found in the

Arabian peninsula.

Molecular genetic data from wolf and dog

populations from around the world suggests that

they belong to a closely related wolf - dog clade

(Vila et al. 1 997). Genetic analysis of mitochondrial

DNA (control region and cytochrome b) of wolves

from the Indian subcontinent has shown that there

are three extant lineages, two of them very

different from the wolf - dog clade. Wolves from

peninsular India considered to be C.l. pallipes

may have diverged from the wolf - dog clade about

500.000 years ago and are different from the

pallipes found in the Middle East. Wolves from

Himachal Pradesh to eastern Nepal (considered

to be C.l. chanco) are basal to the other wolf

clades and may have separated from them about

800.000 years ago, while wolves west of Kashmir

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ECOLOGY AND CONSERVATION OF THE INDIAN WOLF

belong to the widespread wolf - dog clade. Thus,

the peninsular and Himalayan wolf lineages of

India are very ancient and unique to the Indian

subcontinent (Sharma et al., in press).

Almost nothing is known about the ecology

and status of the Himalayan wolf lineage. In these

regions, persecution of wolves is common by

pastoralists (Fox and Chundawat 1992), and it is

likely that the Himalayan lineage would probably

be one of the most endangered canids in the

world. The account that follows is of the

comparatively well-studied peninsular lineage of

the Indian wolf (Plate 1 , Fig. 1 ).

Adult male and female Indian wolves weigh

between 19-25 kg and 17-22 kg respectively.

Adult wolves measure between 103 and 145 cm

from nose tip to tail tip, and between 57 and

72 cm at shoulder height. Coat colour is sandy

brown with black hair tips. Some wolves have a

more rufous tinge to their coats. Adult wolves

can be distinguished from juveniles in the field

by their size, behaviour, and white markings that

develop above the eyes, on the chin and under

the throat. Colour variation is rare among Indian

wolves, black wolves being reported only from

Ladakh. In much of the wolfs range in India,

summer temperatures up to 47 °C are not

uncommon, while winter temperatures

occasionally approach 0 °C. In summer, most of

the fur is shed, only sparse long hair remains on

the wolfs body. This gives the wolf a scrawny,

long-legged appearance.

Though the wolf is believed to have evolved

as a temperate species, the Indian wolf is aptly

adapted for living in semi-arid and hot

environments. The small body size reduces food

demands, permitting it to sustain its populations

on smaller ungulates, lagomorphs and rodents;

the shedding of under fur and behavioural

thermoregulation permits this canid to live in hot

and arid regions. The Indian wolf still needs ample

drinking water, and in that sense is not truly

adapted to desert living like the chinkara ( Gazella

bennettii).

Status, Distribution and Habitat Needs

C.l. pallipes is considered endangered in

India and features on Schedule 1 of the Indian

Wildlife (Protection) Act of 1972 and of CITES

(Shahi 1982). Killing or trading of C.l. pallipesis

prohibited by law in India. Shahi (1982) in his

preliminary survey estimated the wolf population

in India to be about 800 individuals that were

patchily distributed over peninsular India (Fig. 1 ).

More intensive surveys in the state of Gujarat

and Rajasthan have shown that wolf distribution

is continuous in these states. The population of

wolves in these two states was estimated to be

between 450 and 620 (Jhala and Giles 1991).

Analysis of recent surveys, coupled with the

dispersal capability of wolves (Mech 1 995) makes

it likely that the wolf population within the states

of Gujarat, Rajasthan, Haryana, Uttar Pradesh,

Madhya Pradesh, Maharashtra, Karnataka and

Andhra Pradesh is continuous (Fig. 1).

The current estimated distribution of wolves

covers a much larger area (Fig. 1 ) than was reported

earlier. This does not imply range extension by the

wolf, but is a result of more intensive surveys and a

better understanding of wolf distribution. The wolf

is still far from safe in most of its range and occurs

at low densities (about 1 wolf per 100-200 sq. km,

Jhala and Giles 1991). High density wolf populations

(up to 5 wolves per 100 sq. km) are found to occur in

some habitat pockets and preserves. Breeding packs

ranging from 4 to 14 wolves have been reported

from the Bhal, Dwarka, Banni, and Abdasa area in

Gujarat and Kutch; Kumbhalgarh, Gudda-Bishnoi,

and Pali-Barmer and Jodhpur areas in Rajasthan;

Nannaj, Rehukuri, Nasik, and Phaltan areas in

Maharashtra; Neoradehi in Madhya Pradesh;

Rollapadu in Andhra Pradesh; Melkote and

Ranibennur in Karnataka; and Mahuadaur,

Hazaribag and Palamau areas in Bihar (Fig. 1 ). Such

high-density habitats are extremely important for

wolf conservation, since these pockets serve as

successful breeding and recruitment areas from

where wolves disperse to occupy marginal habitats.

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Fig. 1 : Distribution of the wolf Cams lupus pallipes in India. Locations of known breeding populations

of wolves, areas where wolf studies have been conducted and areas where wolf attacks

on children have been reported are shown

In the light of current information and in

concurrence with Ginsburg and MacDonald ( 1 990),

a population estimate of about 2,000-3,000 wolves

for the Indian peninsula seems more realistic

(Jhala 2000) and may well be a conservative

underestimate.

Since wolves are believed to have evolved

in boreal forest systems as predators of large

ungulates, it is rather surprising that the Indian

wolf rarely lives in forests and prefers scrubland,

grassland and semi-arid pastoral/agricultural

landscapes. The Indian wolf probably evolved

during the drier spells of the Pleistocene to exploit

a relatively unoccupied niche as a top carnivore

of the arid zones.

The eastern population of C.l. pallipes ,

found in Orissa, Bihar and parts of West Bengal,

is an exception and occurs in moister forested

habitats (Shahi 1 982), but even here wolves are

not reported where thick forests occur. Wolves

occur on the periphery of protected forest areas

as is seen in Kumbhalgarh and Kailadevi

Sanctuaries in Rajasthan; Panna, Kanha, and

Bandhavgarh National Parks in Madhya Pradesh;

and Gir in Gujarat. These peripheral areas are under

heavy biotic pressure from surrounding human

populations and are reduced to scrub forests. It

is these scrublands that wolves primarily use.

Wolves do not seem to be restricted in their

movements at night and even approach very close

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to human dwellings and villages (Jhala 1991). In

Maharashtra, radio-collared wolves were

observed foraging in the Ojhar township of Nasik,

at night. However, during the day, wolves are

extremely selective in their habitat use, limiting

themselves to habitat pockets that offer visual

cover and shade, away from human disturbance.

Such habitat pockets are crucial for denning and

as rendezvous sites (areas where pups are kept

after they leave the natal den), and could be in the

form of broken terrain, rocky outcrops, rivulets,

and grass or scrub patches. In Velavadar National

Park and the rest of the Bhal area, wolves preferred

to use moderately dense Prosopis chilensis

patches (Jhala 1991; Jethva and Jhala, in prep.),

while in Nanaj area wolves used forestry

plantations (Kumar and Rahmani 1 995); in Banner

and Pali districts of Rajasthan ‘ Aorans’ (sacred

groves) were crucial wolf habitats. In Ojhar-Nasik

and near Jodhpur, airforce and military bases

provide refugia for denning and rendezvous sites

amongst a semi-urban landscape. In the Bhal area,

where wolves were studied through telemetry for

over eight years (Jhala 2001), core areas of wolf

territories where dens and rendezvous sites were

located were between 5-10 sq. km and were

characterised by low human disturbance, good

cover and presence of fresh drinking water (Jethva

2002; Jethva and Jhala, in prep.).

Wolves living in the western dry zone of

India were considered to be nomadic, primarily

following pastoralists during their annual

migrations. However, telemetry studies in the Bhal

and Kutch area of Gujarat, and in Ojhar in Nasik

district of Maharashtra, have shown that Indian

wolves are territorial like all the other subspecies

of wolves. It seems likely that under certain

ecological conditions territorial boundaries are

more fluid. Territories of Indian wolves can be

quite large, with profound seasonal use of only

some areas within them, giving an appearance of

“nomadism” to a casual observer. The area

covered by a wolf that predated on children, in

eastern Uttar Pradesh in 1996, was over

1 ,000 sq. km (Jhala and Sharma 1 997). Food, water,

and availability of habitat for denning and

rendezvous sites are factors determining territory

size (Fuller 1989, Jhala 1991). Wolves subsisting

on wild prey in areas of high prey densities were

observed to have small territories (100 sq. km),

while wolves subsisting primarily by scavenging

and by predation on domestic livestock had larger

home ranges (250-300 sq. km) covering the grazing

grounds of several villages. Territoriality acts as

a spacing mechanism to limit the number of packs

in an area. Prey density and prey size regulate

pack size and territory size. These two factors act

in synchrony to socially regulate wolf density in

an area.

Social Organisation

In canids, sociality increases with body size

(Moehlman 1990, 1992). Wolves being the largest

of the canids, have a highly developed social

system. Wolf society is organised around the

breeding or alpha pair, the alpha pair and its

offspring living together as a pack. Thus, a pack

is normally a family unit that may have been

established by related or unrelated wolves. The

pack stakes out and defends a resource territory

from other wolf packs. Territorial defence is done

by scent marking, howling and by actual strife

between neighbouring packs (Mech 1970). As

juvenile wolves mature, they either tend to

disperse from their natal pack or stay back as

helpers to their parents. Indian wolf pups may

disperse as early as 7-8 months of age (Fig. 2).

Dispersers wander in search of mates and available

habitat to establish their own territories and packs,

while helpers bide their time till they become

breeders themselves by replacing and/or

displacing their parent (Packard and Mech 1 980).

Telemetry data has shown packs to be quite

tolerant to intruders. Single non-pack members

were observed in close proximity to pack members

and even shared kills on some occasions. Two

packs were observed to intermingle without any

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Jhala, Yadvendradev: Canis lupus pallipes

Plate 1

Fig. 1: The Indian wolf represents an ancient lineage of Canis lupus that is unique to peninsular India and

parts of Pakistan. Considered to be endangered, its numbers are believed to be between 2,000 to 3,000.

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12

N

GO

JC

o

CO

0.

Dispersal

Mating

_J 1 — I 1-

Whelping Rendezvous sites

Natal Den

Join the Pack

4 -

Aug. Sep. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May. Jun. Jul.

Months

Fig. 2: Major annual events and the average monthly pack size (with standard deviation)

of three wolf packs in the Bhal region of Gujarat

strife on two occasions and at one time shared

the same core area for a rendezvous site. This

may be an adaptation to a different set of

ecological conditions, like living off clumped food

resources (MacDonald 1979, Pisapio and

Theberge 2000) in the form of domestic livestock

carcasses, or having limited access to undisturbed

habitats in a sea of human modified landscapes.

Food Habits, Predation and Consumption

Wolves are the top predators of

blackbuck {Antilope cervicapra) and chinkara

(Gazella bennettii), both medium-sized to small

ungulates, in much of the arid and semi-arid areas

of India (Jhala 1991, 1993, Sharma 1978). Food

habits of wolves have been studied from seven

different regions in India (Table 1). The majority

of the wolf population in India occurs outside

wildlife reserves, while wild ungulates are mostly

limited to the reserves. Most of the studies on

food habits reported here (Table 1) have been

done in protected areas and, therefore,

overestimate the contribution of wild prey to the

wolfs diet. It would be safe to conclude that the

majority of wolves in India subsist on small size

livestock, primarily goats and sheep (Shahi 1982,

Jhala and Giles 1991).

Reduction in body size reduces food

demands and permits Indian wolves to subsist

even on small prey like hare {Lepus nigricollis )

and rodents. Besides these, wolves also eat

locusts, other insects, reptiles, birds, and

vegetable matter, like the pods of Prosopis

chilensis and fruits of Zizyphns spp. (Sharma 1 978;

Jhala 1 993; Jethva and Jhala, in press). In Kutch,

wolves were also recorded to prey on donkeys

and camel calves. Wolves of Velavadar National

Park subsisted almost exclusively on wild prey,

even though domestic livestock were present in

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Table 1 : Food habits of wolves reported as percent occurrence of prey in scats or percent

of kills detected from various studies in India

ECOLOGY AND CONSERVATION OF THE INDIAN WOLF

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i = sample size

Protected areas

[

Average computed with the exception of Rollapadu due to its low sample size

ECOLOGY AND CONSERVATION OF THE INDIAN WOLF

the area (Jhala 1993). Consumption by wolves

preying on blackbuck was estimated at 4.62

(SE 0.11) kg/wolf/kill. Wolves killed blackbuck at

an average interval of 3. 5 (SE 0.5) days (Jhala 1993).

Daily consumption by wolves estimated by three

independent studies was 1.33 (Jhala 1993), 1.008

(Kumar 2000), and 1 .80 kg/wolf (Jethva and Jhala,

in press). Jethva (2002) continuously monitored

three radio-collared wolf packs in the Bhal for

periods ranging from 148 to 342 hours (total 1994

hours) and quantified feeding and intake rates of

these packs. Wolf predation was biased towards

adult male blackbuck which contributed about 70%

to the total biomass consumption of these wolf

packs (Jethva and Jhala, in press). Predation on

domestic livestock (cattle calves) was limited to

8%, while scavenging off cattle carcasses

contributed 14% to the biomass consumed (Jethva

and Jhala, in press). Wolf predation was a major

limiting factor responsible for shaping the age

and sex structure of the blackbuck population in

the Bhal (Jhala et al. 200 1 ).

Larger packs (6 to 12 wolves) were recorded

in areas where wild ungulate prey was abundant,

e.g. in Velavadar National Park, Gudda Bishnoi,

Nannaj, Dwarka and Rollapadu. In areas where

domestic livestock formed the major component

of the diet, pack size ranged from 1 to 4 individuals

(Jhala and Giles 1991). Large numbers are more

likely to be detected by vigilant pastoralists and

their dogs, and may prove to be a disadvantage

while hunting domestic prey. Moreover, most

large domestic livestock kills are underutilised due

to human disturbance to feeding wolves and a

major portion of the kill is lost to scavengers like

dogs and vultures. Utilisation of kills in wilderness

areas by wolves is high due to lack of human

disturbance.

Wolves are capable of hunting prey much

larger than themselves. This is primarily achieved

by hunting in packs. Wolves when hunting in

packs use different “strategies” like stalking and

rushing or chasing. Most chases do not last more

than 0.5 km and several of the chases are

unsuccessful. Long chases of over 1 km were

observed when wolves hunted blackbuck calves.

Large prey is usually eviscerated and dies due to

shock and blood loss. Wolves sometimes chase

prey towards other pack members waiting in

ambush. The technique of hunting blackbuck and

chinkara by a lone wolf is by stalking, followed

by a quick rush. If the prey is caught, it is normally

killed by a bite on the throat or nape. The prey

usually dies due to asphyxia. The majority of kills

were made at night, however, during winter and

monsoon, when the days were not very hot,

wolves were seen hunting during daylight hours.

Wolf predation is targeted towards ungulates that

are old, very young or in poor condition.

Breeding

C.l. pallipes is the only subspecies of

wolf that breeds in winter. Births throughout the

wolf’s range seem to be well synchronised. Births

in Gujarat (n=22 litters), Uttar Pradesh (n= 1 ), and

Maharashtra (n=4) occurred between December

15 and January 15. Mating occured sometime

during October to November (Fig. 2), between the

dominant male and female of the pack. The mating

pair is locked in a copulatory tie that lasts for 20-

30 minutes (Sheldon 1992). The gestation lasts

for 62-63 days (Mech 1 970). The breeding (alpha)

female begins excavating dens about a month to

15 days prior to whelping. It takes 2-10 days to

dig a complete den, and one or more dens are

simultaneously excavated. The alpha male

sometimes assists in the excavation but the female

does most of the digging. Dens are excavated in

dry river embankments. Sometimes, fox and

porcupine holes are enlarged. Hollow trunks of

Salvadora oleoides , stacks of harvested cotton

stems, and rock crevices are also used (Jhala 1991,

Kumar 2000, Sharma 1978). There seems to be site

fidelity for denning. Four study packs in the Bhal

region and two packs in Kutch excavated dens in

the same area (1 km radius) for 4-7 consecutive

years, even after the alpha females of three packs

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were replaced. A wolf den in the Gudda Bishnoi

area near Jodhpur excavated in the hollow trunk

of a Salvador a tree, showed evidence of use over

several years. Wolf dens that we observed had

oblong entrances with an average diameter of

48 cm (n=8, SE 14.5). The tunnels were about 3 m

long with a single chamber at the end measuring

about 1 . 1 m by 0.7 m, having an average height of

0.25 m and was at a depth of 0.6 m from the surface.

The mother confines herself to the vicinity

of the den 5-7 days prior to birth. In wild wolves

studied in the Bhal, Kutch, andNasik, mean litter

size was 4.8 (n=28 litters) and ranged between 4

to 6 pups. Pups are bom blind, develop blurred

vision by the age of 1 5 days and by 1 8-20 days,

occasionally emerge from the den to play and

nurse. At the age of about 30-40 days, the pups

are normally moved to another den. They are

moved between 4-5 rendezvous sites between the

age of 40 days to their joining the pack at the age

of 5-6 months.

Canid milk is quite dilute (Oftedal 1 984). The

lactating female needs to drink a lot of water so as

to produce milk. Denning sites and rendezvous

sites are so selected that fresh water is always

available within a radius of 2 km. Water availability,

remoteness from human disturbance, visual cover

and shade, seem to be the critical parameters for

selecting rendezvous and denning sites. At these

rendezvous sites, pups may continue to use dens

that are excavated by adult wolves and by the

pups themselves up to the age of 5-6 months (till

June/July). This behaviour has not been reported

in other wolf subspecies and may be an adaptation

in the Indian wolf to escape high ambient summer

temperatures. After the pups left the natal den, they

were rarely observed using a single den at

rendezvous sites but were distributed in two or more

dens. This behaviour was prominent in Kutch, where

persecution of pups by herdsmen was severe.

Rarely were all the pups killed when shepherds

smoked wolf dens other than the natal den.

Nine rendezvous sites were used by the

Velavadar pack over 2 breeding years. During the

summer (March to early June), the rendezvous

sites in and around a stream bed were intensively

used. The pups would spend the hot hours of the

day in cool depressions dug out in the stream

bank. The depressions were 25-30 cm deep and

25-50 cm wide. These dug out depressions were

also used by adult wolves for lying up. The pups

were restricted to an area of 250 m radius at their

first rendezvous site. Wolf tracks converged from

all directions along well used trails. The area smelt

strongly of wolf odour and urine. Pup and adult

wolf scats accumulated in and around the area.

Kill remains like bones and hides were often strewn

around. The description of rendezvous sites of

timber wolves in North America (Joslin 1 967, Mech

1 970) matches extremely well with the rendezvous

sites of Indian wolves. Wolf pups were taken to

feed on blackbuck kills as far as 3 km from the

rendezvous site, by the age of 2.5 months in

Velavadar National Park.

Conflict with Humans

Since the majority of the wolf populations

in India live outside wildlife reserves, in human

dominated landscapes, they subsist primarily on

livestock. A major occupation of the people in

much of the wolfs range consists of livestock

rearing. Large herds of cattle, sheep, and goats

graze the semi-arid landscape. Most of these

livestock are malnourished and die of disease and

starvation. Since humans rarely consume cattle

in India, much of these carcasses are available for

scavenging by dogs, vultures, jackals, hyenas,

and wolves. Besides scavenging, the wolf also

predates on livestock like goats, sheep, and cattle

calves. Wolf predation severely affects the

economy of the pastoral communities that barely

manage to eke out a living from the highly

overgrazed and degraded landscape of semi-arid

India. The pastoral community invests

significantly in measures to protect their stock

from wolf predation. These measures include

night vigils, maintaining guard dogs, building

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thorn corrals, and bringing the stock back to the

village each night.

In areas with good wolf populations,

average territory size would be about 1 50 sq. km.

Considering an average stable pack size of four

wolves, their density in such areas would be 2.7

wolves per 100 sq. km. Further considering that in

these areas wolves would depend on small

livestock (goats and sheep) to obtain about 50%

of their food requirements (Table 1 ), a pack would

predate on about 65 small livestock per year, i.e. a

loss of about 43 goats/sheep per 100 sq. km.

Considering an average price of Rs. 700 per small

livestock, wolf predation would result in a loss of

Rs. 30,000 per 1 00 sq. km. This would be a rough

estimate of the cost of conserving wolf

populations outside of protected areas. In several

wolf areas in India, the State pays monetary

compensation for livestock loss to wolf predation,

as in Maharashtra. This practice helps negate wolf

persecution by pastoralists to some extent.

However, wolf predation is extremely difficult to

authenticate, and paying of compensation has its

difficulties in identifying false claims.

A more severe form of conflict occurs when

wolves attack humans. These attacks are of two

types: a) attacks by rabid wolves and b) predatory

attacks on children by non-rabid wolves (Linnell

et al. 2002). Attacks by rabid wolves are common

in Maharashtra, Karnataka, Kutch, Rajasthan and

Andhra Pradesh, and occur sporadically

throughout the wolfs range in India. Rabid wolf

attacks are rarely lethal by themselves, but could

prove fatal if treatment is not provided in time.

Attacks on children by wolves have been reported

since British times (Blanford 1891, Lister 1917,

Lydekker 1 897, Pocock 1939). In recent times, such

incidents have been reported from Hazaribagh in

Bihar (Shahi 1982, Rajpurohit 1999), Anantpur in

Andhra Pradesh, and Pavagadh in Karnataka.

Jhala and Sharma (1997) confirmed attacks on

children by a wolf in Jaunpur, Pratapgarh and

Sultanpur districts in eastern Uttar Pradesh.

Between March and October 1996, 76 children

between the ages of 2 and 9 years were attacked.

Of these, over 50 attacks proved fatal. A generality

that emerges from Shahi (1982) and Jhala and

Sharma (1997) is that in wolf range areas where

there is high human density (>600 per sq. km)

with poor economic status, poor child care, with

little or no wild prey around, and with low or

effectively guarded livestock populations, wolves

could potentially attack children. Radio-telemetry

data from three different areas in western India

suggests that wolves come into contact with

humans very often. It would be extremely easy

for wolves to attack children in these areas.

However, there are no authentic reports of wolf

attacks on humans in these regions, in spite of

their high wolf densities. Our data suggests that

attacks on children are extremely rare, considering

the opportunities for attacks available to wolves,

and should be viewed within their special

ecological and socio-economic context (Jhala

2000). However, if and when such attacks do occur,

the responsible problem wolves should be

controlled immediately (either eliminated or

captured), so as to prevent a public backlash

against the species (Mech 1995).

Threats and Conservation

It is indeed surprising that in spite of

heavy biotic pressures on the semi-arid habitats

by humans and livestock populations, and severe

human - wolf conflicts, the wolf continues to

survive in India. The tolerant religious and cultural

attitudes of people (Boitani 1 992) towards all forms

of life, coupled with a low density of firearms and

the absence of systematic use of poison are

perhaps responsible for the continued survival

of wolves in most parts of the wolfs range in

India. The old value system of reverence towards

life forms, and attitudes that resulted in the

conservation of natural resources is, however, fast

changing in rural India. Goals are becoming

monetary and actions exploitative. “Wastelands”

and communal grazing lands are rapidly being

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developed for agriculture and industry. Dry

farming, which is conducive to the survival of the

wolf, is being rapidly replaced by irrigation and

intensive farming with multiple cropping. Such

intensive agriculture areas no longer support wolf

populations (Jhala and Giles 1991).

Wolves were severely persecuted during

the British rule in India. Bounties offered for

wolves were higher than those offered for

leopards (Richards 1914). This attitude persisted

till the 1970s and wolves were eliminated even

from the current Velavadar National Park area to

reduce predation on blackbuck (Ranjitsinh 1982).

Though wolves are still shot illegally in some parts

of their range, hunting currently does not pose a

serious threat to the survival of the species.

Pastoral ists and farmers smoke and dig out dens

to kill wolf pups (Shahi 1982, Jhala and Giles 1991).

Most wolf populations could sustain these losses

due to their high fecundity rates. Recently there

have been cases of poisoning of entire wolf packs

in Rajasthan and Kutch. This is an alarming trend,

since poison can wipe out wolves and the entire

carnivore guild from an ecosystem, and needs to

be dealt with severely. Poison has been the

primary cause of wolf extermination throughout

the world. Human attitudes and persecution of

wolves is related to the amount of livestock

damage caused by wolves. In Kutch, where

wolves subsist primarily on livestock, attitudes

are more hostile and human-caused mortality of

wolves higher in comparison to the Bhal where

wolves subsist on wild prey. Schemes like

Maharashtra state’s compensation for wolf-killed

livestock help reduce persecution of wolves to

some extent, but create claim-authentication

difficulties.

The major threat to surviving wolf

populations in India is loss of habitat resulting in

depletion of natural prey densities and non-

availability of appropriate denning and

rendezvous sites. Wolves do breed in suboptimal

habitats, but recruitment is negligible due to

human-caused mortality of pups. In such areas,

human - wolf conflict is heightened as wolves are

forced to subsist on domestic livestock. As

human values become more monetary and

societies opt for quick short-term economic gain,

such “wastelands” become prime targets for

development of industry and intensive agriculture.

Intensive agriculture, especially when irrigated

by deep bore wells powered by electricity in the

arid and semi-arid areas, is unsustainable in the

long run. Once ground water reserves are depleted

by pumping out, intensive agriculture fails.

Unfortunately, this process may take decades and

irreplaceable damage would have been done to

endangered fauna and the ecosystem drastically

altered. More permanent damage is likely to occur,

to flora and fauna adapted to arid conditions, by

large irrigation schemes like the Narmada Project,

which will alter the land use patterns and

ecosystems over several landscapes. In the rush

towards short-sighted and quick economic gain,

society and decision makers are not willing to

consider even meagre compromises to address

and incorporate conservation goals into the long-

term planning process.

Recent research has identified yet another

threat: canine distemper and rabies. Distemper had

wiped out the entire litter of pups for 2 consecutive

years (1993-94) in Velavadar National Park.

Distemper is contagious and affects pups and

juvenile canids, causing mortality in most of those

affected. Rabies is of special concern since rabid

wolves are known to attack people throughout

their range (Linnell et al. 2002). A rabies outbreak

in Kutch, in 200 1 -02, killed most members of my

study packs. The wolves, however, showed good

resilience and within a year repopulated these

vacant territories. This was possible due to their

dispersal ability (Mech 1995) and because wolf

distribution is continuous with good source

populations. Feral dogs, along with other wild

canids like jackals, spread these diseases. Wolf-

dog interactions are common in India. Wolves

interact with feral dogs at kills and feeding sites.

Wolves occasionally kill and eat dogs (Jhala 1991).

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Besides distemper and rabies, dogs also transmit

other diseases like parvovirus, hepatitis, and a

multitude of other infections to wolves (Jhala

1 99 1 , Goyal et al. 1986, Mech 1970).

Indian wolves can hybridise with dogs in

captivity. However, none of the wild wolves (n=45)

screened by us had dog mitochondrial

haplotypes, nor did dog samples from wolf

territories have wolf haplotypes. This suggests

that hybridisation events may be extremely rare

in the wild and do not pose a threat to the wild

wolf gene pool (Sharma et al, in press).

A proposed national wolf conservation

strategy included the following points:

1) encouraging public support and education,

2) enforcing legal protection, 3) paying

compensation for wolf-killed livestock,

4) conducting surveys of wolf populations and

research on the dynamics of select populations,

5) protecting breeding habitats, and 6) eradicating

feral dogs from wolf conservation areas (Jhala

and Giles 1991). Fig. 1 shows sites that have

confirmed breeding populations of wolves. These

sites are well dispersed throughout the wolfs

range and many of them are in protected areas.

Some of the areas that have confirmed breeding

packs but no legal status should be offered some

level of protection, especially during the breeding

season (December to March). Further

development in these areas should be

discouraged. Wolf conservation is not

incompatible with other land uses. Such areas

could be promoted for multiple use, including

controlled livestock grazing and dry farming. It is

essential that remote habitat patches should be

left intact in these areas for wolves to use as

denning sites, rendezvous sites and resting areas

(Jhala 1 995). These core areas need not be of any

great size, 5-15 sq. km undisturbed patches of

good habitat (cover and fresh water are critical)

seem to be sufficient for a pack to breed

successfully (Jethva and Jhala, in prep.). Public

attitudes towards the wolf in the rest of the world

are rapidly improving. Wolves have in the recent

past colonised areas from which humans had

previously extirpated them. Such recolonisations

have occurred naturally as in Scandinavia,

Germany and France (Promberger and Schroder

1993), or were aided by humans as in Idaho (Fritts

2000) and Yellowstone National Park (Mech et al.

1 995), and in the case of red wolves ( Canis rufus)

(Kelly et al. 2000) and Mexican wolves

(C./. bailey i) (Brown and Parsons 2000).

Though the wolf has probably survived in

the Indian subcontinent for the past 500,000 years,

its continued existence in the next 100 years is

questionable. The wolf is a survivor, and does

not face many of the small population problems

associated with insularization that spell doom for

large carnivores. With the correct attitudes and

actions, we should be in a position to ensure its

future.

Acknowledgements

This study was funded primarily by a grant

from the U. S. Fish and Wildlife Service (USFWS);

grants were also received from the National

Geographic Society, Center for Field Research-

Earthwatch, National Fish and Wildlife

Foundation, Wolf Society of Great Britain, and

Flora and Fauna International. I acknowledge the

support of David Ferguson, USFWS, and George

Furness, Jr., Conservation Treaty Support

Fund, for coordination and logistics. I am grateful

to CRC, Smithsonian Institution for support.

I thank all the graduate students, volunteers

and field assistants who worked with me, for

their sincere and untiring efforts at data collec-

tion. Special thanks are due to D.K. Sharma for

helping with the map preparation for this

manuscript.

JOURNAL , BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003

305

ECOLOGY AND CONSERVATION OF THE INDIAN WOLF

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POPULATION STRUCTURE, COMPOSITION AND ABUNDANCE

OF ELEPHANTS ELEP HAS MAXIMUS IN MINNERIYA NATIONAL PARK, SRI LANKA

( With six text-figures and one plate)

Charles Santiapillai1’2, S. Wijeyamohan3,

Chaminda Wijesundara1 and Rajnish Vandercone1

Key words: elephant, Elephas maximus , Minneriya National Park, Sri Lanka

The Minneriya National Park was established mainly to enhance the long-term survival of the

Asian elephant ( Elephas maximus) in a predominantly agricultural area in Sri Lanka. The ancient

reservoir, after which the Park is named, is seasonally hoi^e to large numbers of elephants because

of the availability of water and the extensive areas of grassland created by fluctuations in the water

level. In a preliminary study carried out from September 2000 to August 2001, a total of 974

elephants were recorded, of which 797 were classified according to age and sex. A quarter of the

observations referred to solitary males. The most frequently observed grouping comprised 5-10

individuals that represent the family unit. The population structure appears to be equally divided

between the adults and the other categories. The observed mean adult malerfemale sex ratio was

1 :2.9, close to the national average of 1 :3. Large groupings of elephant were observed when the drop

in water level in the reservoir resulted in increase in the area of the grazing grounds. The largest

group observed comprised 70 animals. There were also all-male groups, whose composition varied

from 2-6. Most of the animals were observed emerging from the forest to feed on the grasslands

between 1600-1700 hrs in the evening. During the rainy season, as the water level in the reservoir

increased, flooding the grazing grounds, elephants moved to areas outside the Park, causing conflict

with the farming community. The gradual build up of elephants in the Park leading to the observed

maximum of 3 19 animals, translates into a crude density of 3.6 animals per sq. km, which is among

the highest densities recorded in Asia. Therefore Minneriya National Park represents one of the

important areas for elephant conservation in Sri Lanka. But the survival of the elephant outside the

protected area can only be assured if appropriate measures are adopted to reduce the human-

elephant conflict.

Introduction

Despite its small size, the island of Sri Lanka

(65,610 sq. km) supports several viable

populations of elephants estimated to number

between 3,000 and 4,000 animals (Santiapillai and

Jackson 1 990). To conserve the elephant and other

wildlife, Sri Lanka has set aside about 12.5% of

the land area under protection. Outside the system

of protected areas, a combination of high human

'Department of Zoology, University of Peradeniya, Sri

Lanka.

2Email: [email protected]

•Department of Biology, Faculty of Applied Science,

Vavuniya Campus of the University of Jaffna, Vavuniya,

Sri Lanka.

population growth and deteriorating fertility of

the land has led to increased encroachment and

degradation of forests inhabited by the elephant

and other wildlife. This has led to a build up of

elephant numbers within the protected areas, while

outside elephants are finding it increasingly

difficult to move about and adjust their densities

to seasonal changes in vegetation and water

availability across their range. One of the more

recent additions to the system of protected areas

is the Minneriya National Park, opened to visitors

on May 29, 1998. It is known to support high

elephant numbers seasonally in response to

changes in the water availability and grazing areas.

However, given its central location in a

predominantly agricultural area, conflict between

308

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POPULATION OF ELEPHANTS IN MINNERIYA NATIONAL PARK, SRI LANKA

man and elephant has become inevitable. No

studies have so far been carried out on the

ecology of the elephant in Minneriya National

Park to document the seasonal build up of its

numbers. It was to address these issues that a

preliminary study was carried out from September

2000 to August 2001.

Study Area

The 8,889 ha Minneriya-Giritale Nature

Reserve was established on February 12, 1988,

and was subsequently upgraded to a National

Park on August 1 2, 1 997. It is situated next to the

Minneriya-Giritale Sanctuary (6,693 ha), a part of

which was designated as a national biosphere

reserve (809 ha). The entire Minneriya-Giritale

complex is situated in an important agricultural

area in the Polonnaruwa district in the North

Central Province of Sri Lanka (Fig. 1 ). The Park is

named after the ancient irrigation reservoir

Minneriya, built by King Mahasena in the 3rd

century AD. It has a capacity of 2,250 ha when

full and a catchment area of 24,000 ha. The main

source of water is from a diversion of the Amban

Ganga, along the Elahara Channel (IUCN 1 990).

Grasslands and a belt of dry mixed evergreen

forest surround the reservoir. The vegetation in

the grasslands bordering the reservoir consists

of species such as Cynodon dactylon , Brachiaria

mutica , Echinochloa colonum , Paspalum

vaginatum and Digitaria longiflora , many of

which are truly hydrophytic. In addition, extensive

patches of grasslands composed mainly of

Imperata cylindrica (Illuk) and Panicum

maximum (Pohon) are also common. The forest is

dense with species such as Drypetes sepiaria ,

Chloroxylon swietenia , Vi I ex altissima ,

Manilkara hexandra , Limonia acidissima ,

Diospyros ovalifolia and Berrya cordifo/ia. The

fauna includes not only the endangered species,

the Asian elephant Elephas maximus and the

leopard Panthera pardus , but also other mammals

such as spotted deer Axis axis , sambar Cervus

unicolor , barking deer Muntiacus muntjak , wild

pig Sus scrofa and jackal Canis aureus. Both sloth

bear Melursus ursinus and wild buffalo Bubalus

bubalis are very rare in the Park, but domestic

cattle range freely in large numbers.

The Park is also renowned for its rich and

diverse waterfowl, both indigenous as well as

migrant. Resident birds include the painted stork

( Mycteria leucocepha/a), Asian openbill-stork

(Anastomus oscilans), white-necked stork

(Ciconia episcopus), and the lesser adjutant-stork

( Leptoptilos javanicus). The reservoir itself

supports a variety of economically important

fishes such as Labeo dussumieri , Puntius sarana,

P. dorsalis , P. chola, Cyprinus carpio,

Mastacembe/us armatus , Ophicephalus striatus ,

etc. (IUCN 1 990). The annual rainfall for the year

2000 was 1 ,344 mm, and the principal rainy season

extended from October to January. During the

dry season, which lasts from February to

September, the Park receives very little rainfall,

and none at all in May, June and July. The average

annual temperature is 27.2 °C.

Methods

All observations on elephants in Minneriya

were made from a vehicle. We adopted the road-

strip count method of Hirst ( 1 969) to monitor the

fluctuations in elephant numbers. The study area

was visited at monthly intervals during which

observations on elephants were made along a 14

km stretch of motorable forest track, through forest

and grassland habitats. Although some animals

were observed up to a maximum distance of 300

m, most of the observations fell within 200 m. As

the elephants in Minneriya have been known to

emerge from the forest predominantly during the

evenings, all sightings of elephants were made

between 1500 and 1900 hrs. At every sighting,

the location of the elephants, their number,

composition and activity were noted. For census

purposes, the four categories recognised by

Eisenberg and Lockhart ( 1 972) — namely adult,

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POPULATION OF ELEPHANTS IN MINNERIYA NATIONAL PARK , SRI LANKA

Fig. 1 : Map of Minneriya National Park, Sri Lanka, showing the reservoir (shaded area),

grazing grounds (open area) and forest (cross-hatched area)

subadult, juvenile and calf — were adopted. A

group of elephants refers to more than two animals

of any age or sex moving together in a coordinated

manner, while a family unit or herd refers to an

integrated unit of closely related elephants of all

classes excluding adult males (Laws 1 970).

Results and Discussion

Population Structure

Between September 2000 and August 200 1 ,

a total of 974 elephants were sighted from 94

observations of which 797 animals were classified

according to age and sex. Nine of the observations

could not be classified. Of the 94 observations on

elephants, 24 (or 25.5%) represented solitary

individuals (adult or subadult males), and 10 (or

10.6%) referred to all-male groups, whose size

ranged from 2-6 animals. A total of 5 1 family units

were identified (Table 1 ), of which 25 (or 49%) had

no adult bulls, while 8 (15.7%) had only one bull,

9 ( 1 7.6%) had two bulls, 4 (7.8%) had three bulls,

3 (5.9%) had four bulls, and 2 (3.9%) had eight

bulls (Fig. 2). The association of bulls with family

units was temporary and was related to the

presence of oestrus females (Short 1966). In an

extraordinary instance, there were 8 bulls

associated with just two cows. Observations by

Douglas-Hamilton (1972), Croze (1974), Laws et

al. (1975) and Poole and Moss (1981) have largely

dismissed the traditional view of a single bull being

an integral part of a family unit to the exclusion of

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POPULATION OF ELEPHANTS IN MINNERIYA NATIONAL PARK , SRI LANKA

Table 1: The composition of the family units of elephants in Minneriya National Park

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POPULATION OF ELEPHANTS IN MINNERIYA NATIONAL PARK, SRI LANKA

Bulls per herd (n= 51)

60i

Number of bulls per herd

Fig. 2: The proportion of adult bulls in family units or herds observed in Minneriya National Park

other adult bulls (Rushby 1965, Sikes 1971, Poche

1974). Nevertheless, as Barnes (1982) points out,

in situations where elephants form very large

groups, it may pay for a bull elephant to exclude

other bulls and remain with a group, given the

opportunities available for mating with different

females that come into oestrus. In such a situation,

there is no need for the bull to move out in search

of cows in heat. But in Minneriya National Park,

given the small size of the groups, there would be

no real advantage for a bull to be permanently

associated with a group. In such a situation, the

best strategy for a bull to enhance reproductive

success would be to move from group to group in

search of cows in oestrus. Monthly changes in

structure and composition of the population of

elephants are given in Table 2. As can be seen

from Fig. 3, the population structure of elephants

in Minneriya appears to be equally divided

between adults and the other categories put

together. Of the total 797 animals that were

classified, 401 (or 50.3%) were adults (males,

including solitary bulls, and females), while 190

(or 23 .8%) were subadults and 206 (or 25.8%) were

young animals (both juveniles and calves). Such

an adult-dominated age structure is characteristic

of several populations of elephants in Sri Lanka

(Eisenberg and Lockhart 1972, McKay 1973,

Nettasinghe 1973, Ishwaran 1981, Santiapillai et

al. 1984, Katugaha et al 1999). An age ratio of

this type is to be expected in a species

Population structure (n = 797)

adults subadults young

Age classes

Fig. 3: The proportion of adults, subadults and young

observed in the population of elephants

in Minneriya National Park

312

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POPULATION OF ELEPHANTS IN MINNERIYA NATIONAL PARK, SRI LANKA

Table 2: Monthly changes in population structure and composition of elephants in Minneriya National Park

characterised by exceptional longevity, long

gestation period and extended inter-calving

intervals.

Sex-ratio

Among Asian elephants, the sex ratio does

not appear to shift significantly from the expected

1:1 ratio until the subadult stage is reached

(Sukumar 1989). The elephant being a

polygynous, sexually dimorphic species, the adult

sex ratio is unlikely to be in parity but varies

considerably from place to place (McKay 1973),

and is usually biased in favour of the females

owing to a higher natural mortality in the males.

As Sukumar (1989) points out, the sex ratio at

stable age distribution will depend on the

magnitude of the difference in mortality rates of

male and female elephants, assuming an equal

ratio at birth. Furthermore, the difference in the

adult sex ratios in an area could be due to either a

high mortality among males or through dispersion

of the animals in space and time (McKay 1973).

Thus, as Katugaha et al. ( 1 999) point out, even

under normal conditions, the sex ratios of

progressive age classes would become

increasingly female-biased in elephant

populations. The observed mean adult

male:female sex ratio in Minneriya, i.e. 1 :2.9, is in

keeping with the national average of 1:3

determined by McKay (1973), Kurt (1974) and

Hendavitharana et al. (1994). However, within the

family units, the ratio becomes slightly more

biased in favour of the females 1 :4.5 (Table 1 ), as

several of them were not associated with bulls.

The adult sex ratio is significantly biased in favour

of the females in the months of September, October

and November, during which the ratio ranged from

1:5 to 1:7.7 (Table 2). However, as Sukumar (1989)

points out, since at any time, a proportion of the

cows would be either pregnant or in lactation

anoestrus, not all the adult females would be

available for mating with the bulls. Hence, the

operational adult sex ratio would not be as skewed

as the one observed in the population. The

observed adult sex ratios seem to indicate that

the elephant population is not subject to very

high mortality as a result of either poaching or

conflict with man. Given the fact that females do

not carry tusks and only a small proportion of the

bulls (less than 7%) are tuskers in Sri Lanka,

poaching for ivory is not the major cause for the

disparate sex ratio. In Minneriya, there were only

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POPULATION OF ELEPHANTS IN MINNERIYA NATIONAL PARK, SRI LANKA

around the Minneriya reservoir. The largest

grouping consisted of 70 individuals. In addition,

there were ten all male groups. These groupings

appear to be temporary associations between

sexually mature elephants held together by weak

social bonds. They exhibit short-term cohesion.

Such all male groups with rapidly shifting

composition of individuals have also been

recorded in Africa (Croze 1974). The maximum

number of bulls seen together in Minneriya was

six (range 2-6).

The composition of one of the groups (No.

43 in Table 1) that were observed in Minneriya

National Park was rather strange. It had eight bulls

and two cows. According to Dr. Phil Kahl (pers.

comm.), who had studied the phenomenon of

musth in African elephants, a possible explanation

for such a strange grouping is that the older of

the females was a cow in oestrus; the younger

female, probably a previous calf of hers that was

staying on with the mother. The 8 bulls were

probably “suitors” hoping to mate with the adult

30

25

20

0

o

c

0

l—

§ 15

o

£

10

5

0

Fig. 4: Group size frequency diagram for elephants in Minneriya National Park

Group sizes (n = 974)

Group size

4 tuskers: 2 juveniles and 2 adults. Both adults

were single tusked, one being right tusked and

the other left tusked, and so could easily be

identified.

Group size

The most frequently observed grouping

comprised 5-10 individuals that represented the

family units or herds (Fig. 4). A number of family

units associate temporarily to form the larger

groupings seen in the Park. Family units vary in

size from two (mother-calf units) to nine (three

adult females with their offspring). The mean

group size including solitary individuals is 1 0.4. If

solitary individuals (i.e. bulls) are not taken into

account, then the average group size increases to

13.6 (range 2-70). Of the 51 groups numbering

more than three individuals, 25 (or 49%) had no

adult bulls. The association of adult bulls with

family units was common in the months from

August to November. Large groups of elephants

were observed following the build up of numbers

314

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POPULATION OF ELEPHANTS IN MINNERIYA NATIONAL PARK, SRI LANKA

Santiapillai, Charles et al. : Elephas maximus

Plate 1

Fig. A: A herd of elephants around the biggest bull in Minneriya National Park, Sri Lanka

Fig. B: An encounter between two bull elephants in Minneriya National Park, Sri Lanka

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003

315

PICS: S. WIJEYAMOHAN

POPULATION OF ELEPHANTS IN MINNERIYA NATIONAL PARK , SRI LANKA

female. One of them might have been a bull in

musth, in which case he would most likely have

been in “consort” with the oestrous female.

The mean group size of elephants varies

over time and space, and some of the variation

could be related to the availability of food

(Leuthold 1976, Barnes 1983, Dublin 1996). In

Minneriya it appears that elephants range in small

groups during the dry season when food is of

poor quality. Olivier (1978) correlates the decline

in mean group size with a decrease in the habitat

quality. It appears that smaller units of elephants

are perhaps better adapted to exploit a patchily

distributed resource than are larger ones (White

et al. 1993). Of the 33 family units having 10 or

more animals that were recorded during the study,

3 1 (or 94%) were encountered during the rainy

season, and only 2 (or 6%) were seen in the dry

season. As Dublin (1996) argues, the formation

of larger groups allows elephants to interact, to

determine dominance hierarchies and to re-

establish bonds. In Minneriya, there is a gradual

build up of elephant numbers from August to

November as animals move in from outside when

many of the smaller water holes dry up in

the neighbourhood. The large Minneriya reservoir

provides a year-round supply of water. Even at

the peak of the dry season, grasslands around

the reservoir become grazing areas for hundreds

of elephants. The surrounding mixed evergreen

forest provides a variety of plants on which the

animals browse. As the rainy season progresses,

the water level in the reservoir increases

substantially, leading to the flooding of the very

grasslands that support large numbers of

elephants in the dry season. As the water level

increases, the elephants disperse to other areas

outside the Park. It is during such times that the

area experiences an increase in the conflict

between man and elephant. It is also the time when

paddy plants have been planted extensively, and

these provide rich feeding areas for the dispersing

elephants. As Laws (1974) argues, the group size

is a measure of the ecological health of an elephant

population, given that aggregations of family units

and bull groups could be the result of stressful

conditions. The stress may arise through

nutritional deficiency or through the loss of

matriarchs following hunting (Eltringham 1977).

The habitat diversity of Minneriya National Park

makes it unlikely that elephants may suffer from

nutritional deficiency. None of the animals that

were observed in the Park were in poor condition.

Besides, as the Park itself is situated within an

agricultural landscape, the elephants, especially

the bulls, have the opportunity to supplement

their diet with highly nutritious food plants

cultivated by man. Therefore, it is social factors

more than nutritional deficiency and loss of

matriarchs from hunting, that may determine group

sizes.

Feeding activity

The distribution of elephants in Minneriya

National Park appears to be aggregated, which

suggests that the area is not uniformly attractive

to them. Much of the elephant feeding activity

was centred round the grasslands. As all the

elephants were observed in the open grasslands

between 1 500 and 1 900 hrs, grazing was the most

important activity recorded. As Katugaha et al.

(1999) point out, groups of elephants usually

spend a large part of their time during the day

within the forests, under shade, and so are not

noticeable until they move to the grasslands in

the late evenings to graze. Although elephants

spend considerable time in the woodlands, trees

and shrubs make up only a small proportion of

their food (Buss 1961). In Sri Lanka, grasses are

the most important food of elephants. But

elephants alter their food habits in relation to

season. Fig. 5 summarises the feeding activity

pattern of the elephants observed in Minneriya.

The peak of the activity was observed between

1600 and 1700 hrs, when the highest number of

elephants, 388 (39.8%) were recorded. The activity

fell during the next hour ( 1 700- 1 800 hrs) and picked

up once again from 1 800-1900 hrs.

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317

POPULATION OF ELEPHANTS IN MINNERIYA NATIONAL PARK, SRI LANKA

Seasonal abundance

As can be seen from Fig. 6, the number of

elephants utilising Minneriya National Park

increased from August, reaching a peak in

October 2000 when a total of 319 animals were

recorded, and declined abruptly after November.

The 8 month period from December to July was

characterised by a great reduction in elephants

or a total lack of them. Such a build up of elephant

numbers in Minneriya is the result of two related

phenomena: seasonality of rainfall, and the

seasonal establishment of extensive grazing

grounds as a result of the release of water for

cultivation. Following the monsoonal rains in

November, the reservoir begins to fill with water,

and as the water level increases, the floodplains

become inundated, making the area unsuitable

for elephants from December to April. This is the

time of lowest abundance in elephants. As water

is released for cultivation in May, the declining

water level in the Park substantially increases

Activity pattern (n = 974)

45

1500-1600 1600-1700 1700-1800 1800-1900

Time (hrs)

Fig. 5: Activity pattern of the elephants (both bull

groups and herds) in Minneriya National Park

i o

the carrying capacity of the Park. Elephants move

in from outside. The gradual build up leads to

Minneriya supporting elephants at a crude

density of 3.6 animals per sq. km. This is among

the highest densities recorded in Asia. In Africa,

Douglas-Hamilton (1973) recorded elephant

density of 5 animals per sq. km in Lake Manyara

Elephant abundance (n = 974)

350

Sep. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May Jun Jui. Aug.

Months

Fig. 6: Changes in elephant abundance in Minneriya National Park from September 2000 to August 2001

318

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, I00(2&3), AUG. -DEC. 2003

POPULATION OF ELEPHANTS IN MINNERIYA NATIONAL PARK , SRI LANKA

National Park, Tanzania. Elsewhere in Sri Lanka,

the values estimated for elephant density range

from 0.17 per sq. km in Gal Oya National Park in

the east (McKay 1973), 0.46 per sq. km in Block I

of Ruhuna National Park in the southeast

(Santiapillai et al. 1984), and 0.12 per sq. km in

Wilpattu National Park in the northwest

(Eisenberg and Lockhart 1972). These density

values highlight the importance of Minneriya as

a conservation area for elephants in Sri Lanka.

As Seidensticker ( 1 984) points out, the water level

relative to the floodplain and the slope of banks

in reservoirs will determine the capacity of a

catchment area to support elephants. The

seasonal adjustment of water levels in Minneriya

reservoir has resulted in the provision of grazing

grounds for elephants, thereby making the

Minneriya National Park one of the crucial

conservation areas in north central Sri Lanka.

Assuming a minimum elephant population

of about 300 during the peak period, and an

average weight of 1,800 kg for each animal, the

elephant biomass in Minneriya amounts to 540

metric tonnes and the average biomass density

is 0. 1 6 tonnes per sq. km. However, as elephants

spend relatively long periods of time in very small

areas, the ecological densities of elephants in

grasslands can be substantially higher. Although

artiodactyls such as wild pig, water buffalo,

spotted deer, barking deer and sambar are present,

their numbers are low, and their populations are

thinly distributed across the Park. Hence the

elephant remains the most important terrestrial

herbivore. Its importance in Minneriya stems from

its enormous size, intemperate appetite and its

high mean age of survival, which as Watson and

Bell (1 969) point out, enable the species to make

relatively massive interventions in terms of a

diversion of energy flow in an ecosystem.

Conclusion

Elephants have been the raison d’etre for

the establishment of Minneriya National Park.

Despite its small size, it is one of the important

conservation areas in Sri Lanka that is able to

sustain large numbers of elephants seasonally. It

is an important area for the survival of a number

of populations of elephants residents outside its

boundaries, as it provides food, cover and water.

The annual arrival and association of elephants

in Minneriya during the rainy season makes the

Park an entrepot where gene exchanges between

unrelated bulls and family units are possible.

Such genetic exchanges would improve the

genetic fitness of the population and help keep

inbreeding depression to a minimum. Furthermore,

as Dublin (1996) argues, elephants may aggregate

periodically in order to maintain and strengthen

bonds or establish dominance hierarchies within

kin groups. The fact that elephants at times were

either low in number or not observed at all

indicates that they, if not migratory, do undertake

at least extensive movements within a large home

range (Wyatt and Eltringham 1 974).

The farming community in the periphery

also exploits the rich grazing grounds: their herds

of domestic buffalo are allowed to graze inside

the Park. There are some 200 domestic cattle

competing with elephants. Fortunately, there were

no signs of overgrazing caused by cattle, as has

been reported in Gal Oya (McKay 1973), to an

extent that elephants must scarify the ground in

order to feed on the short grass. Nevertheless,

the number of domestic cattle needs to be

controlled in order to prevent overgrazing in the

future. Otherwise, heavy grazing will probably

lead to a degradation of this rich grazing area and

replacement of native grass by the less palatable

Imperata cylindrica. Given the proximity of the

Park to agricultural and human settlements,

the long term survival of the elephant in

Minneriya National Park can only be assured if

management authorities ensure not only the

welfare of the ecologically most dominant

herbivore and its habitat, but also that of the

people who compete with it for the limited

resources of the land.

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POPULATION OF ELEPHANTS IN MINNER1YA NATIONAL PARK , SRI LANKA

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321

THE SOUTHERN KIANG EQUUS K1ANG POLYODON

{ With six text-figures and two plates)

G. Neumann- Denzau and H. Denzau1

Key words: Equus kiang polyodon, skull data, morphology, distribution, threats, north Sikkim,

south Tibet

The Tibetan wild ass or kiang can be divided into 3 subspecies: the Eastern kiang ( Equus

kiang holdereri ), the Western kiang ( E.k . kiang) and the Southern kiang ( E.k . polyodon). So far, the

Southern kiang was only briefly known, mainly from sparse and inconsistent material, based on

Hodgson’s 19th century collection in the British Museum. Now additional skulls of polyodon ,

collected in north Sikkim in 1938/39 by the expedition of E. Schaefer, have been discovered in the

Zoological Museum in Berlin and for the first time examined by the authors. The data prove that the

Southern kiang is indeed a separate subspecies with shorter, but relatively broader and more box-

shaped head. Mounted specimens show differences in colouration and about 10-20% lower withers

height than other kiangs. All available sightings of kiangs south of latitude 32° N since 1 774, were

brought together, analysed, locations determined and the extensive data used to draw a sound new

kiang distribution map for Sikkim, Nepal and south Tibet. The distribution boundaries and

populations of the Southern kiang, especially towards the west, need further investigation. Possibly

there are not more than a few hundred Southern kiangs left, making it an endangered subspecies,

which urgently requires more attention. Only better knowledge can help to protect the smallest

kiang and its habitat adequately.

Introduction

The kiang or Tibetan wild ass has been

regarded by some authors as a distinct species

{Equus kiang), while others regard it as a

subspecies of Equus hemionus. Even modern

molecular genetic studies have not led to any

agreement. While Groves and Ryder (2000) stand

for the separation as a species, Schreiber (pers.

comm, in 2002) suggests keeping the question

open till data for its closest relative, the dziggetai

{Equus hemionus hemionus ), are available for

comparison. Eisenmann (1986) is unable to

discriminate between skulls belonging to dziggetai

{E.k hemionus) and kiang, so that she is tempted

to consider the kiang as a subspecies of Equus

hemionus. After examining a larger database ( 1 60

skulls: 35 kiangs, 29 dziggetais, 32 kulans, 37

onagers and 27 khurs), she concludes in Schreiber

et al. (2000) that “the present osteological

database supports a single-species concept for

all other hemiones” (other than hemippes , which

were omitted from discussions). One of the reasons

for regarding the kiangs as subspecies of Equus

hemionus is that the offspring of a male hybrid

(kulan x kiang) and a female kulan proved to be

fertile (Pohle 1983, 1986). This argument, according

to clarification in Groves and Ryder (2000), is not

valid.

We want to review the available data and

add some new skull data as well as geographical

locations regarding the poorly known southern

subspecies of the kiang {E. kiang polyodon

respective E. hemionus polyodon). Since the

discovery and description of the Southern kiang

was accompanied by several historical errors, it is

necessary to explain some details of the relevant

publications chronologically.

History

'Im Brook 8, 24321 Panker, Germany. Early material: After a brief description of

Email: [email protected] a dead specimen along with the first naming for

322

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003

SOUTHERN KJANG

Equus kiang by Moorcroft (1824) and another

more detailed description of the same specimen

by Moorcroft and T rebeck (1841), Hodgson ( 1 842)

postulated that a second wild member of the horse

family existed in Tibet, which he believed at that

time to be neither identical with the kiang of

Moorcroft nor the hemione (of Pallas) but with

the Asinus : “ Asinus equioides, Mihi. Species

wants verification, spoken of by Moorcroft and

others”, in fact, Moorcroft had earlier mentioned

sightings of wild horses, wild asses, as well as

hybrids of both, on his first visit into Tibet in

1812 (Gupta 1987). Some years later Hodgson

( 1 847a), after obtaining some kiang specimens for

examination, described these animals under the

genus Equus , sub-genus Asinus , as “ Asinus

polyodon mihi” with the remarks, “very common

in all parts of Tibet,” and, “there is, I believe, no

species of wild horse in Tibet, and only one species

of wild ass, viz. the kiang above described ... I

think the kiang may prove a new species, and 1

have named it polyodon from its singularly

anomalous dentition, having 7.7 molars in the

upper jaw.” In a drawing of a skull and a row of

upper cheek teeth, he pointed to the location of

the additional tooth (today known as wolf tooth).

Hodgson (1847a) mentioned five kiangs (not

clear if complete specimens or just skins), one

kiang skull and later two fresh kiang specimens

(provided with skulls) at his disposal. He

published dimensions of a male and a female

kiang, and data of a female kiang skull (all in

comparison with a tanghan or Tibetan pony), but

without informing the reader where exactly his

kiang specimens came from (subtitle of his

attached Plate 6: “ Asinus polyodon mihi. The

kiang of East Tibet”), as in those days only one

kind of kiang was believed to exist all over the

high altitude region.

After having received comments regarding

similar dental anomalies among other equids,

Hodgson (1847b) compared the teeth of three

kiang skulls (all young, but no one less than 4

years old) with those of domestic horses and

concluded: “That they have done so (highest

authorities had uniformly given 6/6 for the Equine

formula) is a fact sufficient to excuse and justify

my insisting on the extra tooth commonly found

in the kiang, and not, I believe, commonly found

in the Equus”

Gray ( 1 849) received three kiang specimens

sent by Hodgson to the British Museum:

“unfortunately they were so destroyed by insects

during their passage from India, that it was

impossible to preserve any part of them except

the skull and the bones of the limbs.” He described

three skulls, but referred to them first (1849) as

Equus kiang , and later ( 1 852) as Asinus hemionus

and Asinus kiang. He obviously faced a problem

in placing the kiang among the equids.

In the Catalogue of the British Museum

(Gray 1852, p. 273), the three kiang skulls

presented by Hodgson are inadequately marked

as “two skulls, lower jaw wanting.” A scaled

drawing of a skull (not the same as in Hodgson

1 847a) is shown in Gray’s Fig. 2 in Table 37.

The two special catalogues of Hodgson’s

collection presented to the British Museum (Gray

1 846, 1 863) do not mention any kiang skull, only a

kiang skin (presented in 1858). However, skulls of

two tanghans and another domestic equid (listed

as “mule of Tibet” or “ Equus caballus var.

domestica ”) also from 1 858, are mentioned. Before

the second catalogue of Hodgson’s collection was

published by Gray in 1863, Gerrard (1862) had

already published another catalogue in which he

misidentified the two tanghan skulls and listed

them as kiangs. This error caused confusion

regarding the identity of the Southern kiang, for

over a century, as even Lydekker (1916) repeated

it.

Lydekker (1916) listed, in his catalogue of

the ungulate mammals in the British Museum

(BM), five kiang skulls presented by Hodgson to

the Museum between 1 848 and 1858. Three skulls

from 1848 were numbered as 48.6.1 1.16(976a),

48.6.11.1 7(976b) and 48.6. 1 1 . 1 8(976c); and two

from 1 858, as 58.6.24. 1 19(976h) skull and skin, and

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SOUTHERN KIANG

58.9.24. 150(976g). For the 1848 skulls, Lydekker

gave the locality as “probably Hundes district of

Tibet”, and for the 1 858 animals, “upper Sikhim.”

Another kiang skull and skin in Lydekker’s

catalogue, BM79. 1 1 .2 1 . 1 82(976j), transferred from

the Indian Museum in 1879, are also connected

with Hodgson’s name and with the doubtful

locality “Hundes?”. In 1 89 1 , a skin (and skull, the

latter not mentioned by Lydekker) collected by

Mandelli from Sikkim, were given the number

91.10.7.176. Lydekker described all kiangs as

“ Equus kiang”

One has to take into account (see Hunter

1 896) that Hodgson collected huge amounts of

zoological material from Nepal and Tibet during

his time at Kathmandu ( 1 825- 1 843) and later during

1 845-1858 from Darjeeling (in those days part of

Sikkim), and presented the collections to the

British Museum. L. Mandelli was a tea planter

and ornithologist in Darjeeling.

Introduction of the southern subspecies:

Trumler ( 1 959) was the first to notice a difference

between some of the skulls of Hodgson’s BM

collection and other kiang skulls of western or

eastern race, and named a new subspecies which

he called Equus kiang nepalensis , unfortunately

using 58.6.24. 1 1 9 as holotype and 58.6.24. 1 50 as

paratype. Groves and Mazak (1967) investigated

the skulls of the Hodgson collection from 1858

again, and concluded that Trumler’s holotype and

paratype belong to horses (Tibetan ponies/

tanghan). When we checked the old catalogues it

became evident that Gerrard ( 1 862) and Lydekker

(1916) had listed the tanghan skulls presented by

Hodgson (Gray 1 863) erroneously as kiang skulls,

and Gerrard’s location “Nepal” and Lydekker’s

location “Upper Sikhim” were unproven

speculations. But the exact origin of Hodgson’s

true kiang material still remained unknown. Gerrard

mentioned “Thibet” and Lydekker, “probably

Hundes.” Groves and Mazak ( 1 967) believed that

it came “most likely from the area of Tibet north of

the Sikkhim border”, while the skull 9 1 . 1 0.7. 1 76

(Mandelli coll.) is doubtless from Sikkim. Groves

and Mazak (1967) named the Southern kiang as

Asinus kiang polyodon Hodgson, 1847.

Eisenmann and Shah (1996) also did not

believe that certain equid skulls of Trumler’s study

belonged to kiangs at all. They wrote that “the

skulls labelled Kiang nepalensis trumler in the

British Museum collection (never trust a label!)”

are those of E. caballus.

Southern Kiang Material in Various Museums

London: 4 skulls (Id1, 3 9) and 2 skins of

Hodgson’s collection (dated 1 848, 1 858 and 1 879);

and 1 skin and 1 skull (d*), partly broken, collected

by Mandelli in 1891.

Calcutta (=KoIkata): A kiang skull

forwarded in 1838 by G.T. Lushington to the

Indian Museum in Calcutta (Blyth 1863, origin:

Tibet) could perhaps also belong to the southern

subspecies, as the basilar length of this adult

specimen is only 425 mm (our measurement),

although its exact origin and history is unknown.

Blyth (1863) also mentioned 3 kiang skins

presented to the museum in Calcutta by

Dr. Archibald Campbell, 2 of them (mare and foal)

mounted. Dr. Campbell, the Superintendent of

Darjeeling, joined J.D. Hooker on one of his

journeys (1 848-1 849) to Sikkim and south Tibet. It

can only be speculated that he obtained the kiangs

from the range of the southern subspecies. While

searching for details, we found the following

remark in Campbell’s Diary (Campbell 1 852), dated

October 25, 1 849, Lachoong: “We purchased three

good skins of the kiang of Thibet to-day, a male,

female, and young one, and sent them to Doctor

O’Shaughnessy at Darjeeling for the Asiatic

Society’s Museum. The men who sold them were

Thibetan hunters. People who live by hunting in

Thibet are called ‘Hurpo’; they are very numerous;

they eat the kiang, and all other animals, use the

gun, make their own powder, and are good

marksmen; they cultivate and graze sheep

occasionally; but live mostly by the chase.” The

place Lachoong (=Lachung, 27.7° N, 88.7° E) is

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SOUTHERN KIANG

located in Sikkim, about 30 km off the Southern

kiang’s distribution boundary. From these lines

in Campbell’s Diary we come to know that he did

not hunt kiangs himself, but bought them. This,

perhaps, allows us to draw the conclusion, that

the kiang specimens in Hodgson’s collection,

some of them definitely known to be procured by

Dr. Campbell as well, were also purchased at

markets or from hunters in Sikkim, without

knowing the exact locality.

Leiden: The catalogues of Jentink (1887,

1 892) mention a skull of an adult individual Equus

hemionus from Tibet (coll. M. Hodgson, 1 853), as

well as a mounted adult male Equus asinus kiang

from Ladakh, Tibet (coll. M. Hodgson). The

location Ladakh is obviously wrong. This skull,

No. R1666A, has a greatest length of 501.5 mm

(our measurement).

Frankfurt: At the same time as Leiden, the

Senckenberg Museum obtained a kiang specimen

of Hodgson’s 1853 collection, from London

[according to old correspondence between

Horsfield and Temminck, investigated by Smeenk

(pers. comm, in 2002)]. According to Kock (pers.

comm. 2002), there is a skull in the Senckenberg

Museum, without history, which could fit, but it

was not accessible for examination due to

renovations in the building.

Berlin: Ernst Schaefer went on his 1st and

2nd Tibet expeditions (193 1-32 and 1934-36) as a

member of the American Brooke Dolan Tibet

Expeditions (Academy of Natural Science,

Philadelphia). Only the 2nd expedition had

collected Eastern kiangs in east and central Tibet

for museum collections and also for the Zoological

Museum in Berlin (ZMB).

The 3rd Tibet expedition of Ernst Schaefer

went to Sikkim and south Tibet in the years 1938-

39. According to museum documents we studied

in Berlin (Schaefer SHI), a total of 1 5 kiangs were

collected in northern Sikkim and given to the

museum. One kiang was shot on July 28, 1 938 at

Gyakang, the other 14 kiangs were shot to the

east of Lake Gaymtsona So between August 1 2-

24, 1938, and on October 1, 1938. So, the origin of

these specimens is well known. According to the

actual filing cards, besides the 15 skulls, the

museum has 7 skins of Southern kiangs (although

the correspondence in Schaefer SHI mentioned

1 5 skins). For the identity of a mounted group of

3 kiangs see the section on Morphology.

Skull Data

Measurements: Out of the 15 skulls of

Schaefer’s collection, 14 were examined by us in

ZMB, the 15th (unsexed juvenile ZMB 70291) was

totally broken. All skulls were measured, but for

this study we only used the data of the 7 adult

specimens having six fully developed cheek teeth,

i.e. 4 males with numbers ZMB 91104, 91106

(Plate 1 , Fig. 1), 91 107, 83377, and 3 females with

numbers 91 108, 91 1 10, 83379. The juvenile skull

numbers are: male 91 105, 91 1 17, female 83378,

83380,91109,91111,91116.

To allow comparisons with data of other

kiangs, we used here the same measurements as

published by Groves and Mazak ( 1 967) (Table 1 ).

Groves and Mazak (1967) used, besides the 5

Southern kiang skulls of the British Museum, two

skulls (ZMB 91106 and 91110) from Berlin (Groves,

pers. comm, in 2002). This means that two skulls

of Schaefer’s collection were available in ZMB at

that time. The others remained — for reasons

unknown — undetected so far.

Results of comparisons: Single values for

the 3 subspecies of kiangs are only found as

graphics in Groves and Mazak (1967) without

distinction between males and females. To enable

a comparison between our measurements and the

other kiang data, we digitised their figures and

added them to our Figs 1,2 and 3.

A comparison of length measurements of

all kiangs (Fig. 1 , basilar length vs greatest length)

shows that polyodon is significantly smaller than

the other kiang subspecies. While Groves and

Mazak (1967, p. 352) observed a “long basal length

compared to the greatest length” for 3 specimens

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SOUTHERN KIANG

Table 1: Skull measurements

*Numbers refer to numbers on the abscissa in Fig. 4 and Fig. 5.

500.00 — j

Basilar Length

mm

480.00 —

460.00 —I

440.00 —

420.00 —

400.00

460.00

+ 4

Greatest Length mm

— ] T | t | i 1 r |

480.00 500.00 520.00 540.00 560.00

1 =holdereri, 2=kiang, 3 =polyodorr, A-polyodon (Schaefer’s collection)

Fig. 1 : Skull measurements of 3 subspecies of kiangs: basilar length vs greatest length,

based on Fig. 9 in Groves and Mazak (1967), with additional skull data of the Southern kiang

and with linear regression line for all data (n=39)

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Neumann-Denzau, G. and H. Denzau: Southern kiang Equus kiang polyodon

Fig. 1: Skull of Southern kiang (ZMB 91 106) in dorsal, ventral and lateral view

Plate 1

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SOUTHERN KIANG

Neumann-Denzau, G. and H. Denzau: Southern kiang Equus kiang polyodon

Plate 2

Fig. 1: Mounted group of Southern kiangs (Schaefer’s collection)

in the Natural History Museum, Berlin

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of polyodon, the 7 specimens of the Schaefer

collection fit in well with the linear trend of all

subspecies. Therefore, taking the now enlarged

sample size of polyodon into statistical

consideration, their statement cannot be

corroborated.

However, on looking at the relation of skull

breadth to length, it can be noticed that the

measures of polyodon are not the smallest among

the kiangs. The incisor breadth compared to the

palatal length (Fig. 2) is, for example, quite large,

indicating a broad muzzle.

On the other hand, the graph of ‘nasal

breadth vs nasal length’ (Fig. 3) demonstrates

that Western kiangs and Southern kiangs show

no significant difference. The nasal breadth was

taken at the point above the infraorbital foramen.

Table 2 contains the mean values of skull

data of Southern kiangs in Schaefer’s collection,

in addition to skull measurements as published

earlier by Groves and Mazak (1 967). Despite the

uncertainties regarding the origin of some of their

skulls, the polyodon mean values of Groves and

Mazak fit in well with the Schaefer data.

When comparing the differences of the skull

mean values between the kiang subspecies and

the kulan ( Equus hemionus kulan), it becomes

evident that the length measurements (1-5) of

polyodon are almost equal to those of the kulan

(Fig. 4). The breadth measurements 7 and 10 are

Table 2: Mean values of skull measurements of 3 kiang subspecies (in mm ±SE)

* Data of Groves and Mazak 1967, p. 329.

** Schaefer’s collection, ZMB.

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90.00 — |

Incisor Breadth

mm

80.00 —

70.00 —

60.00 —

50.00 —

40.00

i

180.00

• •

• • •«»

• • 4

• •

200.00

Palatal Length

220.00 240.00

• 1,2,3

+ 4

mm

! I

i I

260.00 280.00

1,2,3=unspecified kiang subspecies; 4 =polyodort (Schaefer’s collection)

Fig. 2: Skull measurements of kiangs: incisor breadth vs palatal length,

based on Fig. 7 in Groves and Mazak (1967), with additional skull data of the Southern kiang

about 1 0% larger, the measurements 8 and 1 1 on

the other hand almost 10% smaller. Skull

measurements of the Eastern kiang ( Equus kiang

holdereri) are always the largest.

Compared to the nominate form {Equus

kiang kiang), the skull of the Southern kiang

proves 8-10% smaller in length measurements 1-4

(Fig. 5). Measurements 8, 1 1 and 12 reveal certain

differences among the 3 kiang subspecies. It is

obvious that in the Southern kiang, not only the

incisor breadth (7), but also the diastema breadth

(6), the orbital breadth (9) and the occipital breadth

(10) are proportionally larger than in other kiang

subspecies. Due to these parameters we can

characterise the Southern kiang as more broad-

headed. This fact is the opposite of Trumler’s

erroneous statement, terming the heads of the

Southern kiangs as rather horse-like, “slim and

long.”

Trumler (1959), after examining the skulls of

kiangs, distinguished 3 different subspecies with

the help of the ‘Stimbreitenindex’. His index is

identical with the ‘cephalic index’ of Osborn

(1912): frontal width at posterior borders of orbits,

multiplied by 100, divided by basilar length. He

found a cephalic index of 44-46 for the Eastern

kiang, 46-49 for the Western kiang and only 43 for

the Southern kiang. We determined a cephalic

index of 48.1 ±1.5 for the 7 polyodon skulls of

Schaefer’s collection. The low value of Trumler

for the Southern kiang is due to the erroneous

use of some tanghan skulls.

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80.00 —

Nasal Breadth

mm

70.00 —

60.00 —

50.00

40.00

o

a d

+ o

a a ® •

+ o ^ •

Oa

A

■H-

O

o

1

2

3

4

Nasal Length

mm

160.00 180.00 200.00 220.00

1=/70/c/erer/, 2=kiang, 3~polyodon, 4 -polyodon (Schaefer’s collection)

240.00

Fig. 3: Skull measurements of kiangs: nasal breadth vs nasal length,

based on Fig. 12 in Groves and Mazak (1967), with additional skull data of the Southern kiang

We found the shape of the face in lateral

view slightly more box shaped in the Southern

kiang and more inclined in the other kiangs.

Among the 7 skulls of adults of the Schaefer

collection, 5 have at least one 7th tooth (premolar

PI) in the upper jaw. Groves and Mazak (1967)

examined the presence of PI in different equid

species and they too found a high ratio (8 in 11)

of kiangs with presence of PI.

Morphology

In the ZMB exhibit, there is a mounted kiang

group (male, female and foal) which could not be

identified so far, as none of the museum documents

mention this kiang group (Plate 2, Fig. 1 ). However,

with the help of a newspaper article (A.C.L. 1 940)

we could confirm that a stallion, a mare and a foal of

the 1938-39 expedition were indeed selected for

dermoplastic modelling. J.M. Dolan (1999)

misidentified this group as a member of the

holdereri type from Schaefer’s 2nd expedition.

The two adults within the mounted

Southern kiang group in the ZMB have shoulder

heights of 1 1 3 cm and 1 1 5 cm (our measurements).

The height at shoulder of dead specimens

is given by Hodgson (1847a) as 3 ft 9 in. (1 14.3

cm) for a male, and 3 ft 5 in. ( 1 04. 1 cm) for a female

(printing error?), whereas Bailey (1910) gives 48.5

in. ( 1 23 .2 cm) for a female. The mounted kiang in

Leiden (collected by Hodgson) stands 1 1 7 cm tall

at the shoulder (our measurement).

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SOUTHERN KIANG

Skull Measurement No.

1

2

3

4

1= holdereri , 2=kiang, 2>=polyodon; 4-polyodon (Schaefer’s collection)

Fig. 4: Percent difference in average skull data of 3 subspecies of kiangs in comparison to Equus

hemionus knlan at null axis, based on Table 2 [for measurement numbers see Table 1 ; kulan data

taken from Groves and Mazak (1967)]

Although data taken on dead or mounted

animals may differ slightly compared with data of

live animals, it can be concluded from the available

data that the Southern kiang has an average

shoulder height between 1 1 0- 1 20 cm (thus one of

the smallest living wild equids).

Groves (1974) writes: “These southern

kiangs, south of the upper Brahmaputra, are much

smaller than the big north-eastern ones, only 1 GO-

115 cm high.” He characterised the Western kiang

as 135 cm high and of very dark colour and the

Eastern kiang as 140 cm high and of light colour.

Schaller (1998) wrote: “I have observed the

three supposed subspecies and noted no marked

difference in size or colour. Although slight

regional variation in kiangs may exist, an

acceptance of subspecies seems premature.” He

photographed a group of 7 Southern kiangs in

1995 (erroneously printed: 1985) in the Chigo Co

area, south of Lhasa (Schaller 1998, p. 169).

Besides his photograph, we have seen very few

pictures of the Southern kiang, as in Shah (1994,

Plate 2) with a group of kiangs in the Bamchona

area in northern Sikkim, a picture of a dead female

and a foal in Bailey (1910, Plate A), and a group of

animals in Schaefer (1950, opp. p. 128). A black

and white film with the German title ‘Geheimnis

Tibet’ shows some sequences with herds of

kiangs, filmed by Schaefer’s expedition in north

Sikkim and south Tibet in 1 938-39.

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Mounted animals and skins in different

museums are quite old and faded in colour and

have not been thoroughly examined for this study.

However, taking all impressions together, the

Southern kiang appears to have a shorter brown

elongation from the back down to the shoulder

than the other kiangs, and thus larger white

portions on the chest, a larger white wedge from

the belly upwards to the shoulder and the outer

sides of the front legs white. The white field on

the flank is indistinct.

Geography

Historical sightings and descriptions: Only

a very few kiangs of the southern range have

ever been caught alive. Turner ( 1 800) mentioned

hearing that 4 kiangs were once in Warren

Hastings’ possession. It can be presumed that

these were brought down by Bogle, who travelled

to Teshu Lumbo (near Shigatse/ south Tibet) in

1774, commissioned by Hastings, the first

Governor-General of Bengal, with an order to

collect wild animals and seeds besides other rare

goods.

Bailey ( 1 9 1 0, 1 9 1 1 b) had reared two foals of

the Southern kiang with the help of ponies as

foster-mothers during his time as a trade agent in

Gyantse. The Dalai Lama used to keep tame kiangs

on a ‘Wild Ass Meadow’ between his summer

residence and the state palace of his mother on

the western outskirts of Lhasa (see map in Waddell

‘ \-holdereh , 2=kiang, 3=polyodon ; 4 -polyodon (Schaefer’s collection)

Fig. 5: Percent difference in average skull data of 3 subspecies of kiangs in comparison to the Western kiang

( Equus kiang kiang) at null axis, based on Table 2 [for measurement numbers see Table 1]

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SOUTHERN K1ANG

1904). These semi-tame kiangs, three mares,

allowed Walton (1905) to approach to within

twenty yards (18 m) of them. Waddell (1905)

mentioned that two of the kiang mares from the

captive stock (of unknown origin, maybe even

crossbreeds of different captive subspecies) of

the Dalai Lama were taken as a present to King

Edward VII. One mare was drowned while crossing

the Tsangpo (Brahmaputra), the other landed

safely in England in January 1905 and died,

according to Dolan (1999), in 1915. A skull from

“near Lhasa” (BM 5.6.20.1, listed by Lydekker

1916), possibly that of the drowned animal (same

presenter for live and dead specimen: G.R.

Macdonald), was examined by Groves and Mazak

(1967) and described as follows: “though most

like holder eri, does not fit satisfactorily into the

race because of its (the specimen’s) very long

basal length, proportional to greatest length: a

feature observed in polyodon as well. It may

therefore be suggested that the specimen is a

member of basically holdereri population affected

by some intergradation with polyodon .”

We investigated many old travel reports in

the area south of latitude 32° N and determined

the locations of historical kiang sightings as far

back as possible. Recent references were also

included. The locations, coordinates and

references of kiang observations supposed to

belong to the southern subspecies range are

summarised in Table 3a, the information of other

possible kiang sightings, in Table 3b. We have

included available data from the year 1774

onwards. The coordinates are given, with a few

exceptions, as precisely as kiang observations

could be localised.

Geographical distribution: Our map (Fig. 6)

shows the recorded kiang sightings, according

to Tables 3 a and b, with our interpretation of the

distribution boundaries.

The northern boundary of the Southern

kiang’s geographical distribution is either the

upper Brahmaputra (Tsangpo) or perhaps the

Nyenchen Tanghla Mountain Range to the north

of that river. It can be seen that polyodon might

have its western and northern distribution

boundaries at 88° E and at 29° N (except one

location east of Lhasa). The kiangs west of 88° E

and between 28° N and 29.5° N need to be specially

investigated (our suggestion), for finding out

which subspecies they really belong to. Another

suggestion is that if Southern kiangs inhabit

Bhutan at all, they should be looked for in the

extreme northeast.

If the gap in the kiang distribution south of

the Brahmaputra, between 87°-88° E, proves to be

true, it could be explained by the glaciation history

in the area of the Upper Arun and its tributaries.

Evolution of subspecies is usually an effect of

long-term isolations between populations. The

separation of the Southern kiang from the other

kiangs was possibly caused by glacio-tectonic

forces during the Pleistocene or Late Glacial

period. Kuhle (2001) explains that big ice sheets

must have existed in south Tibet due to the

damming effect of the Himalayan mountain wall,

which found their drainage via the steep south

slopes. The glaciation of the Arun river valley,

between Mt Everest and Kangchenjunga, is

confirmed by glaciogeological findings.

Our map (Fig. 6), which is based on more

recorded kiang sightings than any previous

distribution map, is in agreement with the map of

Schaller (1998, p. 164), regarding a continuous

east-west extension from the western towards the

eastern kiangs. While Schaller also connects the

southern distribution range with that of the other

kiangs, Groves (1974, p. 95) as well as Denzau

and Denzau ( 1 999, p.50 and back cover) keep the

southern subspecies geographically apart. Groves

( 1 974), in his map, left a big gap between the eastern

and western subspecies, but allowed the range of

the Eastern kiang to meet that of the Southern

kiang along the Brahmaputra. It seems that Groves

( 1 974), as well as Groves and Ryder (2000), when

including the Lhasa district in the geographical

range of holdereri , have used the skull BM

5.6.20.1, mentioned earlier, for this claim. Our

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Table 3a: Locations of sightings of southern kiangs

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SOUTHERN K1ANG

careful investigations show that the distribution

boundary of holdereri does not touch Lhasa, but

remains about 1 10 km further north. The sighting

of a kiang east of Lhasa and north of the

Brahmaputra (as reported to Hodgson in 1832)

remains uncertain regarding the subspecies

question and is perhaps a doubtful point in our

Southern kiang distribution map (Fig. 6). However,

we did not want to exclude any information. With

the help of our documentation it will be easier to

re-interpret the subspecies distribution data in

future, if more facts are ascertained.

Present Situation

Official conservation status (for Equus

kiang in general): in China it is listed in first

category of the State Key Protected Wildlife List

(Wang 1998), in India in Schedule 1 of the Wildlife

(Protection) Act, 1972 (Agrawal 1994). The

Southern kiang is listed as ‘DD’ (Data Deficient)

in the '2000 IUCN Red List of Threatened Species’

(Hilton-Taylor 2000), but deserves a threatened

classification as endangered feEN C2a\ since the

total population is estimated to number less than

2,500 mature individuals, severely fragmented,

with no subpopulation estimated to contain more

than 250.

Partial population estimates: Today’s

population density of the Southern kiang is largely

unknown and only a few authors have published

their observations.

Schaller (1 998) mentioned, “During a wildlife

survey in October 1 995 we attempted to delineate

current distribution between the Bhutan border

and the Yarlung Tsangpo. According to local

people, the kiangs were exterminated in most areas

between the 1960s and 1980s. In that eastern part

of their range we found kiangs around Chigo Co

in three populations totalling probably no more

than 200 animals; and others persist just south of

Yamdrok Co.”

Ali (1981) had reported two kiang groups

(consisting of 10 and 7 individuals) during his

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SOUTHERN KIANG

84 86 88 90 92 [°E]

* locations according to Table 3a 9 Z

** locations according to Table 3b

*** additional sightings according to Fig. 10.1 in Schaller (1998) ♦ 4

ssmaa Kiang distribution

Fig. 6: Recent and historical sightings of kiangs in southern Tibet, Nepal and Sikkim

(between 27°-32° N and 84°-93° E) with proposed distribution boundaries for the Southern kiang

ecological studies in north Sikkim in summer 1978

and summer 1979.

Lachungpa (1994) summarised her few

kiang observations during 9 visits (4 without any

kiang sightings) to north Sikkim during a period

of 5 years (September 1989 - September 1994). Her

list shows a total of 52 kiangs in 5 groups (group

size ranging from 4 to 16). She estimated no more

than 10-40 seasonally free-ranging Southern

kiangs in Sikkim.

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SOUTHERN KIANG

Shah (1994), while conducting a survey in

north Sikkim between October 29 and November

15,1 994, counted a total of 74 kiangs in 14 groups

(group size ranging from 1 to 48). Taking additional

sightings by army personnel into account, she

estimated a total population of 74-120 Southern

kiangs in Sikkim and the adjacent areas of China.

Interviewing the local nomads, she came to know

that kiangs are sighted all year round.

Threats: Mahapatra (1998): “In 1962 the

Indian Army was invited over to Sikkim, and has

remained in border areas inaccessible to most

researchers. Landmines have been laid in many

patches in these areas. This has not only

prevented study of rare animals like the kiang,

but also led to decimation of its population.

According to army officials, the three-strand

barbed wire fencing around landmined areas has

been damaged at many places in Dongkung-Chho

Lhamo. But these patches of lush green grass

attract kiangs and locals have often reported

seeing kiangs being blown apart by landmines.”

Also, Raj (1999) describes the minefields

between China and India along the Sikkim border

Refer

A.C.L. (1940): Besuch im Zoologischen Museum in der

Invalidenstrasse. Berliner Jllustrierte Nachrichten,

24.10. 1940, 2. Beiblatt zu Nr. 250: 4.

Agrawal, V.C. (1994): Tibetan Wild Ass or Kiang.

Pp. 161-162. In: The Red Data book on Indian

animals. Part 1: Vertebrata. Zoological Survey of

India, Calcutta.

Ali, S.M. (1981): Ecological reconnaissance in Eastern

Himalaya. Tiger paper 8(2): 1-3.

Anon. (1999): Kiang: The Tibetan wild ass.

(www.south-asia.com/kingmah/greenwat.htm

downloaded 18. Dec. 2001).

Avasthe, R. & A. Jha(1999): Mammals of Sikkim. WWF

Sikkim, Gangtok. 97 pp.

Bailey, F.M. (1910): A young kyang or Tibetan wild ass.

J. Bombay nat. Hist. Soc. 19: 731 +P1. A.

Bailey, F.M. (1911a): Notes on the game animals from

near Gyantse and the Chumbi Valley. J. Bombay

nat. Hist. Soc. 20: 1028-1032.

Bailey, F.M. (191 lb): A quiet day in Tibet. Blackwoods

Magazine 189 (No. 1144): 270-275.

as the main casualty factor for rare wildlife

including the kiang: “villagers have reported

seeing herds of wild animals charging across the

Tibetan border after being fired at by Chinese

soldiers only to meet their end in the Indian

minefields.” Lachungpa (1994) mentioned the

sighting of a lame kiang with the remark “land-

mine casualty.”

Natural calamities include high snowfall in

winter and predation of foals by wolves. Domestic

livestock is a serious grazing competitor (Shah

1994).

Acknowledgements

We want to thank the following persons for

valuable help: Manfred Ade, Hannelore

Landsberg and Irene Thomas (all ZMB Berlin),

Chris Smeenk (Leiden) and Rina Chakraborty

(Kolkata) for making museum collections

accessible, Ashim Mukherjee (Kolkata) and

Wolfgang Frey (Germering) for providing rare

literature, and Colin Groves (Canberra) and Vera

Eisenmann (Paris) for additional information.

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TAXONOMY OF UNGULATES OF THE INDIAN SUBCONTINENT

Colin Groves'

Key words: Artiodactyla, Perissodactyla, taxonomy, Indian subcontinent, Equus khur ,

Moschus cupreus, Muntiacus vaginalis , Cervus wallichii , Cervns hanglu ,

Capricornis thar , Tetracerus quadricornis , Gazella bennettii salinarum

The ungulates of the Indian Subcontinent are remarkably diverse taxonornicaily and, despite their

conspicuous nature, much remains to be learned about them. Their diversity appears to have been

considerably underestimated, at both specific and subspecific level. Here, I argue that in historic

times the Subcontinent has been home to 46 ungulate species. Three of these species ( Equus

hemionus , Rhinoceros sondaicus and Bos javanicus) are probably extinct. Six taxa, commonly-

regarded as subspecies of other species, I here raise to specific rank, or have done so very recently

elsewhere: Equus khur , Moschus cupreus , Muntiacus vaginalis, Cervus wallichii, Cervus hanglu

and Capricornis thar . Twenty-four species are polytypic; of these, eight are more widespread

species represented by a single Subcontinental subspecies, but the remaining 16 show subspecific

diversity within the Subcontinent itself. I describe one new subspecies, Gazella bennettii salinarum,

and recognise three subspecies (for which names already exist) in Tetracerus quadricornis , a

species previously thought to be monotypic.

Introduction

Over the hundred plus years of its

existence, the Bombay Natural History Society

has seen vast changes in its political and social

setting, its clientele and its very' rationale for

existence. It has survived the demise of shikar

and of British India, the dismembering of the polity

that nurtured it, and even the change of name of

the city that forms its base. In the meantime, the

Journal of the BNHS has progressed from being

a vehicle for documenting the fauna and flora of

the Indian Subcontinent to its present status as

the premier voice for knowledge, understanding

and conservation of the Subcontinent’s wildlife,

both nationally and internationally.

In the midst of all these changes, we

should not assume that the Society’s original

mission, that of documenting the fauna and flora

of the Subcontinent, is complete. There is much

to be learned, as I hope to show', about even the

largest species of fauna, the ungulates. Again,

’School of Archaeology and Anthropology,

Australian National University,

Canberra, ACT 0200, Australia.

Email: [email protected]

while the original three countries of the

Subcontinent (India, Nepal and Bhutan) have

become divided into six, with the creation first of

Pakistan and Sri Lanka and later of Bangladesh,

these six countries share a common wildlife

heritage, and 1 treat them together here.

The object of this centennial

contribution, then, is to gather together

information on the taxonomy of the ungulates of

the Subcontinent, and to try to document exactly

what it is that we know or do not know about this

small but important comer of its biodiversity.

Taxonomy

Taxonomy, the science of biological

classification, is siill undergoing a wide-ranging

rethink of its basic premises. A classification is

an '‘information retrieval system”, but exactly what

information is it that we want to retrieve? Most

practising taxonomists today would say that they

want to try and incorporate information about

evolution into their classifications, because this

allows us to make predictions: if certain taxa share

a common ancestor, their behaviours, their

ecology and other aspects of their biology will be

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JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003

TAXONOMY OF INDIAN UNGULATES

modifications of those of this ancestor. About

how we might incorporate evolutionary

information, however, there is as yet no consensus,

although it can be argued that a new one is

emerging.

Genera and higher categories: It is by now

widely accepted that taxa of the genus-group and

above should represent phyletic lineages. To place

species A and B in one genus, and C in another, is

to say that A and B share a common ancestor

which is not shared by C.

The remaining controversy concerns how

we should decide what ranks we should use. At

present, this is purely arbitrary, often based on

no more than tradition. Goodman et al. (1998), in

the context of Primate taxonomy, have proposed

to link taxonomic rank with time since divergence,

and recently Groves (2001), citing some data on

fossil ungulates, suggested slight modifications

to this. A genus, in this scheme, would have a

time depth of about 4-8 million years, a family, of

18-24 million. Subfamilies, tribes and subtribes

would remain categories of convenience, as

would subgenera: they would be used if

subdivisions were needed, regardless of the exact

times.

The Goodman et al. (1998) proposals are

well worth considering; they would bring a much-

needed objectivity into higher-category

taxonomy. For the moment, however, we need more

information on splitting times of lineages; this is

a task for the future, and in what follows I have

not made any drastic proposals for altering

conventional genera.

Species: It is generally not possible to apply

the well-known Biological Species Concept,

whereby species are defined as being

reproductively isolated. If two populations are

sympatric and do not interbreed, this is of course

unassailable evidence that they are different

species; but what if they are allopatric, as closely

related populations usually are? There is in this

case no objective criterion beyond the opinion of

some “expert” of whether they would or would

not be likely to interbreed if their ranges were to

meet.

Many taxonomists now adopt the

Phylogenetic Species Concept, whereby species

are defined as being diagnosably different: that

is to say, that every individual can be

differentiated on at least one heritable feature

(see Groves 2001). This substitutes ‘genetically

isolated’ for ‘reproductively isolated’, and

switches the focus to the pattern (which we can

observe) and away from the process (which we

can only infer, and which may have brought it

about). Under this concept, some species are

recognised that have hitherto been regarded as

only subspecies: a small price to pay, one might

say, for objectivity. In what follows, application

of this fresh view of species has led to a drastic

revision of species boundaries in Equus, Cervus,

Capricornis and Nemorhaedus.

Subspecies: Some taxonomists do not think

it worthwhile recognising subspecies: two

populations are either diagnosably distinct, in

which case they are different species, or they are

not. Yet I think there is still value in distinguishing

two populations which are different as a whole,

but do overlap to some extent. Their gene pools

are distinct, and they are of conservation concern

(for example).

Exactly how distinct should their gene-

pools be to merit subspecific rank? We must not

try to be too rigid about this; the rule that says

that 75% of individuals of one subspecies must

be distinguishable from all individuals of other

subspecies (the 75% rule) is a good guideline.

We can say no more than that.

It is mandatory, of course, that

subspecies be geographic segments of a species.

The category should not be misused to designate

conspicuous morphs (colour, karyotype, etc.) that

occur within populations.

Synonymy: Among junior synonyms, only

those described for the Subcontinent are listed

here. Author, date and type locality, but not full

bibliographic reference, are given.

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Order Proboscidea

Elephants

FAMILY ELEPHANTIDAE

Genus Elephas Linnaeus, 1758

Elephas maximus Linnaeus, 1 758. Asian elephant

Elephas maximus maximus Linnaeus, 1758.

1758 Elephas maximus Linnaeus. “In Zeylonae

paludosis ad amnes” (in the marshes of

Ceylon by rivers). Restricted to rain forests

of Sabaragamuva by Deraniyagala (1939).

1797 Elephas indicus G. Cuvier. Deraniyagala

(1955) states that this was based on a

specimen from Ceylon (Sri Lanka), but an

examination of Cuvier’s description does

not bear this out.

1797 Elephas asiaticus Blumenbach. Asia,

chiefly Ceylon.

1845 Elephas indicus bengalensis de Blainville.

Ganges in Bengal.

1845 Elephas indicus ceylanicus de Blainville.

Ceylon.

1940 Elephas maximus vilaliya Deraniyagala.

Manampitiya, floodplain of Mahavili,

eastern Ceylon.

1950 Elephas maximus dakhunensis

Deraniyagala. Travancore.

Deraniyagala (1955) distinguished two

subspecies in Sri Lanka and two on the mainland

of the Subcontinent. Elephas maximus vilaliya ,

which he calls the giant Ceylon swamp elephant,

is restricted to the Mahavili floodplain, and is

distinguished from the smaller nominotypical

E.m. maximus , which is the widespread form, by

its larger size and associated skull shape

differences. The food supply on the Mahavili

floodplain is exceptional; one must be cautious

about recognising a separate subspecies based

almost entirely on size and size-dependant

features under circumstances where phenotypic

plasticity is so likely to be involved. Sri Lankan

elephants differ from Indian in their general

tusklessness, as male Indian elephants generally

have tusks (see next para). Most Sri Lankan

elephants are rather small, but Mahavili ones can

be larger than mainland ones.

Deraniyagala’s mainland subspecies were

differentiated on the basis of tusk development and

the degree of depigmentation on the ears and face.

In E.m. dakhunensis 96% of the males are tusked,

and depigmentation is “heavy and white”, but in

E.m. bengalensis only 51% of males have tusks,

and depigmentation is “scanty and white”; in

Southeast Asian subspecies over 90% of males are

tusked, and depigmentation is scantier and grayer

(less white) the further south one goes.

The information on the proportion of

tusked males is interesting but cannot be of much

value for differentiating subspecies, given that it

concerns males only and the percentage

differences are so slight (nowhere near the

‘seventy-five percent rule’ described above). The

amount of depigmentation may be more cogent.

All Asian elephants undergo some facial and ear

depigmentation with age, and it is, in my

experience, notably heavier at any age in Indian

than in Southeast Asian elephants, but there is a

great deal of overlap between northern and

southern Subcontinent elephants. For the

moment, no subspecies are recognised in the

region, though Indonesian/Malaysian

{E.m. sumatranus ) and probably mainland

Southeast Asian {E.m. birmanicus ) subspecies

would be worth recognising at least provisionally.

Order Sirenia

Seacows

FAMILY DUGONGIDAE

Genus Dugong Lacepede, 1799

Dugong dugon { Muller, 1776). Dugong

1776 Trichecus dugon Muller. Cape of Good

Hope to Philippines.

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TAXONOMY OF INDIAN UNGULATES

1799 Dugong indicus Lacepede. Indian Ocean.

There appears to be no study to examine

whether any geographic differentiation exists

between Red Sea, Indian Ocean and Australasian

dugongs.

Order Perissodactyia

Odd-toed ungulates

FAMILY EQUIDAE

Genus Equus Linnaeus, 1758

Equus kiang Moorcroft, 1841. Kiang or Tibetan

wild ass

The kiang has a large head and thick muzzle;

relatively long mane; and long hairs not restricted

to tail tuft but extending some way up either side

of tail. The demarcation pattern between

contrasting dark (reddish) body blocks and white

underside is oblique from stifle (knee joint) to

croup (rump), and the white rump patch is infused

with the reddish tone of the haunch. The dorsal

stripe is thin and never bordered with white; it

extends to tail tuft. A dark ring round hoof. Ear

165-178 mm long.

Skull resembles E. hemionus , except that

incisors tend to sit more vertically in the jaws

(except in aged individuals, in which alveolar

recession tends to reveal the oblique roots), and

highest point on cranial profile is often directly

above the posterior rim of the orbit, instead of

behind it.

Equus kiang kiang Moorcroft, 1 84 1

1841 Equus kiang Moorcroft. Eastern parts of

Ladakh.

Size large, skull length 492-537 mm (n=15).

Range in Subcontinent: Ladakh.

Equus kiang polyodon (Hodgson, 1 847)

1847 Asinus polyodon Hodgson. Tibet, just

north of the Sikkim border (fixed by Groves

and Maz&k, 1 967).

1959 Hemionus kiang nepalensis Trumler.

“Nepal”; more probably the region of Tibet

just north of the Sikkim border (see Groves

and Mazak, 1967, who discuss the status of

the skin and skull of the type specimen).

Size very small, skull length 473-494 mm

(n=7), but nasals and toothrow as long as in other

kiangs. Range in Subcontinent: Sikkim.

Equus hemionus Pallas, 1775. Onager or Asian

wild ass

Mane very short, “clipped”; dorsal stripe

thick, bordered with a white line on either side;

white of rump is not infused; demarcation

between the reddish flank block and whitish

underside runs parallel to the body outline, before

turning sharply up towards the dorsal stripe. A

dark ring round hoof. Nasal bones relatively

straight. Skull resembles E. kiang.

Equus hemionus blanfordi (Pocock, 1947)

1947 Microhippus hemionus blanfordi Pocock.

Sham Plains, Quetta, Baluchistan.

This grayish subspecies was unusual in

E. hemionus in that the dorsal stripe did not reach

the tail-tuft. Skull length 484-498 mm (n=2). St is

now extinct; I have seen only two specimens, one

from the type locality, and the other from Kandahar

in Afghanistan.

Equus khur Lesson, 1827. Khur or Indian wild

ass

1827 Equus khur Lesson. Little Rann of Kutch

(fixed by Groves and Mazak, 1 967).

1862 Asinus indicus George, 1869. India.

The khur is sharply distinct from

Equus hemionus , and we cannot now tell whether

the two intergraded; at any rate, the extinct

Baluchistan subspecies of E. hemionus (see

above) was not intermediate, except possibly in

the failure of the dorsal stripe to extend to the tail-

tuft. In Equus khur the coloured blocks on flank

and haunch are very small, so the predominant

colour is white, and lower 45% or more of flank is

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TAXONOMY OF INDIAN UNGULATES

whitish; demarcation on the lower haunch slants

upward from front (stifle) to back. Dorsal stripe

with a clear white border on either side; this

becomes obfuscated with age, but probably never

entirely disappears. White wedge between

haunch- and flank-blocks nearly or fully reaches

spine. Legs pure white. White zone on muzzle

occupies nearly 40% of snout-to-ear distance.

Dorsal stripe fades out halfway down taii. No dark

ring round hoof. Nasal bones raised (making

whole facial profile strongly concave); skull short

[length, male 493-5 1 9 mm (n=3), female 468-5 1 1

mm (n=6)], with small teeth, noticeably high-

crowned. Choanae small. Orbits high.

Metapodials less elongated than E. hemionus. Ear

very long, 1 87-2 1 0 mm.

The khur still occurs in the Little Rann of

Kutch. I have seen a specimen from Thar Parkur,

Sind, where it is now extinct.

FAMILY RHFNOCEROTIDAE

Genus Rhinoceros Linnaeus, 1758

Rhinoceros unicornis Linnaeus, 1758. Indian/

Great one-homed rhinoceros

1758 Rhinoceros unicornis Linnaeus, 1758.

Rookmaaker (1998) shows that this name

was based on the same specimen from

which Albrecht Durer’s famous woodcut

was taken. He suggests restricting the type

locality to Assam.

1817 Rhinoceros indicus G. Cuvier.

1830 Rhinoceros asiaticus Blumenbach.

1867 Rhinoceros stenocephalus Gray. Asia.

1876 Rhinoceros jamrachi Sclater. Manipur.

Contra Rookmaaker ( 1 983), 1 would consider

that the original description, that of Jamrach

(1874), does not truly count as a

“publication.”

The historical distribution of this species

extended from the upper Brahmaputra, on (and

beyond) the Burmese border, along the Ganga-

Brahmaputra system to the Indus and its

tributaries, as far west as the Khyber Pass

(Rookmaaker 1980, Rookmaaker 2000).

In the multivariate analysis of Groves

(1993a), Nepalese and Assamese skulls are

different. Those from Assam tend to have

narrower zygomatic breadth but wider occiput and

interorbital region. A single skull from Koch Bihar

was more similar to those from Nepal. These

findings should be tested on larger samples.

Rhinoceros sondaicus Desmarest, 1822. Javan/

Lesser one-homed rhinoceros

Rhinoceros sondaicus inermis (Lesson, 1 840)

1840 Rhinoceros inermis Lesson. Sunderbans.

Broad across the zygomatic arches, and

high occiput (Groves and Chakraborty 1983,

Groves 1993a). The subspecies from the

Sundarbans and neighbouring districts (including

the Sylhet and Chittagong districts of Bangladesh,

and Manipur, possibly also Bhutan and Sikkim:

see Rookmaaker 1 980) became extinct in the 1 890s.

A skull from Moraghat, Bhutan Duars

(where the species has been extinct since the

1870s), differs slightly from Sundarbans

specimens, especially in its large teeth (Groves

1967).

Genus Dicerorhinus Gloger, 1 84 1

Dicerorhinus sumatrensis (G. Fischer, 1814).

Sumatran/Asian two-homed rhinoceros

Dicerorhinus sumatrensis lasiotis (Sclater, 1 872)

1872 Rhinoceros lasiotis Sclater. Chittagong.

A large subspecies with large teeth and

very broad, high occiput; molar teeth are large

(Groves and Chakraborty 1983, Groves 1993a).

Formerly occurred in the northeastern states of

India and eastern Bangladesh, as far west as the

Sankosh river, and North Cachar, Cachar and

Hailakandi districts in Assam; it apparently still

occurs in Manipur and Nagaland (Choudhury

1997).

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TAXONOMY OF INDIAN UNGULATES

Order Artiodactyla

Even-toed ungulates

FAMILY SUIDAE

Genus Sus Linnaeus, 1758

Sus scrofa Linnaeus, 1758. Eurasian wild pig

(Indian wild boar of Prater, 1971)

Sus scrofa cristatus Wagner, 1839

1839 Sus cristatus Wagner. Near Calcutta (fixed

by Groves, 1981).

1868 Sus aper var. aipomus Gray. Nepal. After a

nomen nudum of Hodgson ( 1 842).

1868 Sus aper var. isonotus Gray. Nepal. After a

nomen nudum of Hodgson ( 1 842).

1843 Sus indicus Gray. Nepal (fixed by Groves,

1981).

1847 Sus affmis Gray. Nilgiri Hills.

1851 Sus zeylonensis Blyth. Ceylon.

1860 Sus bengalensis Blyth. Bengal and Kutak.

Skull large (skull lengths of males in

different populations average from 379 mm in

Nepal to 414 mm in southern India), broad and

high-crowned; mainly black, brindled with white,

with a long mane all along the back. From the sub-

Himalayan tract, from Punjab east to Nagaland,

and south to Bihar, Madhya Pradesh and

Kolhapur, Maharashtra.

Groves (1981) recognised a southern Indian

and Sri Lankan subspecies, Sus scrofa affmis , but

this differs from S.s. cristatus only in its larger

size, and considering the known environmentally

induced plasticity of size in pigs, the inference

that the difference is genetic cannot be sustained.

In addition, the wide variation in size in Sri Lanka

covers both Indian “subspecies”, and more.

Sus scrofa davidi Groves, 1 98 1

1 98 1 Sus scrofa davidi Groves. Sind.

A small-sized subspecies (adult males

average 365 mm), with a low-crowned skull; colour

light brown or yellowish, with long thick mane.

From the dry country of northwestern India (Pune,

Gujarat, Rajasthan) and Pakistan (Sind).

Sus salvanius (Hodgson, 1847). Pygmy hog

1847 Porcula salvania Hodgson. Sikkim Terai.

1863 Sus lilliputensis Gray. Nomen nudum.

A tiny species, shoulder height 230-305 mm

in males, 200-216 mm in females. Dark brown with

subterminal light hair bands, no mane, very short

tail, and relatively long hindlegs compared to

forelegs. The inner “false hoofs” are short

compared to other pigs. A detailed description is

given by Ghosh (1988), who supports restoring

the genus Porcula Hodgson, 1847, for this

species. Known in the past from the grass jungles

of eastern Nepal, Sikkim, northern Bengal, Bhutan

and western Assam; now known for certain only

from Assam.

FAMILY TRAGULIDAE

Genus Moschiola Hodgson, 1 843

Moschiola meminna (Erxleben, 1777). Indian

chevrotain or Mouse-deer

1777 Moschus meminna Erxleben. Ceylon.

1842 Tragulus mimenoides Hodgson. Nepal

Terai.

1843 Meminna indica Gray. Renaming of

meminna.

1843 Meminna malaccensis Gray. Supposedly

from Singapore.

The Indian mouse-deer, which lives in Sri

Lanka, southern India and Nepal, has been poorly

studied taxonomically, in contrast to the related

Southeast Asian genus Tragulus in which dozens

of subspecies have been described.

FAMILY MOSCHIDAE

Genus Moschus Linnaeus, 1758

According to the latest revision (Groves Qt

al. 1995), four species of musk deer occur in the

Subcontinent; three of these (the exception being

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TAXONOMY OF INDIAN UNGULATES

Moschus cupreus) extend into Sichuan and

Yunnan, in China. Some new subspecies remain

to be described in M. leucogaster; these may

possibly prove to be distinct species. All species

live at about 2,500 to 4,200 m above msl, in the

Alpine zone. The undescribed species inhabiting

Uttar Pradesh (areas now in Uttaranchal), Himachal

Pradesh and western Nepal apparently lives in

high-altitude forest at 2,500 to 3,000 m. The

ecological differentiation between the two

overlapping species of the Himalayan slopes,

M leucogaster and M. fuscus , is unclear. The

Subcontinent has no lower-altitude species

restricted to the lower montane forest zone,

analogous to M. berezovskii of China.

Moschus chrysogaster Hodgson, 1839. Alpine/

Golden-bellied musk deer

1839 Moschus chrysogaster Hodgson. “Nepal”:

probably from the Tibetan plateau.

1915 Moschus cacharensis Lydekker,

From eastern Nepal, Sikkim and Bhutan,

entirely in the plateau zone, above the tree-line.

Dark brown, intensely speckled with buff to

golden ochre. Throat with a noticeable pair of

yellowish longitudinal stripes or spot-rows; rump

yellowish, paler than back; ears dark brown,

speckled buff, outside, with pale tips; orange-

ochrey inside and at base. Limbs becoming

paler distally, until shanks are whitish. Underside

paler, often yellowish; an orange eye-ring. Large

size: metacarpal length 109-118 mm, meta-

tarsal 128-138 mm. Hoofs are more elongated

than other species, 27-29 mm (in other species,

24 mm).

Moschus leucogaster Hodgson, 1839. White-

bellied musk deer

1839 Moschus leucogaster Hodgson. “Nepal,”

probably from the Himalayan slopes.

1839 Moschus saturatus Hodgson. “Nepal,”

probably from the Himalayan slopes.

Eastern Nepal, Sikkim and Bhutan, on the

Himalayan slopes. Dark brown to agouti red

above, with a yellowish tone on buttocks; belly

pale grey; throat wholly whitish, or with a white

sternal patch; limbs dark externally, with white

speckling on inner surfaces and often on digits.

Ears blackish, with very fine whitish speckling

except at the tip which is wholly dark. Size large

likeM chrysogaster.

Moschus sp. Kulu/Indian forest musk deer

The musk deer from Uttaranchal (erstwhile

Uttar Pradesh) and Himachal Pradesh (Garhwal,

Tehri, Chamba, Kulu, Kedamath, Barinda Pass,

Jaunsar) and western Nepal (as far east as

Mustang) may be specifically distinct. Grubb

(1982) cites field evidence from M.J.B. Green,

R.M. Mitchell and others that there may be

geographic overlap, with some elevational

difference between light and dark musk deer, within

Nepal, the grey ones being found in dense oak/

rhododendron forests. Colour is pale grey, with

at most a poorly defined browner, speckled saddle

area; ears are bordered with white; the rump is

entirely dark; the belly is white.

Moschus fuscus Li, 1981. Dwarf musk deer

From Assam west to the Mount Everest

region of Nepal, again on the Himalayan slopes.

Colour very dark above and below, the head, ears

and legs being black, neck often lighter; throat

dark often with two incomplete yellow “collars”;

rump with ochrey tones, but black on buttocks.

The smallest species: skull length less than 150

mm, nasals under 46 mm; metacarpal length 88-

101 mm, metatarsal 126-135 mm.

Moschus cupreus Grubb, 1982. Kashmir musk

deer

From Kashmir, at over 3,000 m above msl.

Colour grey-brown, often vaguely spotted, with

a coppery-reddish dorsal saddle; rump dark,

grizzled grey; underside light grey; throat white;

lower segments of limbs whitish. Ears dark

brown, white at base, with frosted rims. Large

species.

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TAXONOMY OF INDIAN UNGULATES

FAMILY CERVIDAE

Genus Muntiacus Rafinesque, 1815

Muntiacus vaginalis (Boddaert, 1785). Red/

Indian muntjac or Barking deer

Though conventionally placed in a single

species, the red muntjac should preferably be split

into at least two. Differences are marked and

consistent, and their different karyotypes should

reduce the fertility of hybrids between them

(although one possible wild hybrid was described

by Groves and Grubb, 1 990). The present species,

found in suitable country throughout South Asia

and mainland Southeast Asia north of the Isthmus

of Kra, differs from Indonesian/Malaysian

M. muntjak in its chromosome number (2n = 6 in

the female, 7 in the male, vs 2n = 8 in the female, 9

in the male of M. muntjak ), and in the relative lack

of contrasts in colour, especially the absence of

the median dorsal darkening of M. muntjak. The

black-legged muntjac of the southern borders of

China and neighbouring regions may constitute

a third species of the red muntjac group.

Descriptions of subspecies are after Groves

and Grubb (1990).

Muntiacus vaginalis vaginalis (Boddaert, 1785)

1785 Cervus vaginalis Boddaert. Bengal.

1827 Cervus moschatus Hamilton Smith. Not of

de Blainville, 1816 (= M. muntjak).

1833 Cervus ratwa Hodgson. Nepal.

1840 Cervus me las Ogilby. Himalayas.

1 845 Cervus styloceros Schinz. Renaming of melas.

Dark reddish dorsally, somewhat paler on

flanks; nape slightly greyer; forehead and occiput

light orange-brown, rest of face grayish; ear-backs

reddish at base, remainder dark grey; limbs dark

brown to grey; underside paler; groin and line on

front of hindlegs, to hocks, white. Largest of the

three subspecies of the Subcontinent, averaging

about 200 mm skull length. Antlers 80-120 mm

long; pedicels relatively long, averaging 125 mm.

From Shevaroy Hills north to Nepal and Bhutan.

Muntiacus vaginalis aureus (Hamilton Smith, 1 826)

1826 Cervus aureus Hamilton Smith. Some part

of southern India.

71844 Cervus albipes Wagner. Supposedly from

Bombay and Poona. This name may actually

be a senior synonym of malabaricus.

1872 Cervus tamulicus Gray. Deccan.

Pale yellowish on body and limbs; nape

greyer; forehead and occiput pale orange-brown,

rest of face light orange; ear-backs orange at base,

becoming grey, and tips and rims may be dark

grey; underside paler; a line on front of hindlegs,

to hocks, white. Medium sized, skuil length

averaging 192 mm. Antlers short, 70-100 mm,

pedicels about 1 10 mm. From Kumaun and Kheri,

southeast to the Deccan.

Muntiacus vaginalis malabaricus Lydekker, 1915

1915 Muntiacus muntjak malabaricus Lydekker.

Nagarhole.

Colour of body and limbs a washed-out

reddish, with much greying on nape and back;

underside drab; white area on inner and lower limb

segments extending round to front of pasterns,

restricting red to a narrow anterior band. Small, skuli

length averaging 178 mm. Antlers short, 60- 1 00 mm,

with short pedicels, only 100 mm. From Sri Lanka

and southern India northwest as far as Pune.

Genus Cervus Linnaeus, 1758

This genus has usually been taken to

include, as subgenera or species-groups, Rusa

and Rucervus (and sometimes also Axis and

Hyelaphus). Groves and Grubb (1987) pointed out

that Elaphurus is the sister-group of Cervus s.s.

(=sensu stricto), so if Elaphurus — a widely-

recognised genus — is to be retained, to preserve

the monophyly of Cervus it is necessary to keep

Axis, at the very least, separate (they included

Hyelaphus in Axis ); and, according to the

arguments of Grubb ( 1 990) and Geist ( 1 998), Rusa

and Rucervus should aiso be recognised as valid

genera for the same reason.

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TAXONOMY OF INDIAN UNGULATES

The genus, thus restricted, has traditionally

been considered to contain two species: Cervus

elaphus (red deer, wapiti, shou, Kashmir stag) and

C. nippon (the sika, of Japan and China). Geist

(1998), however, has considered the

interrelationships of the “elaphine deer” in some

detail, and stressed the consistent differences in

both morphology and behaviour between red deer

and wapiti, which he clearly regards as distinct

species ( Cervus elaphus and C. canadensis

respectively). Information on other elaphine deer

is not adequate to determine their taxonomic status

so clearly, but the two taxa from the Subcontinent

are neither wapiti nor true red deer, though perhaps

closer to wapiti. They belong to what Geist regards

as the “primitive group”, whose antlers are

characterised by the bez tine being larger than

the brow tine and by having a sharp bend at the

3rd tine (like wapiti in both characters), while

retaining a primitive 5-tined plan; extra branching

occurs in well-developed antlers, as is

characteristic of all elaphine deer, but in these

primitive taxa it takes the form of a transverse

branching of the 4th tine, and occasionally of the

5th as well

In these primitive elaphines, as Geist shows,

no special summer coat is grown, so in summer

they are just a faded version of the winter colour.

Their small rump patches and short manes are

also primitive, yet the very short tail, only half as

long as the ear, is an advanced feature.

The rutting call, which is so different in

wapiti and red deer, is different again in the

primitive group. It begins in red deer fashion, with

a closed-lips roar, then opens out as the lips are

retracted to end in a wapiti-like bugle. Geist sees

the group as adapted to saltatorial running on

steep slopes in high mountain forests and among

shrubs.

There is no sense in attempting to keep

these primitive elaphines in the same species as

wapiti, with which they admittedly share a few

primitive retentions; still less as in the same

species as red deer. Their primitive features are

shared with a third taxon, macneilii from Sichuan.

They nonetheless differ consistently from each

other, and the radical step of regarding them as

separate species seems unavoidable. Excellent

colour photographs of them can be seen in Dolan

(1988).

Cervus wallichii G. Cuvier, 1823. Shou

1823 Cervus wallichii G. Cuvier. Nepal;

restricted to Mansarowar Lake, Tibet, by

Lydekker(1915).

1841 Cernus affinis Hodgson. Supposedly from

Sal Forest, Nepal, but more likely from

Sikkim.

1850 Cervus tibetanus Hodgson. Lingmo, Phari,

Dingcham, Tibet.

1851 Cervus nariyanus Hodgson. Western

Tibet.

These are, according to Geist (1 998), short-

legged, broad-hoofed deer with long narrow ears

and a large square muzzle. In winter, light sandy

brown with a grey face, in summer fading to slate-

grey. The large white rump patch extends upward

on the croup and surrounds the short, white tail,

and is sometimes partially divided by a very

weakly-marked median line of the body colour;

there may be a blackish edging infero-iaterally.

Chin and lips are grey or fawn; belly and inner

surfaces of hindlegs grey. The short mane is the

same colour as the body. Skull length of stags

432-48 1 mm (n= 1 0); antler length (straight) of prime

stags 800-1,040 (n^8), brow tine length 272-368

mm; the tip of the beam is longer than the 4th tine.

This species occurred in Sikkim on the

Tibetan frontier (Chumbi Valley), and still occurs

in Bhutan (Dolan 1988). It is uncertain whether

the Tibetan (Upper Tsangpo) population (true

wallichii ) is the same as that of the Chumbi Valley

{affinis), but it seems likely that the features of

the rump patch, the only described difference, are

variable..

Cervus hanglu Wagner, 1844. Kashmir stag

1844 Cervus hanglu Wagner. Kashmir.

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1847 Cervus casperianus Gray. Kashmir.

1859 Cervus cashmeriensis Adams. Kashmir.

In winter, dark liver-brown, the legs and

chest rather darker, and the face, neck and back

usually lighter; in summer, fading to a relatively

light tone with a sharply contrasting dark chest

and limbs. Many females are spotted in summer.

The narrow rump patch does not extend, or hardly

extends beyond the tail root, is invaded by a

wedge of body colour which may extend onto

the tail, and is bordered infero-laterally by a black

band. Chin, lips and inside of ears are white;

belly, groin and inner surfaces of hindlegs

whitish; a dark area on posterior belly. The

metatarsal gland is creamy to light red. There is

dark curly hair between the antler pedicels. Smaller

than Cervus wallichii : skull length of stags

359-415 mm (n=8), but antlers relatively larger,

straight length 752-1 ,053 mm (n=12), with shorter

brow tine (229-315 mm); the tip of the beam is

shorter or barely longer than the 4th tine. From

the Vale of Kashmir and neighbouring regions,

up to 3,600 m above msl; now restricted to

Dachigam and Srinagar City Forest National

Parks.

Genus Rucervus Hodgson, 1838

Regarded as a distinct genus, rather than a

subgenus of Cervus or of Rusa, by Grubb ( 1 990).

The two species of the Subcontinent differ sharply

in antler form, in the development of the lachrymal

pit in the skull, and in the metatarsal tuft of R. eldii.

However, they are clearly sister species, and the

genus Panolia , erected for R. eldii, is not

warranted.

Rucervus eldii (McClelland, 1842). Eld’s/Brow-

antlered deer

Rucervus eldii eldii (McClelland, 1 842)

1842 Cervus eldii McClelland. Manipur.

1843 Cervus frontalis McClelland. Renaming of

eldii.

1843 Panolia acuticornis Gray. Manipur.

1845 Cervus lyratus Schinz. Manipur.

1901 Cervus eldi cornipes Lydekker. Manipur.

A distinctive subspecies, differing from

those in the Indo-Chinese region by its elongated,

spreading hoofs with bare, comified skin on the

backs of the pasterns. It is restricted to the masses

of floating vegetation called ‘phumdi’, in Logtak

Lake, Manipur, where there are still under a

hundred individuals, though it breeds well in

captivity.

Rucervus duvaucelii (G. Cuvier, 1823).

Barasingha or Swamp deer

Rucervus duvaucelii duvaucelii (G. Cuvier, 1823)

1823 Cervus duvaucelii G. Cuvier. Northern India

(restricted to Kumaun by Groves, 1 983).

1834 Cervus bahrainja Hodgson. Nepal.

1835 Cervus elaphoides Hodgson. Substitute

for bahrainja.

1837 Cervus smithii Gray. Northern India

(restricted to Kumaun by Groves, 1983).

1843 Cervus dimorphe Hodgson. Sal forests of

the Morung, Nepal.

1850 Cervus euceros or euryceros Gray. India

(restricted to Kumaun by Groves, 1 983).

1868 Cervus eucladoceros Falconer. West bank

of Ganges, south of Hardwar, United

Provinces.*

A swampy-ground form, with splayed

hooves and naked pasterns. Antlers long, slender,

not compressed or palmated. Short nasals, rostrum

slender (not deep). Little size difference between

sexes; tail long, slender; ears large, rounded, with

thick white hair inside. From Kumaun, Kheri, and

southwestern Nepal; formerly extended east to

Chitwan.

Rucervus duvaucelii branderi Pocock, 1943

1943 Rucervus duvaucelii branderi Pocock.

Mandla, Madhya Pradesh.

*area now in Uttaranchal

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The hard-ground form, with ‘‘well-knit” feet

and hair-covered pasterns. Body size smaller, with

some sexual dimorphism, male with dark rutting

pelage and prominent neck ruff; antlers very long,

with long brow tine, many branches, branching high

up the beam, with long anterior branch. Long nasals,

short broad snout. Tail somewhat shorter; ears large,

rounded, with thick white hair inside. Prominent

white spots along dorsal stripe, especially during

moult. Restricted to Kanha National Park.

Rucervus duvaucelii ranjitsinhi (Groves, 1983)

1983 Cervus duvauceli ranjitsinhi Groves.

Guwahati, Assam.

A swampy-ground subspecies with

spreading hooves and bare pasterns. Long

nasals, short snout, slender from side to side but

deep. Antlers short, thick, branching low down,

with short anterior branch, somewhat compressed

and tending to be palmate. Heavily built, with

females rather small. Tail short; ears small, pointed,

without much white hair inside. White median

dorsal spots not prominent. Assam.

Genus Rusa Hamilton Smith, 1 827

Regarded as a distinct genus, rather than a

subgenus of Cervus , by Grubb (1990).

Rusa unicolor { Kerr, 1792). Sambar (Sambur)

Rusa unicolor unicolor (Kerr, 1792)

1792 Cervus axis unicolor Kerr. Ceylon

(restricted by Hamilton Smith, 1 827).

1792 Cervus axis major Kerr. Ceylon.

1799 Cervus albicornis Bechstein. Substitute

for major.

1816 Cervus niger de Blainville. North India

(restricted by Pocock).

1823 Cervus ar is tot el is G. Cuvier. Nepal.

1823 Cervus leschenaultiG. C uvier. Coromandel.

1827 Cervus hippelaphus Hamilton Smith.

Bengal. Not of Erxleben {Mil) = Cervus

elaphus.

1831 Cervus jar ai Hodgson. Nepal.

1841 Cervus heterocerus Hodgson. Nepal.

1841 Cervus nepalensis Hodgson. Nepal.

1843 Axis pennantii Gray. India.

Antlers are relatively long, with short

brow tines. The two terminal antler tines tend to

form an equal fork, although this is variable;

the relationship between their lengths is also

variable, but the posteromedial tine is frequently

the longer. From Sri Lanka and most of the

mainland, except the far Northeast. Pocock

(1943a) separated Sri Lankan and mainland sambar

into different subspecies, an action described by

Groves and Grubb (1987) as “based on scant

evidence of a size difference” and, even were it

fairly consistent, it would not be very convincing

evidence for different subspecies, given the

known environmental effects on body size among

deer.

Rusa unicolor equina (G. Cuvier, 1823)

1823 Cervus equinus G. Cuvier. Sumatra.

Antlers are shorter, and brow tines are

longer. The anterolateral tine always continues

the line of the beam and is longer than the

posteromedial tine. From the northeast of the

Subcontinent; Pocock (1943 a) mentions the Garo

Hills.

Genus Axis Hamilton Smith, 1 827

Axis axis (Erxleben, 1 111). Chital or Spotted deer

1777 Cervus axis Erxleben. Banks of the Ganges.

1792 Cervus axis maculates Kerr. Banks of the

Ganges.

1829 Cervus axis var. indicus G. Fischer.

1829 Cervus axis var. ceylonensis G. Fischer.

Ceylon.

1831 Cervus nudipalpebr a Ogilby. Banks of the

Ganges.

1842 Axis major Hodgson.

1842 Axis minor Hodgson.

1905 Cervus {Rusa) axis zeylanicus Lydekker.

Ceylon.

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Pocock ( 1 943b) did not admit the mainland

and Sri Lankan populations as distinct

subspecies.

Genus Hyelaphus Sundevall, 1 846

This is usually placed as, at most, a

subgenus of Axis , but Grubb ( 1 990) regards it as a

full genus, retaining the primitive antler plan of

the Cervus group.

Hyelaphus porcinus (Zimmermann, 1780). Hog-

deer

Hyelaphus porcinus porcinus (Zimmermann, 1 780)

1780 Cervus porcinus Zimmermann, 1780.

Bengal.

1883 Cervus minor Sclater. India. Not of Hodgson

(1842).

From Sind and the Ganga north into Uttar

Pradesh (erstwhile United Provinces), Nepal and

northeastern India.

Hyelaphus porcinus oryzus (Kelaart, 1 852)

1 852 Axis oryzus Kelaart. Ceylon.

Pocock (1943b) separated the Sri Lankan

subspecies on the basis of being smaller, with

poorly developed antlers, without spots in the

adult and with no marked seasonal change in

colour. He admitted that the Sri Lankan population

might actually be introduced (and the absence of

hog-deer from southern India gives pause for

thought). Whether there are any valid genetically

based differences between them should be tested

by future research.

FAMILY BOVIDAE

Genus Bos Linnaeus, 1 758

Bos gaurus Hamilton Smith, 1 827. Gaur or Indian

bison

In historic times, gaur occurred in Sri Lanka

(Knox 1681). The name guavera Kerr, 1 792, which

was based on Knox’s description, was

suppressed by the International Commission on

Zoological Nomenclature (1985) in order to

preserve the priority of the familiar name gaurus.

The subspecies as listed here, and the use

of the name laosiensis in place of the more familiar

readei for the Southeast Asian subspecies, are

based on a revision in preparation by Grubb and

Groves.

Bos gaurus gaurus Hamilton Smith, 1 827

1827 Bos gaurus Hamilton Smith. Mainpat,

Sarguja Tributary States, India.

1827 Bosgour Hard wi eke. Mountainous district

of Ramgurh and tableland of “Sirgoojas.”

1837 Bibos subhemachalus Hodgson. Sal

Forests, Nepal.

1837 Bibos cavifrons Hodgson. Substitute for

subhemachalus.

1842 Bibos concavifrons Roulin. Substitute for

cavifrons.

1846 Bos gaur Sundevall. Substitute for gaurus.

1851 Bos asseel Horsfield. No locality.

Indian gaur are rather smaller (mean skull

length of males, 542 mm) than Southeast Asian

gaur, but with relatively longer nasal bones (222

mm) and wider occiput. Northern Indian gaur do

not differ in size from the from the peninsular ones,

but have a markedly narrower span across the

horns in males (mean values are 795 mm in northern

India, 846 mm in the south). The range of this

subspecies reaches Nepal and Sikkim.

Bos gaurus laosiensis (Heude, 1901)

1901 Gauribos laosiensis Heude. Camoun, in

the mountains separating Laos and Tonkin.

1903 Bos gaurus readei Lydekker. Myitkyina,

Burma.

Skulls from Assam resemble Southeast

Asian gaur. The Southeast Asian subspecies is

larger than Indian gaur (skull length averages 587

mm in male), with relatively short nasals (mean

228 mm); and the occiput is narrower. The horn

tips are less turned in relative to the horn span.

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Specimens from Bhutan and Chittagong are

intermediate between the two subspecies.

Bos javanicus d’ Alton ( 1 823). Banteng

1 898 Bos sondaicus birmaniciis Lydekker. Burma.

There have been constant reports of

banteng in Manipur, but there is as yet no

confirmation.

Bos mutus (Przewalski, 1883). Wild yak

1883 Poephagus mutus Przewalski. Western Nan

Shan, Ganssu.

Yak enter Indian territory in Ladakh.

Bubalus arnee (Kerr, 1 792). Wild buffalo or Ama

The subspecies of the Subcontinent, and

their characters, are after Groves (1996). The

question of whether the Sri Lankan wild buffaloes

are wild or feral is unresolved.

Bubalus arnee arnee (Kerr, 1792)

1792 Bos arnee Kerr. Restricted to Kuch Behar

by Harper (1940).

1807 Bos ami Blumenbach. Mountains of north

Hindustan.

1852 Bubalus arna var. macrocerus Gray. After

a nomen nudum of Hodgson (1841). “India”

(probably Nepal).

1912 Bubalus bubalus septentrionalis

Matschie. Kuckri-Muckri, an island in the

Bay of Bengal, off the Sunderbans.

Formerly occurred from the Sunderbans into

Nepal, Madhya Pradesh and Andhra Pradesh; still

occurs in Raipur and Bastar districts (Madhya

Pradesh) and Kosi Tappu Reserve (Nepal), but in

very small numbers, probably less than 200. Skull

length usually under 570 mm, hom span under

1,200 mm. Black, with contrasting white limbs

below knees and hocks, and white muzzle; tail

reaches down to hocks.

Bubalus arnee fulvus (Blanford, 1891)

1 89 1 Bos arnee fulvus Blanford. Mishmi Hills.

Brahmaputra Valley, formerly from Koch

Behar to Mishmi Hills and south to Chittagong

Hills; still occurs sporadically along the

Brahmaputra and in Manas (Choudhury 1997).

Larger, skull length usually over 570 mm, hom

span over 1,100 mm. Grey or brownish grey, and

less contrastingly white on limbs; tail falls short

of hocks.

Genus Boselaphus de Blainville, 1816

Boselaphus tragocamelus (Pallas, 1766). Nilgai

1766 Antilope tragocamelus Pallas. Plains of

peninsular India.

1777 Antilope albipes Erxleben. India.

1777 Antilope picta Pallas. India.

1827 Damalis risia Hamilton Smith. Substitute

for picta.

1837 Tragelaphus hippelaphus Ogilby.

Substitute for picta.

1846 Portax tragelaphus Sundevall. Error for

tragocamelus.

There is very little geographic variation in

male nilgai skulls, from localities as far apart as

Gir, Rajasthan, Bhopal, Kanpur and Patna. A skull

from Bengal, however, is overall one of the

smallest, with the shortest horns, but the widest

in bizygomatic breadth [157 mm, cf. 1 3 1 - 148 mm

(n=20) from other localities].

Genus Tetracerus Leach, 1 825

Tetracerus quadricornis (de Blainville, 1816).

Chowsingha or Four-homed antelope

Unlike the nilgai, the chowsingha has well-

marked geographic variation, and we can

distinguish three subspecies. The metric

differences are given in Table 1 .

Tetracerus quadricornis quadricornis (de

Blainville, 1816)

1816 Cerophorus (Cervicapra) quadricornis de

Blainville. Plains of peninsular India.

1825 Antilope chickara Hardwicke. Bengal,

Bihar and Orissa.

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TAXONOMY OF INDIAN UNGULATES

1828 Antilope tetracornis Hodgson. No locality.

A relatively large form with narrow nasal

bones. Four long horns. Colour yellow-fawn

above, creamy or creamy-fawn below, this tone

often confined to midline; forelimbs markedly

blackened down anterior surface; median dorsal

region darker than rest of upperside; tail long,

with bushy white tip. Nose diffusely darker. I have

seen specimens from Rajasthan, Gujarat,

Maharashtra, Madhya Pradesh, Bihar, Bengal, the

eastern Godavari in Andhra Pradesh, and Sandur

in Karnataka.

Tetracerus quadricornis iodes Hodgson, 1 847

1847 Tetracerus iodes Hodgson. Sal Forests of

sub-Himalayan region.

1847 Tetracerus paccerois Hodgson. Same

locality.

Similar in size, but with wider nasals; horns

smaller. Colour variably fawn above, light fawn

below, this tone often confined to a very narrow

midline streak; foreleg with at most a vague, dark

brown line, which may be interrupted at knee;

median dorsal region not darkened. Nose diffusely

darker. The four skins and skulls I have seen are

from Nepal (3) and Champa ( 1 ).

Tetracerus quadricornis subquadricornis (Gray,

1843)

1839 Antilope sub-4-cornutus Elliot. Southern

Mahratta country.

1843 Tetracerus subquadricornis Gray.

1847 Teracerus subquadricornutus Hodgson.

Emendation of sub-4-cornutus.

Smaller in size, with very broad nasals; only

a single pair of horns, which are rather long.

Colour varying from red-fawn to more olive above;

underside whitish in midline, pale yellow laterally;

foreleg line very vague or absent; median dorsal

region slightly darkened; midline of nose may be

sharply darker; tail short. The four male and two

female skulls and 9 skins are from Palkonda Hills,

Madras(= Chennai), Dharwar and Kuckanalla.

Genus Procapra Hodgson, 1 846

Procapra picticaudata Hodgson, 1846. Tibetan

gazelle or Goa

1846 Procapra picticaudata Hodgson. Hundes,

Tibet.

Like yak, goa enter the Indian region in

Ladakh. The species is monotypic.

Genus Gazella de Blainville, 1816

Gazella bennettii (Sykes, 1831). Chinkara or

Indian gazelle

Groves ( 1 993b) revised this species in Iran,

and the accumulation of specimens from India

now enables me to do the same for the species in

the Subcontinent, including the description of a

new subspecies. Apart from the three described

subspecies, I have good-sized samples from two

other regions, the Salt Range (including Punjab

and Haryana) and the Ganga Valley, and I here

describe the first of these as a new subspecies.

Their measurements are given in Table 2.

Gazella bennettii bennettii (Sykes, 1831)

1831 Antilope bennettii Sykes. Deccan.

1843 Antilope hazenna I. Geoffroy St. Hilaire.

Malwa.

Table 1: Differences in measurements of Tetracerus quadricornis subspecies

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Table 2: Comparative measurements of Gazella spp.

TAXONOMY OF INDIAN UNGULATES

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Preorbital length 92.7 ±4.79 (5) 102.5 (2) 95.5 (2) 93.3 ±4.55 (6) 95.6 ±3.36 (5) 89 (1)

Toothrow length 55.7 ±2.06 (7) 58.7 ±2.52 (3) 56.0 ±4.55 *(4) 54.7 ±3.59 (7) 56.6 ±3.05 (5) 56.7 ±2.08 (3)

TAXONOMY OF INDIAN UNGULATES

From Ganga Valley and its vicinity (Muttra,

Gwalior, Jhansi, Etawah, Hamirpur, Allahabad,

Rewa, Jabalpur, Nimar, Asirgarh, Palamau,

Jagodih, Hazaribagh) and the Deccan

(Hyderabad, Haturna, Indore, Bhopal,

Khandesh). 14 skins, 3 head-skins, 4 1 full or partial

skulls (including frontlets and horns). Head, neck,

limbs and most of flanks dull reddish brown;

median dorsal region and lower flanks (abutting

the white underparts) are abruptly darker, tawny.

Winter coat is distinctly longer than summer, but

the colour does not differ. The Ganga Valley

gazelles are of much larger skull size, with more

widely spreading horns (but of equal size), than

those of the Deccan, and it may be appropriate to

separate them into two subspecies if further

material from the Deccan confirms these trends;

but the pelage characters are identical.

Gazella bennettii christyi Blyth, 1842

1 842 Gazella christii Blyth. Thar Desert.

From Kutch and Saurashtra, as far east as

Ahmedabad. 12 skins, 39 full or partial skulls

(including frontlets and horns). Most skins are

very pale, almost silvery drab brown with only

very restricted median dorsal and lower flank

zones being slightly darker than the rest of the

body; a few are slightly richer, with more contrast.

Winter coat hardly longer than summer. More

sexually dimorphic than Gazella bennettii

bennettii : male, but not female, is larger and

longer-horned.

Gazella bennettii fuscifrons Blanford, 1 873

1873 Gazella fuscifrons Blanford. Jalk,

southeastern Iran.

From Baluchistan (Iran and Pakistan) to

Sind and Rajasthan (Bikaner, Jodhpur, Jaisalmer).

16 skins, 36 full or partial skulls (including

frontlets and horns). Colour in winter dark grayish

sandy, often with a distinct brown band edging

the white of the underparts but without much

median dorsal darkening; in summer, brownish

bay-fawn. Coat with very long hair in winter. Size

as in Ganga Valley population of G.b. bennettii ,

but with shorter, broader nasals, especially in

males; males have shorter, more upright horns,

but females have longer horns than any other

subspecies in the Subcontinent. The cranial and

horn metrics of Rajasthan specimens are nearly

identical to those from Baluchistan and Sind, and

pelage characters are the same.

Gazella bennettii salinarum s.s. nov.

Type: BM(NH) 25. 10.3.5, skin and skull of

adult female from Ara, Salt Range, 700 m above

msl. Specimens seen: 3 skins, 1 head-skin, 1 7 male

and 3 female skulls. Haripur, Pind Didan Khan,

Faridkot (skins and skulls), “Punjab” (skull and

head-skin), Barra Jandhi, Sirsa, Gurgaon, Rohtak,

Hissar (skulls only). British Museum (Natural

History), Museum Alexander Koenig (Bonn),

Zoological Survey of India (Kolkata), Royal

Scottish Museum (Edinburgh).

From the Salt Range east as far as Delhi.

Skin rich tobacco-brown, with no

contrasting zone on the midback, but a

contrasting flank band. The largest subspecies

all round, in both sexes, with especially long

nasals. Skull length in males is from 1 90 to 203

mm (only one specimen under 193); that of

G.b. christyi, the next largest subspecies, is

187 to 195 mm (only one specimen above 193).

Skull length in the two females is 186 and 197

mm; in both the two females of G.b. christyi it is

184 mm.

Gazella subgutturosa (Giildenstaedt, 1780).

Goitred gazelle

1780 Antilope subgutturosa Giildenstaedt.

Tbilisi, Georgia.

1 have seen a herd from Pakistan in the

private collection of Sheikhs Khalid and Hassan

al-Thani at A1 Wabra, near Doha, Qatar. They do

not appear distinguishable in external characters

from the nominotypical subspecies of the Iranian

plateau, but this must remain a subjective

judgement until specimens can be examined.

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Genus Antilope Linnaeus, 1758

Antilope cervicapra (Linnaeus, 1758). Blackbuck

On geographic variation in this species, see

Groves (1982). Dr. E.C. Mungall (pers. comm.) is

examining the status of blackbuck from Point

Calimere, which may represent a further

subspecies, as yet undescribed.

Antilope cervicapra cervicapra (Linnaeus, 1758)

1758 Capra cervicapra Linnaeus. Inland of

Trivandrum, restricted by Zukowsky, 1 927).

1776 Antilope rupicapr a Muller. Bengal.

1830 Antilope bilineata Gray. Bengal.

1843 Cervicapra bezoartica Gray. India.

1927 Antilope hagenbecki Zukowsky. Bengal.

From approximately east and south of the

Delhi region, south to Chennai (earlier Madras)

and Karnataka (Rannebenur) and to Vallanadu

Reserve Forests in Tirunelveli district, and east

to Bihar and Bengal. Smaller, with short fine hair;

the dark colour runs all down the limbs to the

hoofs, and the white eye-ring is narrowed above

the eye. Horns are relatively short, not very

divergent, and have a relatively open spiral.

Antilope cervicapra rajputanae Zukowsky, 1 927

1927 Antilope rajputanae Zukowsky. Bahawalpur,

borders of Rajasthan and Panjab.

1928 Antilope centralis Zukowsky. Gwalior.

From west of the Delhi region, to Saurashtra

and Vadodara, Amritsar and into Pakistan. Larger,

with longer, roughened pelage; the male in the

breeding season has a grey sheen; the shanks are

largely white, with little or no extension of the dark

colour from the upper limb segments; the white eye-

ring is broad all around the eye. Homs tend to be

longer, more divergent and more closely spiralled.

Genus Pantholops Hodgson, 1 834

Pantholops hodgsonii (Abel, 1 826). Chiru

1826 Antelope hodgsonii Abel. Tingri Maidan,

Arun Valley, Kooti Pass, Tibet.

1827 Antilope kemas Hamilton Smith. Central

Asia.

1827 Antilope chiru Lesson. Nepal.

This species enters the Indian region in

Ladakh; whether they have ever genuinely

occurred in Nepal is unclear.

Genus Ovis Linnaeus, 1758

Ovis vignei Blyth, 1841. Urial

Geographic variation in this species is after

Schaller(1977).

Ovis vignei vignei Blyth, 1841. Ladakh urial

1841 Ovis vignei Blyth. Astor, Kashmir.

1854 Ovis montana Cunningham. Ladakh.

From Ladakh; in Zanskar, it is sympatric

with O. ammon. Homs of male thick, 230-260 mm

in circumference, rise more steeply from the

head and more corrugated than that of other

urials, and tend to be supracervical (hetero-

nymous), but sickle-shaped and homonymous

occasionally occur. Colour brownish or greyish,

with a grey saddle, occasionally with a dark line

in front.

Ovis vignei punjabiensis Lydekker, 1913. Punjab

urial

1913 Ovis vignei punjabiensis Lydekker. Salt

Range, Punjab.

From the Salt and Kala Chitta Ranges,

between the Indus and Jhelum rivers. Horns of

male more slender ( 1 90-240 mm in circumference),

sickle-shaped, with either tighter or looser curl.

More reddish, with a two-coloured saddle patch

(but this may be white only, or absent). The ruff,

usually black in all urials, is occasionally white in

this subspecies.

Ovis vignei cycloceros Hutton, 1 842. Afghan urial

1842 Ovis cycloceros Hutton. Hazara Hills, near

Kandahar, Afghanistan.

1877 Ovis blanfordi Hume. Hills above Bolan

Pass, near Kelat, Baluchistan.

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From Pakistan, west of the Indus, into

Afghanistan. Horns of male 210-270 mm in

circumference, usually homonymous. Reddish-

buff to yellowish or light brown, with a small black

saddle spot.

Ovis ammon Linnaeus, 1758. Argali

Geographic variation in this species follows

Geist(1991).

Ovis ammon hodgsonii Blyth, 1841. Tibetan argali

1841 Ovis hodgsonii Blyth. Tibet, on Nepal

frontier.

1841 Ovis ammonoides Hodgson. Himalayan

region.

1852 Caprovis b am b her a Gray. Nepal.

1874 Ovis brookei Ward. Ladakh.

In the Subcontinent extends into Ladakh,

Sikkim and, formerly, Nepal. In mature rams, the

long-haired, light-coloured ruff is sharply set off

from the dark shoulders; rump patch distinctly

set off dorsally, surrounds the tail; dark partial

flank band, separating light belly from dark body;

dark stripe from chest to hooves contrasts with

light colour of rest of limbs. Face wholly dark in

young, with light rostrum in old males. Homs of

adult males have less developed combat edge (on

external angle), less everted; of adult females long,

thick, angular, usually about 450 mm long,

occasionally up to 600 mm. Tail tiny, up to about

83 mm including hair, 58 mm without.

Ovis ammon polii Blyth, 1841. Pamir argali or

Marco Polo’s sheep

1841 Ovis polii Blyth. Near sources of Syr Darya,

Pamir.

In the Subcontinent, extends to Hunza,

Pakistan. Neck ruff of mature rams shorter, blends

gradually with body tone; no sharp contrasts of

pelage, grading between dark upper parts and

white underparts and limbs; white of rump patch

usually not sharply set off above, and continuous

with an extensive white zone on thighs, hindlegs,

underside and forelegs; flank stripe narrow, only

slightly darker than body, extends along flank and

tops of limbs. Face light. Horns of adult males

with well-developed combat edge, relatively thin,

widely flaring, arise at a shallower angle; of ewes,

relatively short and thin. Tail longer, 1 20- 1 50 mm

long including hair, 1 00 mm without.

Genus Pseudois Hodgson, 1 846

Pseudois nayaur (Hodgson, 1833). Bharal or Blue

sheep

1833 Ovis nayaur Hodgson. Tibetan frontier of

Nepal.

1835 Ovis nahoor Hodgson. Alternative name.

1841 Ovis burrhel Blyth. Boorendo Pass.

Extends from the Karakoram eastward

through Ladakh all along the Himalayan slopes

from 3,500 to 5,500 m above msl (Schaller 1 977).

Genus Capra Linnaeus, 1758

Capra sibirica (Pallas, 1776). Siberian ibex

Capra sibirica sakeen Blyth, 1 842

1842 Capra sakeen Blyth. Tibetan slopes of

Himalayas.

1844 Aegoceros skyn Wagner. Baltistan,

Kashmir.

1886 Capra dauvergnii Sterndale. Hills north of

Kishenganga river, Kashmir.

1900 Capra sibirica wardi Lydekker. Braldu,

Baltistan, near Baltoro Glacier.

1906 Capra sibirica pedri Lorenz. Gilgit,

Kashmir.

191 1 Capra sibirica filippii Camerano. Lahul.

In the region, ibex are found on high, rocky

cliffs, seldom below 3,000 m, from the Hindu Kush

east only to the Sutlej river. Kashmir ibex resemble

those from the Tianshan ( Capra sibirica

alaiana ), and differ from those from the Altai

(C.s. sibirica ), in their large size and long nasal

bones, and the large white saddle on the back,

but differ in their shorter horns, and the failure of

the saddle to extend to the haunches.

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Capra aegagrus Erxleben, 1777. Bezoargoat

Capra aegagrus blythi Hume, 1875.

1875 Capra blythi Hume. Sind.

1913 Capra falconeri chialtanensis Lydekker.

Chiltan Range, near Quetta, Baluchistan.

Bezoar goats occur on the low desert

ranges of Pakistan west of the Indus, both

sympatric with markhor (as on the Murdar Range,

near Quetta, and in the Gadabar Ghar Range, east

of Loralai) and separately (south of about 30° N).

On some of these ranges, along with the common

scimitar-homed form, occur screw-homed Capra ,

called Chiltan goats by Schaller and Khan (1975).

These were originally described as a subspecies

of markhor by Lydekker ( Capra falconeri

chialtanensis ), but their status has been queried

and discussed by both Schaller and Khan (1975)

and Roberts (1977).

According to Schaller and Khan ( 1 975 ; see

also Schaller 1977), Chiltan goats are found

intermingled in the same herds as scimitar-homed

goats in the Zahri, Koh-i-Maran and Dilband-

Moro ranges and districts; as the only Capra on

the Chiltan Range; and on the Murdar Range,

where goat and markhor are sympatric.

Schaller and Khan (1975) consider that

Chiltan goats are a polymorphic variant of Capra

aegagrus. They note that the horns lack the

markhor ’s posterior keel, and the cross-section

is oval like a scimitar-horned bezoar’s horn;

there is no neck ruff; and the pelage resembles a

bezoar. Where it coexists with scimitar-homed

goats, the two are found intermingled in the same

herds.

For Roberts (1977), however, the Chiltan

goat is a hybrid between bezoar and markhor.

The female resembles markhor, being reddish-

grey with a dorsal stripe and creamy legs with a

pattern on the front; but Roberts admits that

bezoar can be reddish-grey as well, although they

have a dark face pattern and the legs are greyer,

less creamy. The male, he agrees, resembles

bezoar more, becoming more and more grey-white

with age, and some have a black chest and

shoulder-stripe like the bezoar. He says that there

have been some local extinctions of bezoar or

markhor, and implies that early in the 20th century

the distribution of Chiltan goats encompassed

all the ranges where bezoar and markhor were

sympatric.

The type of chialtanensis is a frontlet with

horns, but there is also a nearly complete skull of

a Chiltan goat in the Natural History Museum

(London), BM 67.795. This is as big as a markhor,

and bigger than any bezoar, but has big teeth like

a bezoar (Table 3). In my opinion, the Chiltan goat

is most likely to be a hybrid.

Table 3: Comparison of Chiltan goat with markhor

and with scimitar-horned bezoar

Capra falconeri (Wagner, 1839). Markhor

Geographic variation after Schaller and

Khan (1975).

Capra falconeri falconeri (Wagner, 1839)

1839 Aegoceros (Capra) falconeri Wagner.

Kashmir (restricted to Astor by Lydekker,

1913).

1898 Capra falconeri cashmiriensis Lydekker.

Pir Panjal Range, Kashmir.

1958 Capra falconeri chitralensis Cobb. Chitral.

1958 Capra falconeri gilgitensis Cobb. Gilgit

watershed. Nomen nudum.

From eastern Afghanistan east to Kashmir.

Horns are more curved, wider across the tips; size

is larger, and the coat is longer and silkier, with a

well-developed neck ruff. The “Kashmir” horn

type, moderately divergent and with 2 or 3 twists,

occurs in Afghanistan, Chitral, Dir, and Swat, and

again in Shamsberi, Kaj-i-Nag and Pir Panjal; the

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more flaring, less twisted ( 1 .5 turns) “Astor” type

predominates in intervening localities.

Capra falconeri megaceros Hutton, 1 842

1842 Capra megaceros Hutton. Kandahar,

Afghanistan.

1875 Capra jerdoni Hume. Suleiman Range,

Punjab, Pakistan.

From the northern hill ranges of Pakistan.

Homs are less curved (more closely corkscrewed,

less divergent; size is smaller, and the pelage is

shorter, with (usually) no distinct ruff. Two horn

types, the more open-spiralled Kabul type, and

the more corkscrew Sulaiman type, occur together

widely.

Genus Hemitragus Hodgson, 1 84 1

Hemitragus jemlahicus (Hamilton Smith, 1 826).

Himalayan tahr

1826 Capra jemlahica Hamilton Smith. Jemla

Hills, Nepal.

1833 Capra jharal Hodgson. Nepal.

1836 Capra quadrimammis Hodgson. Nepal.

1845 Capra tubericornis Schinz. Substitute for

jemlahica.

1944 Hemitragus jemlahicus schaeferi Pohle.

10 km southwest of Chuntang, Sikkim.

The range, according to Schaller (1977),

extends from about 40 km west of Banihal Pass, in

the Pir Panjal, east into Bhutan, in cliff country

where they migrate seasonally between forest and

alpine zones.

Hemitragus hylocrius (Ogilby, 1838). Nilgiri

tahr

1838 Kemas hylocrius Ogilby. Nilgiri Hills.

1842 Capra (Ibex) warryato Gray. Nilgiri

Hills.

Found in the highlands of the Tamil Nadu/

Kerala borderlands, presently surviving mainly

in the Nilgiri Hills, High Range and Highwavy

Mountains; in rolling country at 1,200 to 2,600 m

above msl (Schaller 1977).

Genus Capricornis Ogilby, 1837

The taxonomy of this genus follows Grubb

and Groves (in prep.). Himalayan serow are sharply

distinct from those in China or Southeast Asia,

and there is no merit in keeping them in the same

species.

Capricornis thar (Hodgson, 1831). Himalayan

serow

1831 Antilope thar Hodgson. Nepal Himalaya.

1832 Antilope bubalina Hodgson. Nepal.

1842 Nemorhaedus or Kemas proclivus or thar

Hodgson. Nomen nudum.

1908 Capricornis sumatraensis humei Pocock.

Kashmir.

1908 Capricornis sumatraensis rodoni Pocock.

Chamba, Panjab.

1908 Capricornis sumatraensis jamrachi

Pocock. Kalimpong, near Darjiling.

From Kashmir, almost at the Afghanistan

border, east as far as the Lushai Hills, Assam.

This species is black, with a buffy tone as the

black tips wear off and reveal the lighter hair bases;

a long mane, mixed black and white; below, sharply

cream-buff; broadly white over nose, or only on

lip margins; white extends backwards along

jawlines in a ‘V’ shape, or interramal region

completely white, or occasionally white tones are

nearly absent; legs creamy-white from below

knees and hocks.

An erythristic population (subspecies or

separate species?) occurs in the Garo, Mishmi and

Naga Hills. This is close to C. thar in its fairly long

pelage, white hair-bases, and black and white mane;

it is different from Capricornis ntbidus of northern

Burma, which differs in its very short pelage, the

black (not white or buflfy) hair-bases, the black dorsal

stripe, and the very short dark red mane.

Genus Nemorhaedus Hamilton Smith, 1 827

As in Capricornis , the taxonomy of this

genus follows Grubb and Groves (in prep.).

360

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG. -DEC. 2003

TAXONOMY OF INDIAN UNGULATES

Nemorhaedus goral (Hardwicke, 1825). Himalayan

goral

Nemorhaedus goral goral (Hardwicke, 1825).

Brown goral

1825 Antilope goral Hardwicke. Kathmandu.

1827 Antilope duvaucelii Hamilton Smith, 1 827.

1908 Naemorhedus hodgsoniPocock. Sikkim.

From Bhutan west to about Nainital.

Medium brown with black hair tips, giving a

hare-like effect; or slightly greyer, to grey-

brown; or pale or dark fawn; legs browner, to

very bright tan, or white on forelegs only.

Underside paler grey. Throat and chin variably

white, may be interrupted under jaw; lips white.

Dorsal stripe usually weak, may fade behind

withers.

Nemorhaedus goral bedfordi (Lydekker, 1 905).

Grey goral

1905 Urotragus bedfordi Lydekker. Dharmsala.

From Chamba and Kulu, west into Kashmir.

Grey to grey-brown to yellow-grey; legs lighter,

yellower, with dark brown line down front, fading

on pasterns. Underside off-white. Throat and chin

creamy the whole way. Merest trace (if that) of

dorsal stripe.

Refer

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103 pp.

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Asiat. Soc. (Ceylon) 34(91): 126-162.

Deraniyagala, P.E.P. (1955): Some extinct elephants,

their relatives and the two living species. National

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Dolan, J.M. (1988): A deer of many lands - a guide to

the subspecies of the red deer Cervus elaphus L.

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Geist, V. (1991): On the taxonomy of giant sheep ( Ovis

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Geist, V. (1998): Deer of the World: their Evolution,

Genus Budorcas Hodgson, 1850

Takin of India and Bhutan differ from those

of China in their smaller horns and in the brown

rather than golden colour, except for a lighter,

yellow-toned saddle. They become blacker with

age, this colour developing earliest on the

underside, haunches and limbs.

Budorcas taxicolor Hodgson, 1850. Takin

Budorcas taxicolor taxicolor Hodgson, 1850

1850 Budorcas taxicolor Hodgson. Mishmi Hills.

Known from the Mishmi Hills (extending

into Chinese and Burmese territory). Males much

larger than females (biorbital breadth over 190 mm

in males, less than 1 76 in females); teeth relatively

small, maxillary toothrow length 114-122 mm; horns

of males very large, their span 341-411 mm.

Budorcas taxicolor whitei Lydekker, 1 907

1907 Budorcas taxicolor whitei Lydekker.

Bhutan.

From Sikkim and Bhutan. Males as small as

females (biorbital breadth 166-182 mm in both

sexes); maxillary toothrow length 120-124 mm;

horns of males smaller, their span 297-357 mm.

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Behavior, and Ecology. Stackpole Books,

Mechanicsburg, PA, USA.

Ghosh, M. (1988): The craniology and dentition in the

pigmy hog, with a note on the generic status of

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Goodman, M., C.A. Porter, J. Czelusniak, S.L. Page,

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Groves (1998): Toward a phylogenetic

classification of Primates based on DNA evidence

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Asia. Saugetierlcundl. Mitt. 15: 221-237.

Groves, C.P. (1981): Ancestors for the Pigs: taxonomy

and phytogeny of the genus. Sus. Technical Bulletin

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of Pacific Studies, Australian National University.

Groves, C.P. (1982): A note on geographic variation in

the Indian Blackbuck ( Antilope cervicapra

Linnaeus, 1758). Rec. zool. Surv. India 79:

489-503.

Groves, C.P. (1983): Geographic variation in the

barasingha or swamp deer ( Cervus duvauceli).

J. Bombay nat. Hist. Soc. 79: 620-629.

Groves, C.P. (1993a): Testing rhinoceros subspecies by

multivariate analysis. Pp. 92-100. In: Rhinoceros

Biology and Conservation (Ed.: Ryder, O.A.).

Zoological Society of San Diego, San Diego.

Groves, C.P. (1993b): The chinkara ( Gazella bennetti )

in Iran, with the description of two new subspecies.

J. Sci. I. R. Iran 4: 166-178.

Groves, C.P. (1996): The taxonomy of the Asian wild

buffalo from the Asian mainland. Z. f.

Saugetierkunde 61: 327-338.

Groves, C.P. (2001): Primate Taxonomy. Smithsonian

Institution Press, Washington, D.C. vii + 350 pp.

Groves, C.P. & S. Chakraborty (1983): The Calcutta

collection of Asian rhinoceros. Rec. zool. Surv.

India 80: 251-263.

Groves, C.P. & P. Grubb (1987): Relationships of living

deer. Pp. 21-59. In: Biology and Management of

the Cervidae (Ed.: Wemmer, C.M.). Smithsonian

Institution Press, Washington, D.C.

Groves, C.P. & P. Grubb (1990): Muntiacidae. Pp. 134-

168. In: Horns, Pronghorns and Antlers (Eds.:

Bubenik, G.A. & A.B. Bubenik). Springer Verlag,

New York.

Groves, C.P. & V. Mazak (1967): On some taxonomic

problems of Asiatic wild asses with a description

of a new subspecies (Perissodactyla, Equidae).

Z. f. Saugetierkunde 32: 321-355.

Groves, C.P., Y. Wang & P. Grubb (1995): Taxonomy

of musk-deer, genus Moschus (Moschidae,

Mammalia). Acta Theriol. Sin. 15: 181-197.

Grubb, P. (1982): The systematics of Sino-Himalayan

musk deer {Moschus), with particular reference to

the species described by B.H. Hodgson.

Saugetierkundl. Mitt. 30: 127-135.

Grubb, P. (1990): Cervidae of South Asia. Pp. 169-179.

In: Homs, Pronghorns and Antlers (Eds.: Bubenik,

G.A. & A.B. Bubenik). Springer Verlag, New York.

International Commission on Zoological Nomenclature

(1985): Opinion 1348: Bos gaurus H. Smith, 1827

(Mammalia, Artiodactyla): conserved. Bull. Zool.

Nomen. 42: 279-280.

Knox, R. (1681): An Historical Relation of the Island

Ceylon in the East Indies. Reprinted 1983,

Navrang, New Delhi.

Pocock, R.I. (1943a): The larger deer of British India.

Part III. The Sambar ( Rusa ). J. Bombay nat. Hist.

Soc. 44: 27-37.

Pocock, R.I. (1943b): The larger deer of British India.

Part IV. The Chital (Axis) and the Hog-deer

(Hyelaphus). J. Bombay nat. Hist. Soc. 44: 169-

178.

Roberts, T.J. (1977): The Mammals of Pakistan. Ernest

Benn Ltd, London. 361 pp.

Rookmaaker, L.C. (1980): The distribution of the

rhinoceros in eastern India, Bangladesh, China and

the Indo-Chinese region. Zool. Anz. 205: 253-268.

Rookmaaker, L.C. (1983): Historical notes on the

taxonomy and nomenclature of the recent

Rhinocerotidae (Mammalia, Perissodactyla).

Beaufortia 33: 37-51.

Rookmaaker, L.C. (1998): The sources of Linnaeus on

the Rhinoceros. Svensk. Linnesallsk.s arssk.,

Uppsala 1996-1997: 61-80.

Rookmaaker, L.C. (2000): Records of the rhinoceros in

Pakistan and Afghanistan. Pakistan J. Zool. 32:

65-74.

Schaller, G.B. (1977): Mountain Monarchs: Wild Sheep

and Goats of the Himalaya. Chicago University

Press, Chicago, xviii + 425 pp.

Schaller, G.B. & S. Khan (1975): The status and

distribution of markhor ( Capra falconeri). Biol.

Conserv. 7: 185-198.

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DISAPPEARANCE OF THE WHITE- WINGED DUCK CAIR1NA SCUTULATA

FROM THE PABLAKHALI WILDLIFE SANCTUARY:

A SAGA OF LARGE-SCALE DESTRUCTION OF MIXED EVERGREEN

FOREST IN BANGLADESH

( With one text-figure and two plates)

Mohammad Ali Reza Khan1

Key words: Bangladesh, Biodiversity Bureau, clear-felling, jhum cultivation,

monoculture, Kaptai Lake, Pablakhali, Rangamati, tribal insurgency, white-winged duck

Bangladesh used to be a country with rich fauna and flora. During the rule of the British, largely

after the 1850s, the first commercial exploitation of natural resources vis-a-vis the forests was

introduced. This included ‘clear-felling’ of trees in the Sal Forests of central and northwestern

parts, and Semi-evergreen or Mixed Evergreen Forests of north and eastern parts of present

Bangladesh. The same tradition was carried forward by the Pakistani authorities who ruled the then

East Pakistan (now Bangladesh) from 1947 to 1971. After the independence of Bangladesh, in

December 1971, the forest officials kept following their predecessors in destroying forests. Forest

destruction became more rampant as the new government had few environmental policy decisions.

The situation was further aggravated by lumber poaching, land grabbing, and settling people in the

reserved forest areas leading to tribal insurgency. All this ultimately brought an end to the existence

of virgin Mixed Evergreen Forest in the northeastern parts of Bangladesh. The white-winged duck

Cairina scutulata is a victim of this process. This paper examines the process of destruction of the

forests and suggests measures to save the remaining habitats of the still surviving biodiversity

through the creation of a Wildlife/Biodiversity Bureau separating it completely from the Forest

Department that traditionally and wrongly acted against the interests of indigenous wildlife in the past.

Introduction

White-winged duck or white-winged wood

duck Cairina scutulata used to be present in a

section of the Mixed Evergreen Forest in the

Chittagong Hill Tracts (CHT) District of

Bangladesh bordering the Indian State of Mizoram

(Husain 1977, 1985, Husain and Haque 1982, Khan

1981, 1 983, 1 986). These reports were based on

observations made prior to 1 980. The population

was estimated to be 25 birds at that time.

Bangladesh has an area of roughly 147, 570

sq. km and lies between 20° 34' to 26° 38' N and

88° 0T to 92° 41' E. It is bounded on the west,

north and east by India, with a small portion of

the southeast comer bordering with Myanmar

(Fig. 1). The Bay of Bengal covers the entire

'Dubai Zoo, P.O. Box 67 Dubai, United Arab Emirates.

Email: [email protected]; markl [email protected]

southern border. Bangladesh occurs at the

confluence of three mighty rivers, the Padma

(Ganga), the Brahmaputra and the Meghna,

forming one of the largest deltas of the world. It

used to have three distinct forest types: Moist

Deciduous or Sal, Mixed or Semi-evergreen, and

the Mangrove Forest in the Sundarbans (Khan

1982).

Around 1970, the country had roughly 1 5%

land area under forests. However, following

independence, in December 1971, there was large-

scale systematic removal of forests by the

government Forest Department whose main aim

was, and possibly still is, to provide revenue to

the government exchequer through cutting and

selling of forest wood. This was followed by illicit

logging, poaching for firewood and illegal

conversion of forested land into agricultural fields

and human habitations, as well as road building

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG. -DEC. 2003

363

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Fig. 1 : Map of Bangladesh showing forest types (left) and Chittagong Hill Tracts (enlarged; right)

DISAPPEARANCE OF WHITE-WINGED DUCK FROM BANGLADESH

and industrial development. There was, and still

is, large-scale conversion of forested land into

monocultures of commercially viable plant species

such as Tectona grandis , Dipterocarpus

turbinatus , Syzygium grandis , Gmelina arborea ,

Hevea braziiensis , Hopea odorata, Michelia

champaca and Mesua ferrea.

Today there is virtually no virgin Sal or

Mixed Evergreen Forest in the country other than

the Sundarbans Mangrove Forest. In a recent

report, IUCN Bangladesh (2000) said that though

a current forest inventory is unavailable, it is

estimated that the forest cover has been reduced

by more than 50% since the 1970s. Estimates in

1990 revealed that Bangladesh has less than 0.02

ha of forest per capita — one of the lowest forest

to population ratios in the world. Presently, less

than 8% of the country is under forest cover —

that too is neither natural nor virgin (Plate 1 , Fig. 1 ).

Bangladesh had its best Mixed Evergreen

Forest in the CHT District on the banks of the hill

river, Kassalong (Fig. 1). The Kaptai Dam was

constructed here to generate hydroelectric power

in the 1 960s, as a result of which a large portion of

the valley forest was inundated. Even then, it had

a viable forest with an immense variety of wildlife

almost up to 1980, as described in old District

Gazetteers and Working Plans of the Forest

Department.

After 1980, bloody insurgency reigned,

continued up to January 1998 and subsided by

February 10 the same year, when tribal insurgents

surrendered their arms as per a peace treaty that

they signed with the Bangladesh Government.

Tribals objected to the settlement of plains-

dwelling Bengali people in the hilly areas of the

northeast.

Chittagong Hill Tracts District has been

bifurcated into Bandarban, Khagrachari and

Rangamati Hill Districts in the recent past.

White-winged duck (WWD) used to be

present in good numbers in the Pablakhali Wildlife

Sanctuary at the heart of Kassalong Valley Forest

under the CHT North Forest Division. It is a

threatened species as per birds to watch-2 of

BirdLife International (former ICBP - International

Council for Bird Preservation) and the IUCN

(International Union for Conservation of Nature

and Natural Resources) Red Data Book. Also it is

in Schedule I of CITES (Convention on

International Trade in Endangered Species of Wild

Fauna and Flora) (Green 1993) and Schedule III of

the Bangladesh Wildlife Preservation Act 1973.

Bangladesh is a party to the CITES. That means

this species cannot be traded, trapped or killed,

nationally or internationally.

Habitat Survey

I conducted a field survey of the known

habitats of the WWD from August 1 5 to 2 1 , 2000,

when a wildlife researcher from Dhaka helped me

in my fieldwork. I had also visited the area from

February 1 8 to 22, soon after the tribal insurgents

from the area surrendered their arms to the

Bangladesh Government on February 10, 1998.

From Rangamati town, that houses both the district

as well Forest Division headquarters, we went to

Mynimukh (22° 59.226' N; 92° 12.226’ E) about 60

km upstream northwards, along the Kassalong

river by a motor launch. From there we travelled

to Guishakhali and then to Rangi Para (22° 59.304'

N; 92° 14.531' E) by motorboat. Mostly settlers

live in these two spots. We met just two tribal

Chakmas here, but were able to talk to a

schoolteacher and a few local woodcutters. Later

on, we moved to Pablakhali Wildlife Sanctuary

(Fig. 1), about 10 km northeast of Mynimukh.

The Sanctuary’s office is at a village called

Amtali, which used to be a small outpost 20 years

ago with only a tiny grocery shop. Everyone in

the area used kerosene oil lanterns. Now it is a

bustling town with nearly three dozen shops, a

police station, high school and paramilitary camp.

Two diesel-powered privately run generators

supply electricity for 5 hours daily, from 5 p.m. to

10 p.m., to the shops at nominal charges. During

the course of the survey, I met the concerned

JOURNAL , BOMBAY NATURAL HISTORY SOCIETY, I00(2&3), AUG.-DEC. 2003

365

DISAPPEARANCE OF WHITE-WINGED DOCK FROM BANGLADESH

forest officials responsible for CHT North and

South Forest Divisions to get permission and

assistance in the field. A Dhaka-based wildlife

biologist, at least one Forest Department person,

one tribal Chakma and one boatman accompanied

us in all our surveys of the forest. They helped us

to find our way through various watercourses

and jungles as well as to communicate with the

tribal people in certain areas.

We walked through forests and/or travelled

by a slow-moving country boat fitted with an

improvised diesel water pump. It followed

watercourses surrounding the forests. We also

searched inundated banks of the Kassalong river,

which is the lifeline for all the forests surrounding

it, and probable roosting-nesting sites of the

WWD. In addition to this direct observation,

several tribal Chakmas, living in all parts of the

forest, plains-dwelling Bengali-speaking settlers

and some 50 people visiting market places were

also interviewed. We showed them the

photograph of a WWD and enquired whether they

had seen that particular duck, locally called ‘Bhadi

Hansh’ by the tribal Chakmas. We told them its

colour and size and explained to them where it

usually lived.

Earlier reports of the WWD were mostly

from a radius of 5 km from Amtali (23° 03.903' N;

92° 14.686' E). We surveyed the neighbourhood

of Amtali during the afternoon and the whole of

the next day, both on foot and by boat. We also

talked to the villagers, fishermen and boatmen,

mostly at night, in the market places where most

men of the area gathered to gossip and shop.

On the 4th day, we travelled through

Kassalong river and its flooded banks, a

marshland where there was a report of the sighting

of a pair of WWD by a passing European student,

and stopped at several prospective WWD

habitats at 23° 05.0 12' N, 92° 1 3.774' E; 23° 05.720'

N, 92° 1 3 .23 1 ' E; and 23° 06.906' N, 92° 1 3 .034' E.

We also prospected at the Nalbonia Beel (Beel =

low-lying marshland in Bengali) where there was

the possibility of its occurrence. We also stopped

and watched birds at Shishak Valley. We halted at

Marishsha-Baghaichari — the last stop for a

motor launch that plies between this and

Rangamati town. It is also the headquarters for

the Baghaichari Police Station. From there we

went to a roughly 5 sq. km marshland named

Ugalchari Beel or Lailla Ghona Beel (92° 12.453' E;

23° 08.467' N) and its neighbourhood, and also

Bot Tali. Ugalchari village is entirely hilly and

dominated by tribal Chakmas, while Bot Tali which

is on the bank of Kassalong river is full of settlers.

This Beel dries up in winter and is being used for

rice cultivation by irrigation. At night we looked

for frogs and talked to local folks.

Next morning we left Marishsha for Baghai

Haat (23° 1 6.690' N; 92° 09.09 l'E). We travel led the

first 20 km of the hill road by bus. Then we walked

for 8 km to reach Baghai Haat where we stayed for

two and a half days. By late afternoon we travelled

another 6 km to Ganga Ram (23° 1 8.073' N; 92° 10.031’

E), on the bank of Kassalong river, and the last stop

on our journey — beyond this there was no road,

and it was not safe for free movement as there was

no law enforcing authority there.

From Baghai Haat we made short field trips

in different directions and met tribal people who

generally congregated in hundreds every Sunday

at Baghai Haat to sell their products and to procure

provisions.

Results and Discussion

We did not come across any WWD not

only in the Pablakhali Wildlife Sanctuary and its

neighbourhood, but also in the entire valley of

the Kassalong river up to Marishsha. These

included three Beels or marshy areas — Nalbonia

Beel, Shishak Valley and Ugalchari Beel. Of these,

only the first one is known to hold some water all

year round while the others dry up during winter

and are used for cultivation of paddy and other

winter crops.

Among the persons interviewed, only one

Chakma villager from Dhoopchari Bazar, close to

366

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DISAPPEARANCE OF WHITE-WINGED DUCK FROM BANGLADESH

Khan, M.A.R.: Mixed Evergreen Forest in Bangladesh

Plate 1

Fig. 1 : There are no virgin Mixed Evergreen Forests in Bangladesh — this is one of the best patches

with regenerated vegetation

Fig. 2: Tribal jhum cultivators have mercilessly cut the trees in the forest, planted paddy,

and not even bothered to remove the logs

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367

DISAPPEARANCE OF WHITE-WINGED DUCK FROM BANGLADESH

Khan, M.A.R.: Mixed Evergreen Forest in Bangladesh

Plate 2

Fig. 3: Clearing reserved forest for housing plains-dwelling settlers and for jhum cultivation

by the tribal settlers

Fig. 4: An officially banned brickfield stands out as a reminder of the ruthless large-scale

destruction of the surrounding Mixed Evergreen Forest

368

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DISAPPEARANCE OF WHITE-WINGED DUCK FROM BANGLADESH

the Sanctuary, said that he had seen a pair of

WWD in Naibonia Beel during the previous

winter. However, we visited the same area and did

not come across any WWD, though we saw a

large flock of lesser whistling-duck Dendrocygna

javanica and a pair of cotton teal Nettapus

coromandelianus in the same area. We also noted

bronze-winged jacana Metopidius indicus and

chestnut bittern Ixobrychus cinnamomeus. There

were many domesticated ducks, mostly mallards

and muscovies.

As it was the breeding season for most of

the fishes, an official fishing ban was in place. A

few people were still fishing in the area for their

livelihood, but the fish catch was too scanty to

feed the local market. So, if there were WWD in

the area, it would have been sighted by both

Chakmas and Bengali fishermen. From Naibonia

Beel up to Ganga Ram, there was no report of

sighting of the WWD.

There is only one recent record of sighting

of a pair of WWD between Marishsha and

Pablakhali Wildlife Sanctuary, by Neville J. Ash

— a British student proceeding towards the

Sanctuary searching for bears — on June 5, 1999

(c. 23° 1 0.000' N; 92° 20.000' E), according to Baz

Hughes (of Threatened Waterfowl Specialist

Group, Siimbridge, UK, pers. comm.). We did not

see the duck there during the present survey.

Our past visits during the late 1970s and

early ’80s, and the present one, revealed a massive

change in the pattern of avifauna distribution in

the Sanctuary. First and foremost is the excessive

abundance of mynas and starlings ( Acridotheres

spp. and Sturnus spp.). We came across too many

common mynas (A. tristis ) and Asian pied

starlings (S. contra) that were very rare earlier and

mostly restricted to human habitations. During

the present visit, they were found in almost every

part of whatever remains of old forests that have

been largely converted into jhum cultivation

through the process of slash and bum.

Another group, the bee-eaters, has

flourished. We saw great numbers of blue-tailed

bee-eaters, small bee-eaters and chestnut-headed

bee-eaters ( Merops philippinus, M. orientalis ,

and M. leschenaulti ) all along the watercourses.

We also saw a blue-bearded bee-eater ( Nyctyomis

athertoni ) sallying for dragonflies along an oxbow

lake. We noted small blue and white-breasted

kingfishers ( Alcedo atthis and Halcyon

smyrnensis ) as very common, but missed stork-

billed and oriental dwarf kingfishers (//. capensis

and Ceyx erithacus ) noted earlier.

Nearly a dozen species of woodpeckers

used to live in this forest. We saw only three

species (fulvous-breasted pied woodpecker

Dendrocopos macei , lesser golden-backed

woodpecker Dinopium benghalense and little

scaly-beflied green woodpecker Ficus

xanthopygaeus ) that are also common outside

the forested area. We failed to notice any hombill.

Blue-throated and lineated barbets ( Megalaima

asiatica and M lineata) seemed to be more

common than the coppersmith or crimson-

breasted barbet (M haemacephala). We missed

greater and lesser racket-tailed drongos (Dicrurus

paradiseus and D. remifer) but black drongos

(D. macrocercus) were very common. We noticed

much fewer flycatchers and warblers than we saw

two decades ago.

During the whole survey period, we had

seen only two black-shouldered kites Elanus

caeruleus at the Sanctuary, and also jungle owlet

Glaucidium radiatum , spotted owlet Athene

hr am a, Eurasian scops-owl Otus scops and brown

hawk-owl Ninox scutulata , but no diurnal birds

of prey. Among herons, only the Indian pond-

heron Ardeola gray 'd was very common, while

the chestnut bittern Ixobrychus cinnamomeus

was not so common. We did not see any cormorant

( Phalacrocorax spp.), little grebe ( Tachybaptus

ruficollis ) or moorhens ( Porphyrio sp. and

Gallinula sp.).

The largest warbler in the area is the striated

marsh-warbler Megalurus palustris and it was as

common as before, but rewarding was the sighting

of quite a few pied bushchats with juveniles

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DISAPPEARANCE OF WHITE-WINGED DUCK FROM BANGLADESH

(Saxicola caprata). Red-vented bulbuls

Pycnonotus cafer outnumbered all the forest

bulbuls. Olive bulbul Iole virescens and red-

whiskered bulbuls Pycnonotus jocosus were also

common. Among flycatchers we saw a few black-

naped monarch-flycatchers Hypo thy mis azure a,

a niltava ( Niltava sp.) and another blue flycatcher

( Muscicapa sp.). Among sunbirds and

flowerpeckers, the purple-rumped sunbird

Nectarinia zeylonica and scarlet-backed

flowerpecker Dicaeum cruentatum dominated the

rest. The common iora Aegithina tiphia and

oriental white-eye Zosterops palpebrosus were

also very common.

Among mammals we observed the Irrawady

squirrel Callosciurus pygerythrus to be the most

common diurnal species. In addition, we noted

Malayan giant squirrel Ratufa bicolor , rhesus

macaque Macaca mulatto , a troop of capped

langur Trachypithecus pileatus , a small herd of 5-

6 elephants Elephas maximus , flying fox Pteropus

giganteus and false vampire bat Megaderma lyra

as common, and pygmy/Indian pipistrelle

Pipistrellus mimus as a very common species.

The reptiles we noted included checkered

keelback Xenochrophis piscator , common skink

Mabuya carinata. Bengal/common Indian

monitor Varanus bengalensis and common garden

lizard Calotes versicolor.

Among frogs, we noted treefrogs

Polypedates leucomystax and P. maculatus ,

common toad Bufo melanostictus , skipping frog

Euphlyctis cyanophlyctis , Indian bull frog

Hoplobatrachus tigerinus, cricket frog

Limnonectes limnocharis , Boulenger’s frog Rana

alticola , ornate rnicrohylid Microhyla ornata and

red rnicrohylid Microhyla rubra.

There appeared an appreciable change in

the sightings of mammals and reptiles. We saw

only one group of langurs and a pair of Malayan

giant squirrels, and one each of Calotes and

Mabuya. However, these species used to be

common or very common in the 1 970s and early

1980s.

Conservation Issues

We may safely conclude that the WWD

has disappeared from the Pablakhali Wildlife

Sanctuary. Even if a stray pair or two are alive in

some pockets, it is only a matter of time before

these will either be trapped, netted, or hunted down

with guns that have become freely available in

the area.

If we try to find the reasons behind the

disappearance of the WWD, we will arrive at the

reasons for the large-scale destruction of other

biodiversity from the region. The present study

points out to several important but chronic causes.

These are:

Jhum cultivation: This is the slash and bum

process, of clearing a patch of natural forest and

planting paddy and other cash crops, carried out

by the tribals. Such a cleared field area is used

alternately every 3rd or 4th year. The moment a

forest is removed from an area, the topsoil is

washed downhill by the monsoon rain. By

repeated jhum cultivation, the soil loses its fertility

and such fields are soon taken over by exotic

plants or invasive local species of no commercial

value. Such problematic species include the twiner

Mikania scandens, shrubs such as Eupatorium

odoratum , Lantana camara and thatching grass

Imperata arundinacea.

Back in 1875, the then Conservator of

Forests in Bengal, Sir William Schlich, noted that

jhum cultivation was the enemy number one for

the existence of natural forests in CHT District

(based on Forest Department reports). The British,

up to 1947, the East Pakistan authorities up to

1971, and the present Bangladesh authorities have

not been successful in dissuading the tribal people

from practising this slash and bum process of

cultivation in all the three Hill Districts (Plate 1,

Fig. 2).

Clear-felling and monoculture: The British,

followed by the East Pakistan, and the present

Bangladesh Forest Authorities, had and have one

common goal in mind — to send more and more

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DISAPPEARANCE OF WHITE-WINGED DUCK FROM BANGLADESH

revenue to the government exchequer by selling

natural trees and other forest produce. This trend

was consistent in the past and is continuing

unabated to date. To do this they utilised the

simple process of ‘clear-felling’ operations, on a

predetermined basis prescribed in the ‘Working

Plan’ prepared by a forest staff member for a Forest

Division. In this process, all timber and

commercially important species of trees are

removed first, followed by planting of saplings of

a single commercially viable species of timber such

as teak ( Tectona grandis ), jarul ( Lagerstroemia

reginae ), dipterocarp (Dipterocarpus spp.), jamun

( Syzygium cumini), etc., as monocultures. Trees

grown under monoculture are also removed on

the basis of short or long rotations ranging from

20 to 60 years. The planting activities were assisted

by tribal Chakmas and forest-villagers. In

exchange for their labour, they would be allowed

to bum all other plants from the monoculture area

and start cultivating jhum rice and other crops.

The jhumias stayed there for the first two years

to take care of government forest while cultivating

their own crop in the area. They also helped remove

the weeds from the plantation. These processes

resulted in the large-scale removal of natural

forest, encouraged jhum cultivation in reserved

forest area and allowed encroachment of forested

land by the forest villagers (Plate 2, Fig. 3). At the

end of it all, wildlife was wiped out from the area.

Construction of Kaptai Dam: Under a fund

from USAID during 1 959-63, the then East Pakistan

Government constructed the Kaptai Hydroelectric

Project with a view to generating 230 MW power

for the country (now Bangladesh, after December

1971). Unfortunately in 1992, the power

production in the project area went down to an all

time low of 30 MW, due to shortage of water and

filling up of both the lower and upper reaches of

the lake by accumulated silt that resulted from

deforestation in the hilly areas. When fully

commissioned in 1962, the Project inundated an

estimated area of 655 sq. km that included 40% of

the best cultivable land in the former CHT District

(Gain 1995). Due to this project the Kaptai Lake

— an artificial one — has been created. This dam

has displaced at least 20,000 plough and jhum

cultivating Chakmas from the Kassalong Valley.

It destroyed an entire ecosystem from the valley

floor up to about 10 m. This was a one time

colossal loss of biodiversity in CHT.

The Chakmas and some settlers from the

1950s and early ’60s allotment of valley lands,

who were affected by the creation of the Kaptai

Dam in 1 965, were ultimately resettled in a section

of the Kassalong Valley Reserve Forest in the late

1960s. This also destroyed the wildlife and its

habitat.

Negative management of wildlife:

Bangladesh Government ended all its

responsibilities of managing the nation’s wildlife

wealth, by promulgating the Bangladesh Wildlife

Preservation Order/ Act 1973 and appointing a lone

Senior Research Officer under the Forest

Directorate. This person is now retiring in the same

post, in a year or two. The Government has also

declared a few areas as National Parks, Wildlife

Sanctuaries, Game Reserves, Bird Sanctuaries, etc.,

under the above Act, as well as Biosphere

Reserves and a World Heritage Site, under

pressure from western donor countries. But all of

these are only on paper and not in practice. The

Government has utterly failed in managing the

country’s forests and wildlife (Plate 2, Fig. 4).

Settlements: Towards the end of the 1970s,

the Bangladesh Government decided to settle

plains-dwelling and Bengali-speaking people en

masse in the Bandarban, Khagrachari and

Rangamati Hill Districts, which traditionally used

to be the home of the tribal people only. I

understand, over a period of a decade or so, the

Government was able to settle about 25,000

Bengali families in the hilly areas, including

government-reserved forests. Each family was

allotted a plot of 5 acres of hilly land, 4 acres of

mixed land and 2.5 acres of paddy land (Gain 1 995).

These settlers first chopped and sold all valuable

timber trees from their allocated areas and then

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371

DISAPPEARANCE OF WHITE-WINGED DUCK FROM BANGLADESH

moved on to encroach upon more and more

government forests as they had very little place

to practise agriculture. It was an easy way of

getting quick money by selling trees and other

forest produce. Ultimately, they started

encroaching upon both government revenue

(khas) and forested lands. The population of the

settlers must have doubled during the past two

decades or so. Their demand for land and forest

is never ending. Also, these people have negative

social interaction with the tribal people due to

cultural conflicts (Gain 1995).

Tribal insurgency: To counter the

settlement of plains-dwelling Bengali people in

the hilly areas of the erstwhile CHT District, a

section of the tribal Chakmas formed a resistance

group called ‘Shanti Bahinr meaning Peace Force.

This tribal insurgency continued roughly from

1980 to 1998. During the insurgency period, the

Government brought in their counter-insurgency

machinery such as the Military, Para Military BDR

(Bangladesh Rifles), Police, Ansars, VDP (Village

Defence Party), and a number of civil

administrations. It is conjectured that during this

long period, all sides and parties involved in the

conflict destroyed forests and wildlife in an

unabated fashion. The trend continued even after

a peace deal was signed between the Shanti

Bahini and Bangladesh Government in early 1998

when the tribals surrendered their amis. When I

visited the area between February i 8 and 22, 1998,

and during the course of the present study, I

documented the rampant destruction of forests

and wildlife in the Sanctuary.

Presently it seems that nobody is in control

of the CHT forests. It has become a free for all

situation, with Chakmas and other tribals randomly

cutting reserved forests and even government

forests of monoculture just created in the 1990s,

considering it to be their right. Taking advantage

of the prevailing situation, the Bengali settlers

are removing as many forest trees as possible to

meet their daily needs as well as in greed. All law-

enforcing authorities are apparently sitting idle.

Possibly, they do not like to get involved in any

conflict. Overall, the social situation seems to be

tense between the tribal Chakmas and Bengali

settlers, as there is no free exchange between the

two communities. Meanwhile, forest destruction

and kill mg/capturing of wildlife are continuing

unhindered.

Recommendations

a. First of all, parties occupying the area must

restore peace and tranquillity in the area without

which there can be no development for either

the Bengali-speaking settlers or the ethnic

tribal people.

b. Specifically for the WWD: If any duck of this

species is captured it must be saved. The way

to do this is to buy back the duck from the

tribal Chakmas or Bengali settlers by bartering

it for domesticated ducks. People who live in

the area should be given material incentives

to protect the WWD in their area.

Domesticated ducks and poultry have good

cash value and a certain number of these can

be given to those families who live in the WWD

area. This will allow the unhindered existence

of the species.

However, there must be a Government or NGO

unit to facilitate these barter deals and oversee

the project, from inception to implementation.

This unit should ensure that sufficient funds

are available to rehabilitate any ducklings

produced. A certain number of ducklings can

even be taken away to a few captive-breeding

centres within the tribal area or outside it, with

a view to bringing back the grown birds for

reintroduction into the former range within the

Sanctuary or its neighbourhood.

c. Jhum cultivation should be restricted only to

areas where jhum fields already exist. Further

denudation of forest must be stopped, as tribals

themselves are not likely to get any land in the

future to practise jhum cultivation if the current

trend continues.

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JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG. -DEC. 2003

DISAPPEARANCE OF WHITE-WINGED DUCK FROM BANGLADESH

d. No new monoculture should be allowed.

Instead, mixed species of indigenous trees

should be planted and nurtured, with emphasis

on fruit bearing and softwood trees that allow

animals to get food and nesting opportunities.

At least one third of the existing reserved forest

should be declared as a Nature Reserve where

forestry, agriculture or settlement activities of

tribals or plains-dwellers should be stopped

for at least half a century. This will allow the

forest to regenerate on its own, and the

biodiversity can get a foothold and revive to a

certain extent.

e. A separate Wildlife or Biodiversity Bureau

should be created under the Environment and

Forest Ministry. All lands declared previously

as National Parks, Wildlife Sanctuaries, Game

Reserves, Nature Reserves, World Heritage

Site, Biosphere Reserves, etc., and to be so

declared in the future, must be handed over to

this new Bureau for their total management.

These areas will have no administrative tie with

the Environment or the Forest Department.

This organisation is also to be banned from

altering existing habitats for commercial

purposes. Instead it must aim at scientific and

sustainable management of biodiversity. It

must try to practise sustainable utilisation of

the natural resources.

f. All zoos, wildlife research institutes and

captive breeding centres existing in the

country must be incorporated under this

Wildlife Bureau.

g. Top-level managers of the Bureau must be

recruited at a national level, and have a wildlife

background. Professionals from other

disciplines such as Botany, Geology, Soil

Science, etc., should also be involved in the

activities of the Bureau. The rest of the

manpower can be recruited locally. Only trained

local people should be involved in the field

level activities in all wildlife/biodiversity areas.

No foreigners should be incorporated in the

activities of the Bureau. However, foreign

advisors can help the Bureau in future planning

and project designing.

h. The Bureau would be responsible for the

development of wildlife/biodiversity curricula

for all levels of education in the country.

L It should encourage ecotourism in all managed

areas.

j. It must enrich spoilt habitats by planting suitable

indigenous tree species, especially those with

soft, fleshy fruits and colourful, nectar producing

flowers, various fig species and softwood trees

suitable for hole-nesting animals.

k. The Bureau must be active in raising public

awareness campaigns. It must popularise

Biodiversity Conservation in the country and

take part in all activities of international bodies

like the CITES Authority, HJCN - the

Conservation Union, WWF (Worldwide Fund

for Nature), and WAZA (World Association

of Zoos and Aquariums).

Acknowledgements

I am grateful to the Bombay Natural History

Society and the Honorary Secretary, Mr. J.C. Daniel

for inviting me to submit this paper. I thank the

Emirates Natural History Group, Abu Dhabi, UAE,

and the Forum of Environmental Journalists of

Bangladesh and Centre for Sustainable

Development (CFSD), Dhaka, Bangladesh for

sponsoring portions of my present study.

Mohammad Mohoshinuzzan Chowdhury, a

researcher from Dhaka was my constant

companion for the entire fieldwork period.

Conservator of Forests, Rangamati, Mr. Altaf

Hossain, and DFO of CHT North Forest Division,

Mr. Mamunur Rahman Chowdhury, provided me

with necessary permission and local hospitality. I

wish to thank them all. I am also thankful to Messrs

Mahftiz Ullah of CFSD, Qamrul Islam, Chairperson

of the FEJB, Shehabuddin Ahmed of the Daily

Star, Mofizur Rahman of the Daily Janakantha,

Chief Conservator of Forests, Mr. Mohammad

Nuruzzaman, all the Foresters and many others

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, J00(2&3), AUG.-DEC. 2003

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DISAPPEARANCE OF WHITE-WINGED DUCK FROM BANGLADESH

whom I wish to thank personally, who provided

me valuable information regarding status of

biodiversity and habitat destruction of the area

covered by this survey. Dr. Baz Hughes, Chairman,

Threatened Waterfowl Specialist Group of IUCN/

SSC, WWT, Slimbridge, U.K. gave me the latest

information on the WWD. I am grateful to all of

them for their generosity.

References

Gain, P. (Ed.) (1995): Bangladesh: Land, Forestand Forest

People. Dhaka: Society for Environment and

Human Development, xii + 163 pp.

Green, A.J. (1993): The biology of the White-winged

Duck Cairina scutulata. Forktail 8: 65-82.

Husain, K.Z. (1977): The White-winged Wood Duck.

Tigerpaper (4): 6-8.

Husain, K.Z. (1985): Present status of the birds of

Bangladesh. Proceedings, the Third East Asian

Bird Protection Conference, Tokyo, Japan,

29-31 May, 1985. Wild Bird Society of Japan,

Tokyo. Pp. 18-23.

Husain, K.Z. & M.N. Haque (1982): The White-winged

Wood Duck Project: Report submitted to the

University Grants Commission, Dhaka. Unpubl.

Iucn Bangladesh (2000): Red List of Threatened Animals

of Bangladesh. IUCN-The World Conservation

Union, xii + 54 pp.

Khan, M.A.R. (1981): The Endangered Birds of

Bangladesh. Newsletter for Birdwatchers 21(12):

4-7.

Khan, M.A.R. (1982): Wildlife of Bangladesh-

A Checklist. Dhaka University, Bangladesh.

Khan, M.A.R. ( 1 983): The vanishing White-winged Wood

Duck. Bangladesh Today , 16-30 June: 43-45.

Khan, M.A.R. (1986): The threatened White-winged

Wood Duck Cairina scutulata in Bangladesh.

Forktail 2: 97-101.

374

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003

THE INCUBATION MOUND AND HATCHING SUCCESS

OF THE NICOBAR MEGAPODE MEGAPODIUS NICOBARIENSIS BLYTH

( With four text-figures and one plate)

K. SlVAKUMAR1 AND R. SANKARAN2

Key words: Incubation mounds, Nicobar megapode, hatching success, megapodes,

Megapodius nicobariensis

Incubation mounds of the Nicobar megapode Megapodius nicobariensis were constructed with

sand or sand with plant materials, such as leaves, twigs and bits of decomposing wood. Usually, the

site selected for a new incubation mound was a fallen log, tree stump, or the decomposing roots of

a tree. Mound construction begins with the birds either digging a pit at the site or by covering the

decomposing log or tree stump with soil and litter. The mean egg-laying interval was 14.91 ±1 .43

days, the average number of eggs laid in a mound was 4.5 ±0.6 eggs, but it significantly varied

between the years and the average clutch size of the Nicobar megapode was 2.75 ±0.35 eggs.

Moderate rainfall in the dry season enhances egg production. Microbial activity appears to be the

primary source of heat within mounds. The size of the mound was positively correlated to the

temperature within the mound. The average incubation temperature was 32.44 ±0.21 °C and the

average incubation period was 74.73 ±0.52 days. There was, however, no significant relationship

between the mound size and hatching success of the Nicobar megapode.

Introduction

Megapodes are unique among birds

because they incubate their eggs in mounds of

rotting leaves or in burrows in geothermally heated

ground (Frith 1 956, Dekker and Wattel 1 987, Jones

1988, Dekker 1990). Perhaps the best-studied

aspects of the Megapodiidae are the incubation

conditions within mounds and communal nesting

grounds (Frith 1956, 1 959, Crome and Brown 1979,

Seymour et al. 1 986, Booth 1 987, Seymour et al.

1987, Dekker 1988, Jones 1988).

Within the Megapodiidae, there exist two

groups, burrow nesters and mound builders with

variations in the incubation and breeding

strategies. Burrow nesting species like

Macrocephalon and Eulipoa lay eggs at

communal nesting grounds where sun or volcanic

activity provides heat for incubation (Dekker 1988,

uS£lim Ali Centre for Ornithology and Natural History

(SACON), Coimbatore 641 108, Tamil Nadu, India.

'Present Address: Wildlife Institute of India,

P.O. Box 18, Chandrabani, Dehra Dun 248 001,

Uttaranchal, India.

Email: [email protected]

1990, Heij et al. 1997). Talegalla, Aephypodius,

Alectura and Leipoa build incubation mounds of

forest litter where organic decomposition provides

necessary heat (Dekker 1 990). Of the 1 3 species

that comprise the genus Megapodius , 10 are

mound builders, two are burrow nesters, and one

nests in both burrow and mound (Jones et al.

1995). The Nicobar megapode Megapodius

nicobariensis , a monomorphic mound building

megapode (Plate 1, Figs 1,2), endemic to the

Nicobar Islands in the Bay of Bengal, builds

incubation mounds of sand, loam, coral bits and

rotting vegetation, within which eggs are laid.

Incubation mounds of the Nicobar megapode vary

in type, size and location (Dekker 1 992, Sankaran

1995, Sankaran and Sivakumar 1999, Sivakumar

and Sankaran, in press). Some mounds have a

greater admixture of vegetative material, while

others have a greater amount of sunlight falling

on them, which suggests that the source of heat

varies between mounds. In this paper, we address

the questions consequently raised: which sources

of heat provide the most stable incubation

conditions? Do sources of heat that create suitable

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003

375

INCUBATION AND HATCHING IN MEGAPODIUS NICOBARIENSIS

incubation conditions within mounds vary with

mound type, location and dimensions? And do

heat sources and mound dimensions have a

bearing on the number of pairs that use a mound,

the number of eggs laid, and hatching success?

Study Area

The Andaman and Nicobar Islands (6° 45'

to 13°41' N, 92° 12' to 93° 57' E), in the Bay of

Bengal, arch from Arakan Yoma, Myanmar in the

north to Sumatra, Indonesia in the south (Saldanha

1 989; Fig. 1 ). These islands cover an area of 8,249

sq. km with a coastline of 1 ,962 km. The Andaman

group with more than 325 islands (21 inhabited)

covers an area of 6,408 sq. km. The Nicobar group

with over 24 islands ( 1 3 inhabited) covers an area

of 1 ,84 1 sq. km (Singh 1981, Saldanha 1 989).

We studied the ecology of the Nicobar

megapode between December 1 995 and July 1996,

December 1996 and June 1997, September and

October 1997, and February and May 1998. The

study period includes three dry seasons (peak

period of egg laying) and part of one wet season.

Our study area was on the coast at the southern

tip of Great Nicobar Island. The intensive study

area was a narrow strip of forest, of width varying

between 40 and 300 m and length about 4 km,

which was bisected by a disused metalled road,

ending at the light house at Indira Point. The

beach forms the boundary to the study area in

the east, and wetlands or forests that are

inundated during the monsoons form the

boundary to the west. The soil within this strip

of coastal forest was sandy and loamy, and

the dominant trees were Barringtonia asiatica ,

Fig. 1 : Southern part of the Great Nicobar Island showing the study site

376

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003

INCUBATION AND HATCHING IN MEGAPODIUS NICOBARXENSIS

Sivakumar, K. and R. Sankaran: Nicobar megapode Megapodius nicobariensis Plate 1

Fig. 1 : Nicobar megapode on the mound it has built

Fig. 2: Another incubation mound of the Nicobar megapode

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003

377

PICS: RAVI SANKARAN

INCUBATION AND HATCHING IN MEGAPODIUS NICOBARIENSIS

B. racemosa, Terminalia bialata , Terminalia

catappa , Syzygium samarangense , Thespesia

populnea and Macaranga spp. The study area

had dense stands of Pandanus tectorius and

P. odoratissimus in patches, and the road was

fringed by stands of Lea angulata , L. grandifolia ,

and Dracaena spp. There were a few patches of

open ground with little vegetation. The forest

forming the boundary of the study area to the

west had wet clayey soil and was covered with

Areca spp. as well as trees like Ixora barbata ,

Pongamia pinnata , Alstonia kurzii, Adentania

paranina , Aisandra butyracea , Horsfieldia irya ,

Myristica andamanica , and Celt is timorensis.

Methods

At the start of the study or whenever a new

mound was constructed, detailed drawings of the

mounds were made to scale using measuring tapes

and a compass, and salient characteristics such

as living trees and dead logs or tree stumps were

plotted on these. The basal circumference, height

and diameter of the mounds were measured once

a month. Mounds were uneven in shape with a

cone- like appearance. The mound size, expressed

as volume, was derived from the equation for the

volume of a cone: 1 /3 Tcr2h where ‘r’ is the radius

and ‘h’ the height, giving the approximate volume

of the mound.

All the incubation mounds in the study

area were visited at least twice a day, in the

morning and in the evening and occasionally

during midday, to identify whether megapodes

had worked on the mounds, and what type of

activity they had engaged in. At four mounds,

the megapodes were intensively observed,

following the focal animal sampling method

(Altmann 1974), from observation hides.

Observations from the hide usually started

before the arrival of birds at the mound (at 0500

to 0530 hrs) and ended after the birds left the

mound. Observation recommenced at about 1400

hrs and was carried on till dusk. The activities of

the birds were classified into: visit, pit digging,

egg laying, raking, covering, pits-filled, and

random activity.

In 1996, four temperature probes were

implanted at depths between 20 and 75 cm, in

seven mounds that had been selected for intensive

studies. However, after about two months these

probes malfunctioned, probably due to high

humidity and rainfall. In 1997 and 1998, a

temperature probe placed at the tip of a one metre

long steel tube was inserted to depths of 30, 60

and 90 cm to measure the temperature. Using this

method the temperatures were measured once a

month for all the mounds in the study area, every

1 5 days for the target mounds, both in the morning

and evening. Occasionally the temperature of the

mound was also measured during midday.

Microbial activity was measured using a

soil respirometer (PP Systems EGM-1

Environmental Gas Monitor with a SRC-1 Soil

Respiration System), assuming that in those

mounds where microbial activity was high, greater

amounts of C02 would be emitted. The soil

respirometer measures the C02 change in a fixed

volume over a known time and fits a quadratic

equation to the data to arrive at a value ‘SR5 which

is the soil respiration rate in gC02/m3/hr. Soil

respiration was measured once every 1 0 or 1 5 days

for the seven mounds that were under intensive

study. Like temperature, data on the soil

respiration of mounds was collected both in the

morning and evening.

The intensity of light falling on the mound

at different times of the day was measured using

a luxmeter. The amount of Photosynthetic Active

Radiation (PAR) falling upon the mound and PAR

absorbed by the mound were measured using

Sunfleck Ceptometer (Decagon, Pullman, WA).

This was also measured outside the mound. The

gap in the canopy cover above the mound was

measured using a concave mirror that was

uniformly graduated.

Soil samples were collected from the surface

of the mound and then sun dried for an hour.

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INCUBATION AND HATCHING IN MEGAPODIUS NICOBARIENSIS

Percentage of humidity was measured by using

the following formula:

Humidity (%) = [(Wet soil weight - Sun dried

soil weight) / Wet soil weight] x 1 00

During the breeding seasons of 1 996, 1 997

and 1998, thirty-seven mounds were monitored.

When an egg was laid, it was dug out and weighed

to the nearest gram using a spring balance. Eggs

were also measured with Vernier callipers and

marked with a number and date using an HB

graphite pencil. After weighing and marking, the

eggs were reburied in the same egg chamber and

the mound was rebuilt. For identification, a stick

was placed adjacent to the egg chamber. During

1997 and 1998, thirty-four eggs were directly

marked and monitored. The marked eggs were

monitored by rechecking them once every 15

days. At the beginning of 1998, all the mounds

were thoroughly checked with the help of mound

maps of 1997, where the locations of eggs were

clearly plotted. Successful hatching of eggs was

evident from eggshell fragments and pieces of

shell membrane where the eggs had been.

Emergence of the chick from the egg during

hatching and its subsequent activity was

observed by placing a glass plate adjacent to the

egg on the day of egg laying. In 1998, a total of

seven eggs were monitored for the same. Eggs

which remained in the mounds for the entire

breeding season, or those which did not hatch

for 100 days were opened and examined.

Results

Description of incubation mound of the Nicobar

megapode

Of the 38 incubation mounds present in the

study area between 1996 and 1998 (Table 1), three

were type ‘A’ incubation mounds built on an open

spot away from trees, three were type ‘B’

incubation mounds built against a large living tree,

and 25 were type ‘C’ incubation mounds built on

or around a dead log or stump. Of the remainder,

four incubation mounds were type ‘BC’ (built

against the buttress of a partially living tree, or

with a dead log in it), of which two later became

type ‘C’ in 1998 because the trees died out

completely. One incubation mound was type ‘AB’

as it was built in the open with two living

Pandanus palms in it (Table 1). Two incubation

mounds were unusual, as they were built against

the edge of the disused metalled road that bisected

the study area (type ‘R’). Among the 16 incubation

mounds that were newly constructed during the

study period, 13 were type ‘C’ incubation mounds,

one was a type ‘B’ incubation mound, and two

were built against the road (Table 1).

The construction and maintenance of an

incubation mound involves several activities.

Usually, the site selected for a new incubation

mound is a fallen log, tree stump, or the

decomposing roots of a tree. The megapodes

began construction of the new incubation

mound either by digging a pit if the site was over

decomposed roots or by covering the

decomposing log with soil and litter raked in from

the surrounding areas. The process of raking soil

and litter on to the site, or the piling up of soil and

other material over the pits, soon resulted in the

formation of a new incubation mound.

Temperature and soil respiration in and out of

the incubation mound

Mean core temperature at the depth of 50-60

cm in an incubation mound was 31.94 °C

(SE 0.075, n=634). It was higher than the forest

ground where it was 28.72 °C (SE 1.66, n==105)

at the same depth. The mean soil respiration rate

(SR in gC02/m3/hr) on the active incubation mound

was 5.55 (SE 0. 1 3, n=920), which was always higher

than that on the abandoned incubation mound (SR=

2.88, SE 0.92, n=140) as well as the normal ground

(SR = 3 .7, SE 0.86, n= 1 30).

Incubation period and optimal incubation

temperature

Incubation temperature (mound core

temperature) of the 34 egg chambers in 1 6 different

380

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INCUBATION AND HATCHING IN MEGAPODIUS NICOBARIENSIS

Table 1: The history of incubation mounds of the Nicobar megapode in the study area

NC= New construction; PS= Present at start of study, NF= Newly found.

^s mounds change in size overtime, the mean value for all mound size data collected during the study is given.

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381

INCUBATION AND HATCHING IN MEGAPODIUS NICOBARIENSIS

mounds was monitored, and the incubation period

for 30 eggs determined. The remaining four eggs

did not hatch. The shortest incubation periods

were 70 days (n=l egg) and 72 days (n=6 eggs)

and the longest incubation period was 81 days

(n=l egg). The mean temperature of the egg

chamber for successful hatching was 32.44 ±0.2 1

°C (n=30). The mean incubation period of the

monitored eggs was 74.73 ±0.52 days. Though

the data (Fig. 2) indicates that as temperature

decreased the incubation period increased, there

was no significant negative correlation between

the length of incubation period and incubation

temperature (r = -0.3 1 , n=30, p=0.095). Moreover,

as eggs incubated at different temperatures

hatched in almost the same period, it also indicates

that minor fluctuation in the temperature of the

egg-chamber did not affect the incubation period

significantly.

Effects of incubation mound size on incubation

temperature

The effect of mound size on the incubation

temperature in 37 incubation mounds was studied.

The sizes of the 37 incubation mounds varied from

0. 1 5 cu. m to 40.24 cu. m, with a mean size of 4.78

cu. rn (SE 1.19). As mound size increased, the

Table 2: Average temperature (in °C) of the mounds

at various depths

Ambient Surface Upper Middle Deep Deepest

layer layer layer layer

Mean 28.17 27.57 29.91 30.65 31.94 32.51

n 735 745 196 618 634 628

SE 0.065 0.055 0.177 0.074 0.075 0.072

temperature of the mound also increased (Fig. 3 )

at the depth of 30 cm (r =0.162, n=518, p<0.001),

60 cm (r =0.177, n=532, p<0.001) and 90 cm (r

=0.307, n=526, p<0.001). Within a mound there

was some fluctuation in the incubation

temperature, irrespective of sizes (Fig. 3).

Role of sunlight in incubation temperatures

Intensity of the light (lux value)

significantly enhanced the ambient temperature

(r =0.24, n= 1 68, p<0.0 1 ) and surface temperatures

of the mound (r =0.25, n=168, p<0.01)but not that

of the mound core (r =0.053, n=96, p=0.610).

However, there was a positive correlation

between the surface and the mound core

temperature (r =0.23, n=626, p<0.00 1). The mean

ambient and surface temperatures were always

lower than the mound temperatures at different

depths (Table 2),

70 72 73 74 76 78 79 80 81 ?

Incubation period (days)

Fig. 2: The relation between the incubation temperature (°C) and incubation period (days) of egg of the

Nicobar megapode (Standard error of the mean shown as error bar, ‘?’ are unhatched eggs)

382

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG. -DEC. 2003

INCUBATION AND HATCHING IN MEGAPODIUS NICOBARIENSIS

O

o

<D

t—

3

-s=»

CO

u-

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a.

E

0

C

o

"*e-»

CO

3

O

c

CDlOCOCO COCDOOCSi '^f'r-COCDCOh-^OO^

(M(O0lflON®qNT*O)CNjO)r;rOq{Vj

oooooooc\ic\ic6cocDCDcda>cvio

V” xf

Mound size (cu.m)

Fig. 3: Relation between mound size and incubation temperature of the Nicobar megapode (Standard error of

the mean temperature shown as error bar)

Role of microbial activity in mound incubation

temperatures

There was a significant positive correlation

between soil respiration rate and the incubation

temperature of the mound (r =0.15, n=204, p=0.02;

Fig. 4).

Effect of moisture content of the mound on

incubation temperature

Moisture content of the mound soil

highly influenced the intensity of the mound

temperature. An increase in the moisture content

of the soil resulted in an increase in mound

temperature at the different depths studied,

as follows: 30 cm (r =0.272, n= 1 66, p<0.00 1 ), 60 cm

(r =0 407, n=166, p<0.001) and 90 cm depth

(r =0.534, n=166, p<0.001). We did not estimate 1

the soil respiration rate when the moisture

content of the soil was estimated, as a

result of which the influence of moisture on the

microbial activity of the soil could not be

established.

O

o

0

i_

3

•#-»

CO

u

0

Q.

E

0

c

q

CO

.Q

3

O

c

34.00

33.50

33.00

32.50

32.00

31.50

31.00

30.50

3.4 4.3 5.3 5.5 5.7 5.9 6.5 7.3 7.9 8.8

Soil respiration rate (gC02/m3/h)

Fig. 4: Effects of microbial activities on the incubation temperature in the mounds of the Nicobar megapode

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003

383

INCUBATION AND HATCHING IN MEGAPODIUS NICOBARIENSIS

Egg-laying behaviour of the Nicobar megapode

The egg-laying interval between two

consecutive eggs and the clutch size of the

Nicobar megapode was estimated from 28 colour-

marked birds. The mean egg-laying interval

between two consecutive eggs of the Nicobar

megapode was 14.91 ±1.43 days (n=l 1 intervals

in 17 eggs). The average number of eggs laid in a

mound was 4.5 ±0.6 (n=58) but it significantly

varied between the years (Kruskal Wallis H test,

c2 =8.203, df=2, p=0.017). Clutch sizes of the

Nicobar megapode varied between one to four

eggs per season or year. We collected data on the

clutch size of seven colour-marked pairs in 1997

and five in 1 998. Of the twelve colour-marked pairs,

five pairs laid four eggs per year in one or two

mounds and remaining pairs laid two to three eggs

in one or two mounds. In general, the average

clutch size of the Nicobar megapode was 2.75 eggs

(SE0.35,n=12).

The peak period of the egg-laying was

between February and May, during which 86.6%

and 84.7% of the eggs were laid in 1996 and 1997

respectively. Egg-laying was not observed during

the wet season of our study (September and

October of 1 997). The total number of eggs laid in

all the mounds of the study area in the year

1996, 1997, and 1998, were 112, 124, and 35,

respectively.

Incubation mound size and egg-laying

Of the 38 incubation mounds that were

active in the study area in 1996, 1997, or 1998,

eggs were laid in 35 mounds. Of these, only 10

incubation mounds were used in all the dry

seasons between 1996 and 1998 for egg laying.

According to the sizes, mounds were

grouped into the four categories, namely very

small, small, medium and large mounds. The very

small sized incubation mounds (<1 cu. m) had the

least number of eggs laid in them (Table 3). Small

sized incubation mounds (1-5 cu. m) had an

average of 4.05 eggs laid in them, medium sized

incubation mounds (5-10 cu. m) contained an

Table 3: The mean number of eggs laid in different

size classes of mounds throughout the study period

average of 5.93 eggs, while large sized incubation

mounds had the most number of eggs (Table 3).

Hatching success

Hatching success was determined in 32

incubation mounds in 1 997, where one to five eggs

were laid in 1 3 incubation mounds, six to ten eggs

in 6 incubation mounds, and more than ten eggs

in 4 incubation mounds. Five of the incubation

mounds were not used for egg-laying in 1997,

and the number of eggs laid in the remaining four

incubation mounds could not be determined.

Mean hatching success in the incubation mounds

in 1997 was 57.26%. Of the 124 eggs laid in 23

mounds, 10.48% of eggs did not hatch and those

eggs were unearthed in the next season; 29.84%

eggs disappeared or were predated, and the fate

of 2.42% of eggs was not clear (if these eggs

successfully hatched then the hatching success

was 59.68%).

Small incubation mounds had less number

of eggs, while medium sized incubation mounds

were used by the birds for more egg-laying

(Table 4). However, there was no relationship

between the incubation mound size and hatching

success (Table 4).

Table 4: Hatching success of eggs in different sizes

of incubation mound of the Nicobar megapode in 1997

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INCUBATION AND HATCHING IN MEGAPODIUS NICOBARIENSIS

Discussion

Incubation conditions within incubation mounds

Incubation mounds of the megapodes are

amongst the largest structures made by any non-

colonial animal, and represent the harnessing of

the energy produced by microbial respiration

(Seymour et al. 1986, Jones 1989), and/or solar

radiation (Frith 1956, 1959) by concentrating

suitable material to provide optimal incubation

conditions at about 33-34 °C (Dekker 1992). In

some species, microbial respiration and solar

radiation may be used sequentially to create

incubation conditions (Frith 1956, 1959). Seymour

(1985) proposed that heat production and heat

loss tends to stabilise mound temperatures at an

equilibrium state due to the great thermal inertia

of mounds once they cross a certain size and as

they maintain adequate moisture content with the

regular incorporation of fresh organic material into

the mound. This model has gained further support

from other studies (Jones 1988, Jones and Birks

1992).

Mound temperatures usually stabilise

between 32-35 °C (Jones et al. 1995), which is

consistent with that of the Nicobar megapode

(32.44 °C). Incubation temperatures in mounds

show considerable fluctuation, and while the

negative effects of these fluctuations on eggs are

largely offset by a variable incubation period (Booth

1987), there are strategies to balance both heat

loss and gain (Jones 1989, Jones and Birks 1992).

However, the data from the incubation

mound of the Nicobar megapode does not fully

fit with the assumptions mentioned above by

Seymour (1 985). Firstly, the size of the incubation

mound can vary in height from 10 cm up to 2. 1 m

and in basal circumference from 7 to 45 m

(Sankaran 1995). Secondly, the proportion of

organic material in an incubation mound varies

due to location of the mound, and the availability

of materials around it. And thirdly, the gap in the

canopy above the incubation mound varies,

resulting in differences in the amount and duration

of sunlight falling on it. This might indicate that

the heat sources which create suitable incubation

conditions within the mound may vary, with some

incubation mounds appearing to rely more on

sunlight and others on organic decomposition

(Sankaran and Sivakumar 1 999).

Solar energy, however, probably only

optimises the incubation mound temperatures.

Though the canopy above an incubation mound

was less than the canopy above non-mound areas,

direct sunlight fell on the mounds for very short

periods, with the result that both ambient

temperature and incubation mound surface

temperatures were always lower than incubation

mound core temperatures. Thus, the role of solar

energy appears to be restricted to warming the

surface of the incubation mound, whereby

dissipation of heat was reduced.

Microbial respiration is the primary source

of heat harnessed by most mound building

megapodes (Jones et al. 1995). A clear relation

exists between the incubation temperatures and

the organic activity as evidenced from soil

respiration in the Nicobar megapode as well.

Microbial respiration has a linear relation to the

temperature of the incubation mound at deeper

layers. However, incubation mounds with higher

levels of soil respiration did not necessarily have

higher temperatures. Two factors could be

responsible for this. Firstly, there might be

differences in the amount of heat produced by

decomposition due to the kind of vegetative

materials added to the incubation mound.

Secondly, the rate of heat loss probably differs

between incubation mounds, caused by

differences in the proportion of surface area to the

volume of the incubation mound, or to the amount

of moisture content within the incubation mound

(Jones et al. 1995), or the amount of sunlight or

radiation from the beach falling on the mound.

Mound size, egg-laying and hatching success

The optimisation of incubation conditions

in large incubation mounds is reflected in an

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385

INCUBATION AND HATCHING IN MEGAPODIUS NICOBARIENSIS

overall trend of a greater number of pairs using

such mounds, and consequently, a greater number

of eggs being laid in them. However, some small

mounds had a greater number of eggs, and some

larger mounds had fewer eggs, indicating that size

is not the only criterion. The quality of the

incubation mound, and the number of pairs using

an incubation mound, which appears to be

somewhat independent of incubation mound size,

are probably other determining factors.

As optimal temperature was consistently

present in large mounds, one would expect that

eggs in large incubation mounds are more likely

to hatch successfully than in smaller incubation

mounds. However, hatching success of the

Nicobar megapode does not reflect this trend. This

study reveals that there was no significant

relationship between the incubation mound size

and hatching success. Very small and medium

sized incubation mounds showed more hatching

success than the small ( 1 -5 cu. m) sized ones. The

probable reason for the lower hatching success

in the small sized incubation mounds (1-5 cu. m;

Table 4) was the large number of birds that used

them. Greater digging activity and consequently

greater exposure of eggs to the atmosphere was a

possible factor for lower hatching success.

Secondly, more mound activities may attract more

predators, especially monitor lizards. About 30%

of megapode eggs were predated in 1997, when

activities at the mound were also the most. Clutch

size of the Nicobar megapode was lower than other

mound building megapodes (Jones et al. 1995).

Acknowledgements

The Ministry of Environment and Forests,

Government of India, sponsored this study and

the logistic support by the Forest Department,

Andaman and Nicobar islands is gratefully

acknowledged. We thank the World Pheasant

Association for helping to fund the soil

respirometer. We thank Rene Dekker, Darryl Jones,

Lalitha Vijayan, Ajith Kumar, P.A. Aziz, H.S. Das,

N.K. Ramachandran, A. Rajasekaran, B.P. Yadav,

A.K. Biswal, R. Rajyashri, Areendam, and David

for their inputs in this study, and Jugulu Maheto

and Prem Ram who assisted us in the field. We

also thank all the anonymous reviewers for their

valuable inputs in this paper.

References

Altmann, J.C. (1974): Observational study of behaviour-

sampling method. Behaviour 49: 227-285.

Booth, D.T. ( 1 987): Home range and hatching success of

Mallee fowl Leipoa ocellata Gould (Megapodiidae),

in Murray mallee near Renmark. S, A. Australian

Wildlife Research 14: 95-104.

Crome, F.H.J. & H.E. Brown (1979): Notes on social

organisation and breeding of the Orange-footed

Scrub fowl Megapodius reinwardt . Emu 79: 111-

119.

Dekker, R.W.R.J. (1988): Notes on ground temperatures

at nesting sites of the Maleo Macrocephalon

maleo (Megapodiidae,!. Emu 88: 124-127.

Dekker, R.W.R.J. (1990): Conservation and biology of

megapodes (Megapodiidae, Galliformes, Aves).

Unpublished D.Phil. thesis. University of

Amsterdam.

Dekker, R.W.R.J. (1992): Status and breeding biology of

the Nicobar megapode Megapodius nicobariensis

abbotti on Great Nicobar, India. Report, National

Museum of Natural History, Leiden.

Dekker, R.W.R.J. & J. Wattel (1987): Egg and image:

new and traditional uses for the Maleo

( Macrocephalon maleo). In: The value of birds

(Eds.: Diamond, A.W. & F.L. Filion). ICBP

Technical Publication 6.

Frith, H.J. (1956) Breeding habits in the family

Megapodiidae. Ibis 98: 620-640.

Frith, H.J. (1959): Breeding of the Mallee Fowl, Leipoa

ocellata Gould (Megapodiidae). CSIRO Wildlife

Research 4: 31-60.

Heij, C.J., C.F.E. Rompas & C.W. Moeliker (1997):

DEINSEA 3: The biology of the Molluccan

megapode Eulipoa wallacei (Aves, Galliformes.

Megapodiidae) on Haruku and other Molluccan

Islands. Final report, Part 2.

Jones, D.N. (1988): Construction and maintenance of

the incubation mounds of the Australian Brush-

386

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INCUBATION AND HATCHING IN MEGAPODIUS NICOBARIENSIS

turkey A/ectura lathami. Emu 88: 210-218.

Jones, D.N. (1989): Modern Megapode Research. A post-

Frith review. Corella IS: 145-154.

Jones, D.N. & S. Birks (1992): Megapodes: recent ideas

on origins, adaptations and reproduction. Trends

in Ecology and Evolution 7(3): 88-91.

Jones, D.N., R.W.R.J. Dekker & C.S. Roselaar (1995):

The Megapodes. Oxford University Press. 262 pp.

Saldanha, C.J. (1989): Andaman, Nicobar &

Lakshadweep: An environmental impact

assessment. Oxford & IBH Publ. Co., New Delhi.

Sankaran, R. (1995): The Nicobar Megapode and other

endemic Avifauna of the Nicobar Islands (Status

and conservation). SACON Technical Report 2.

Sankaran, R. & K. Sivakumar (1999): Preliminary results

of an ongoing study of the Nicobar Megapode

Megapodius nicobariensis Blyth. Zoologische

Verhandelingen 327: 75-90.

Seymour, R.S. ( 1 985). Physiology of megapode eggs and

incubation mounds. Acta XVIII Congressus

Internationalis Ornithologici 2: 854-863.

Seymour, R.S., D. Vleck, & C.M. Vleck (1986): Gas

exchange in the incubation mounds of megapode

birds. J. Comparative Physiology 156(B): 773-

782.

Seymour, R.S., D. Vleck, C.M. Vleck & D.T. Booth

(1987): Water relations of buried eggs of mound

building birds. J. Comparative Physiology 157(B):

413-422.

Singh, B.K. (1981): Census of India 1981. Series-24.

Andaman and Nicobar Islands. Govt, of India.

Sivakumar, K & R. Sankaran (in press): Incubation

mounds and mound use patterns in the Nicobar

Megapode. Abstract in: Proceedings of the 1st

Pan-Asian Ornithological Congress and XIII

Birdlife Asia Conference, Coimbatore, India.

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003

387

THE SONG OF NIN OX SCUTULATA OBSCURA

( With one text-figure)

Ben King1

Key words: Ninox scutulata , Ninox affinis , sonograms, Andaman and Nicobar Islands

There has been some controversy over the relationship of the two species of Ninox owls occurring

in the Andaman Islands. Sonograms of the calls

relationships.

As Ninox scutulata obscura Hume 1 873 of

the Andaman and Nicobar Islands is almost

entirely dark in colour, there has been some

uncertainty about its relationship with mainland

Ninox scutulata. Further complicating the picture

is the presence of Ninox affinis Beavan 1 867 on

the Andaman and Nicobar Islands. Superficially,

N. affinis resembles mainland N. scutulata much

more than obscura does.

Ali and Ripley (1981) do not mention the

song of obscura in their account of that form.

However, in their account of N. affinis , A.L. Butler

is quoted as saying that the call of affinis is “a

loud craw, something like a Glaucidium’ s note

of positively identified individuals clarify these

and quite different from that of N. scutulata in

Ceylon (=Sri Lanka), which is a flutey disyllable,

as is also that of obscura ,” thus offering an

accurate description of the calls of both species.

Abdulali (1964) also correctly noted the song of

obscura as a “loud dysyllabic coo-ook .”

In March 1991, 1 had the opportunity to

visit North Andaman Island and was able to tape

record both N. scutulata obscura and N. affinis.

Sonograms of both are presented in Fig. 1. The

left diagrams show frequency plotted with time,

showing the upward inflected double note of

N. scutulata and the downward inflected single

note of affinis. The right side of the figure is a

N

X

0

1

t

energy

I

t

energy

t

energy

1

t

energy

0

seconds

1

Fig. 1 : Song of Ninox scutulata obscura of the Andamans compared with songs of N.s. hirsnta from Kerala

and Sri Lanka, and with the song of N. affinis from the Andamans

'Ornithology Department, American Museum of Natural History, Central Park West at 79th St.,

New York, NY 10024 USA. Email: [email protected]

388

JOURNAL , BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003

SONG OF NINOX SCUTULATA OBSCURA

wave form sonogram showing the amplitude of

sound energy emanating from a central axis. The

double note of scutulata is distinct in hirsuta

from Kerala and Sri Lanka, but less so in obscura

in the Andamans. The single note of affinis is

clearly shown here.

I was able to observe both species at close

range in a powerful flashlight beam, wherein the

all-dark underparts of obscura and the streaked

underparts of affinis were conspicuous.

The song of obscura is a mellow hollow

double note whoo-wup with a rising inflection

and accent on the second note and is similar to

that of Ninox scutulata hirsuta of southern India

and Sri Lanka. It differs only slightly in having a

shorter gap between the two notes.

The song of affinis on the other hand, is a

mellow raspy single note woow. Often two notes

Refer

Abdulali, H. (1964): The Birds of the Andaman and

Nicobar Islands. J. Bombay nat. Hist. Soc. 61(3):

482-571.

are uttered (with a gap between them) as a couplet.

The sonograms show the close relationship

of obscura with mainland scutulata and the

distinctness of affinis.

Acknowledgements

I particularly wish to thank S.A. Hussain

for arranging the trip and being a great field

companion. Jeff Groth prepared the sonograms.

John Fitzpatrick and Greg Budney of the Macaulay

Library of Natural Sounds at the Cornell

Laboratory of Ornithology provided tape

recording equipment and other assistance. All the

tape recordings were made by me and will be

eventually on deposit at the Macaulay Library of

Natural Sounds at the Cornell Laboratory of

Ornithology.

: n c e s

Ali, S. & S.D. Ripley (1981): Handbook of the Birds of

India and Pakistan. Vol 3, 2nd Edn., 327 pp.

Oxford University Press, New Delhi.

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389

LEARNING ABOUT VOCAL COMMUNICATION IN BIRDS

T.J. Roberts1

Key words: Birdsong, recordings, Pakistan, sonograms, song function

Birdsong is of value to observers in identifying and locating unseen species. Modern advances such

as magnetic high frequency tape recording and sonograms have led to a greater understanding of the

subject. However, there are limitations to interpretation of sonograms. Reaction of birds to playback

of songs can help distinguish sibling species and even new species. Birdsong serves varied functions

such as in intra-specific recognition, females’ selection of mates, spacing out of territories, and in

colonial breeding species, mate or chick recognition. Variation in the songs of geographically

separated birds of the same species points to the role of learning by imitation in the development

of complexity of repertoire in a young bird. The role of voice in intra-specific recognition is

particularly critical in the case of cryptic nocturnal species with stereotypic songs. Also significant

are flight calls that keep the flock together and alarm calls that warn birds of danger.

As any birdwatcher knows, the quickest

way to find out what species are in the vicinity is

to leam to recognise their songs and calls. With

the great increase worldwide, in studying birds,

we have many new tools to familiarise ourselves

with the nature and meaning of birdsong. The

invention of the lightweight portable tape

recorder, the publication of compact disks with

collections of recorded birdsong and the

development of sonograms (also called

spectograms) which show the frequency band of

bird calls in a plot against time, have all increased

the non-specialist’s knowledge of this fascinating

aspect of bird behaviour.

During the 1970s and 1980s, I was able to

collect recordings of 319 species mostly from

Pakistan but also many from India (Roberts 1991),

and during these years, being keenly focussed

on their calls and song, was stimulated to continue

trying to keep up to date with recent research and

developments in this field.

As with all biological research, the more we

learn, the more questions remain unanswered, and

‘Cae Gors, Rhoscefnhir,

Nr. Pentraeth,

Anglesey LL75 8YU, U.K.

Email: [email protected]

a brief article such as this can do no more than

review current aspects, and recall some examples

from personal experience.

Firstly, the increasing use of published

sonograms in bird books and journals needs to

be better understood by the layman and amateur

such as this writer (Catchpole and Slater 1995,

Slater and Sellar 2000). You cannot differentiate

tone or pitch from a two-dimensional graph. A

sonogram can clearly show the small dialectical

differences between songs of the same species

and details of the make up of the song, which

would otherwise be undetectable to the human

ear. Birds, it is believed from experiments, can

detect higher pitched sounds up to 8,000 cycles

per second, usually called 8 kHz, well beyond the

limited hearing range of humans. Moreover, it is

therefore possible to show side by side on a

printed page, the sonograms of two or more

different birds, for a leisurely and careful

comparison that is not possible by actually

listening to separate voice recordings. We can

easily learn to recognise from the black lines and

smudges on a sonograph, the types of phrases or

strophes that make up a bird’s song repertoire.

Twittering shows closely spaced, rather short,

vertical black smudges while vehement high-

390

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003

VOCAL COMMUNICATION IN BIRDS

pitched calls are shown as longer darker bands in

a vertical plane. Also, the louder the note, usually

the wider is the black mark in a vertical plane. The

sort of calls or phrases that can be verbalised as

“beeze” will appear as a broad horizontal band.

But again these marks should be interpreted with

caution. The rapid ticking calls of some

Bradypterus warblers can appear very similar to

twittering, and one should always look carefully

at the vertical graph line showing increasing kHz,

as the lower down the scale, the deeper in pitch

the call which may otherwise look similar in shape

on the sonogram. Research in the development

of sonograms and study of birdsong has now

stretched over fifty years, and has come a long

way from the pioneering studies by Thorpe ( 1 958),

who showed that the songs of chaffinches

(Fringilla coelebs) which appear to be

stereotyped and similar within that species,

actually vary in small details both between

individuals and especially between different

regions, revealing the importance of dialectic

variation even in species with comparatively

consistent stereotyped songs. In recording the

social calls of a family group of common babblers

( Turdoides caudatus) on the western border

regions of Balochistan, I found their evening

territorial chorusing so different from those in the

Punjab plains, as to be at first unrecognisable. I

hazarded the opinion (Roberts 1992) that this

showed that the population had been

geographically isolated for a long period, so much

so it they probably would not interact with a family

flock from the Indus plains if it could be juxtaposed

nearby, because it is known that this very

sedentary species, living in tight family groups,

maintains its territory against rival groups, by

such evening choruses at the roost site. It is likely

that this development of local dialects is one of

the forces that lead to speciation. In Pakistan many

consider the grey-winged blackbird ( Turdus

boulboul) to be the finest songster, a view shared

by the great Dr. Salim Ali (1973) in his writings. I

taped continuous songs in the Murree foothills

of Pakistan, showing an unbroken sequence of

hundreds of different melodious and dramatic

strophes or phrases, especially in the early part

of the breeding season (Roberts 1 992). A few years

later, on encountering a small isolated population

of grey-winged blackbirds in a patch of deciduous

forest at Mahandri, in Hazara district, I was

astonished at the very limited and poor range of

their songs. This was proof of what was already

known about the importance for juvenile birds to

learn their songs by imitation. Research has more

recently shown that birds develop and learn the

complexities of their full song in the first year of

their lives, and thereafter there is no increase in

variety or complexity (Dowsett -Lamaire 1 979).

What then do we understand of the

purpose or functions of birdsong?

Firstly, it forms an essential ingredient in

intra-specific species recognition, not only for

successful breeding, but in the case of highly

colonial breeding species, for the recognition of

one’s chick, and in the clamour of a large breeding

colony, the voice of one’s mate. We still do not

understand how the Emperor penguin

{Aptenodytes forsteri) can recognise its offspring,

apparently by voice alone, in circumstances where

both parents have to be out at sea, food-hunting

for prolonged periods, and where the chicks form

vast huddled creches, constantly jostling for

better positions in the throng. The calls of the

parents are, to our ear, comparatively short and

raucous and sound remarkably similar. The minute

differences in pitch or phrasing are obviously not

detectable to the human ear and sonograms have

indeed shown that birds can detect such minute

differences.

The development of high frequency

magnetic recording tape, and the ability to include

playback microphones with such portable tape

recorders, has opened up a powerful new tool for

ornithologists. Since it is only males that have

elaborate territorial and mate attracting songs, the

playback of recorded song in the vicinity of a

certain male species can often result in an intense

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG. -DEC. 2003

391

VOCAL COMMUNICATION IN BIRDS

aggressive or curious reaction. So much so, that

this tool of playback and the degree of response

by the live bird is used to recognise and

distinguish sibling species and even new species

(Irwin et al. 200 1 , Collinson 200 1 ). In the view of

some conservationists, there is a downside to this

practice of searching for possible presence of

species by playing their songs when used

indiscriminately during the breeding season.

However, many reputable wildlife tour operators

use this tool with discrimination to enable their

clients to get the best chances of viewing shy

and elusive birds. The power of such recordings

was well demonstrated when KingBird Tours,

during an expedition to northwest China, played

the calls of that skulking and shy bird, the com

crake ( Crex crex ), and eventually had the excited

bird coming out of the tail grass on to an open

road and pursuing the human party, calling

repeatedly (King 200 1 ).

A second vital function of birdsong, which

has long been known, is mate attracting. Studies

of the long and complex songs of some

passerines have shown that there is a clear

correlation between the success of displaying

males with the longest songs and their ability to

attract females, and this may be an important

element in understanding why some species

include a huge range of almost perfect imitations

of other species’ calls in their song repertoire. Dr.

Salim Ali ( 1 972) records mimicry by Lanius schach

of over 30 different species of birds, as well as

realistic imitation of a puppy yelping. Others,

including this author (Dowsett-Lemaire 1979,

Roberts 1 992), have also noted that the mimicry

by some songsters included calls of species they

could encounter only in their winter migratory

territory, not in their breeding grounds. While

recording the display of the Isabelline wheatear

(Oenanthe isabellina) in Baluchistan, this author

was intrigued to hear clearly, the imitated flight

calls, of wood sandpipers ( Tringa glareola ) and

red-wattled lapwings ( Vanellus indicus ), both not

found on their nesting grounds, as well as such

life-like imitations of a shepherd whistling to

urge his flock on and a puppy sqealing, that a

nearby dog became very excited looking for the

source.

Thirdly, song is also used by males to warn

off rival males and to define and maintain

territorial boundaries, as already discussed under

the use of playing back such recordings in the

field. While recording the song of the White’s

mountain or scaly thrush ( Zoothera daumd) in a

wide forested mountain amphitheatre, I was

surprised to learn, over several evenings, how

many widely spaced song posts were used by

this bird, indicating the large area that it hoped to

maintain as its breeding territory (Roberts 1992).

On another occasion, trying to attract a collared

scops-owl ( Otus bakkamoena ) for identification

on a dark night, my teenage son was suddenly

dive-bombed by the irate bird, to our mutual

surprise and fright!

Returning to the importance of voice in

intra-specific recognition, this is especially critical

in nocturnal species, especially those with cryptic

plumage. In our studies on the songs of scops-

owls of the Otus genus, it soon became apparent

that many sibling species which could only be

separated with difficulty in the hand, and which

had previously been separated only as

subspecies, were in fact quite distinct sympatric,

non inter-breeding species (Roberts and King

1986). The same holds true to a larger or lesser

extent for many of the Caprimulgidae and

Acrocephaline warblers, characteristic of skulking

lifestyles, in a restricted visual habitat of reed

beds.

In a short article of this nature, it is not

practicable to discuss the important and varying

roles also played by flight, contact and alarm calls.

The importance of the continuously uttered

contact calls that keep the flock together, to such

gregariously foraging birds as the minivets

( Pericrocotus spp.), as they sometimes erratically

search over the forest canopy, will be apparent to

any keen birdwatcher. Studies of alarm or warning

392

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3). AUG. -DEC. 2003

VOCAL COMMUNICATION IN BIRDS

calls of small passerines have shown that they

are rather low pitched and often very similar among

different species. Such calls being of low

frequency, carry further, and are more difficult to

locate direction-wise, thus concealing the location

of the alarm giver, and also enabling nearby birds

of other species to respond quickly to potential

danger (Catchpole and Slater 1995)

The Crane Family with their lifelong pair

bonds, longevity, and courtship duetting and

dancing have great appeal to many national

cultures. When in flight their ringing calls, as

everyone knows, carry amazing distances.

Dissection of their trachea (windpipe) reveals a

convolution, reminiscent of a French horn, a

gradually evolved anatomical feature which must

partly explain the continuous though precarious

survival of this ancient bird family which needs

wide open spaces far from rival pairs for nesting,

yet can congregate in astonishingly huge flocks

Refer

Ali, SAlim (1972): Handbook of the Birds of India and

Pakistan, Vol. 5. Oxford University Press, Bombay.

Pp. 113.

Ali, Salim (1973): Handbook of the Birds of India and

Pakistan, Vol. 9. OUP, Bombay, Pp. 94-95.

Alstrom, P, U. Olsson & P.R. Colston (1992): A new

species of Phylloscopus warbler from central China.

Ibis 134(4): 329-334.

Catchpole, C.K. & P.J.B. Slater (1995): Bird Song:

Biological Themes and Variations. Cambridge

University Press, Cambridge, U.K.

Collinson, Martin (2001): Greenish Warbler, Two-barred

Greenish Warbler, and the speciation process.

British Birds 94( 6): 278-283.

Dowsett-Lemaire, Franqoise (1979): The imitative range

of the song of the Marsh Warbler Acrocephalis

palustris, with special reference to imitation of

African Birds. Ibis 121(4): 453-468.

Irwin, D.E.. P. Alstrom, U. Olsson & Z.M. Benowitz-

during migration. The overhead calls of even the

smallest skein of cranes act as a clarion call to any

small populations on the ground below.

Lastly, the importance of song in taxonomy

is only recently being appreciated. We live in an

era of taxonomic turmoil with the recent advances

in molecular genetics and DNA sequencing

forming powerful tools in examining inter-species

relationships, and ultimate phylogenetic

relationships are still being disputed. Yet,

increasingly, sibling species are first recognised

in the field as being distinct and often new species,

on the basis of their taped songs and reaction to

playback (Alstrom et al. 1992, Collinson 2001).

Whilst the wealth of new research is teaching us

more about the complexities and meaning of

birdsong and calls, we should never lose sight of

the aesthetic pleasure we get from being able to

pause in our busy life, simply to listen and to

enjoy their chorus.

NCES

Fredericks (2001): Cryptic species in the genus

Phylloscopus (Old World leaf warblers). Ibis

143(2): 233-247.

King, Ben (200 1 ): Tour Reports - 200 1 N W China Tour,

Xinjiang. KingBird Tours , Inc. Newsletter 43: 10.

KingBird Tours, Planetarium Station, New York,

USA.

Roberts, T.J. (1991): The Birds of Pakistan, Vol. 1.

Oxford University Press, Karachi. Pp. 598.

Roberts, T.J. (1992): The Birds of Pakistan, Vol. 2. Oxford

University Press, Karachi. Pp. 617.

Roberts, T.J. & Ben King (1986): Vocalizations of the

Owls of the genus Otus in Pakistan. Ornis

Scandinavica 7(4): 299-305.

Slater, P.J.B. & P.J. Sellar (2000): Understanding

Sonograms. British Birds 93(7): 323-329

Thorpe, W. H. (1958): The learning of song patterns by

birds, with special reference to the song of the

Chaffinch Fringilla coelebs. Ibis 100: 535-570.

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG. -DEC. 2003

393

BIRD SPECIES DIVERSITY ALONG THE HIMALAYA:

A COMPARISON OF HIMACHAL PRADESH WITH KASHMIR

( With two text-figures and two plates)

Trevor Price1’2, Jennifer Zee1, Kartika Jamdar3 and Nitin Jamdar3’4

Key words: Avian diversity, Himachal Pradesh, Kashmir

About 80% of the birds of the Indian subcontinent can be found within the Himalayan region. We

studied variation in bird numbers along the Himalaya. There are more than twice as many species

in the east than there are in the west. Most of the decline in species numbers from east to west

occurs between western Nepal and northern Pakistan, where the trend of the Himalaya follows a

more northerly course. We compared the breeding birds of two sanctuaries across this region of

steep decline in species numbers, Manali in Himachal Pradesh, whose birds are described for the

first time in this paper and Overa in Kashmir (Price and Jamdar 1 990, 1991). Differences in species’

composition are large, e.g. one-third of the high elevation (c. 3,000 - 3,500 m) species were recorded

from only one or other sanctuary. Some of the missing species from one or other sanctuary were not

a consequence of a range limit between the sanctuaries, but rather a result of a gap in the distribution.

Species with patchy distributions included the western tragopan Tragopan melanocephalus whose

range was historically known to be much more extensive, and the Nepal wren-babbler Pnoepyga

immaculata. We conclude that there is considerable variation in species composition along the

Himalaya and this is being accentuated by local population extinctions.

Introduction

Large numbers of bird species live in the

Himalaya (Meinertzhagen 1928, Inskipp and

Inskipp 1985, Martens and Eck 1995). In addition,

species composition varies along the Himalaya

and there are many more species in the east than

in the west (Inskipp and Inskipp 1985). While

this pattern is qualitatively known, it has not been

quantified, and its underlying cause is not

understood. In this paper, we investigate

variation in species number along the Himalaya.

We use a broad scale comparison based on

published species range maps to document

general patterns, and then a more detailed

comparison of two sites in the Northwest

Himalaya (in Himachal Pradesh and Kashmir)

where we have conducted fieldwork over the past

‘Section of Ecology, Evolution and Behavior,

Division of Biological Sciences, 0116

University of California at San Diego,

La Jolla CA 92093-01 16, USA.

2Email: [email protected]

J5-A Samata, Gen. Bhosale Marg, Mumbai 400 021, India.

4Email: [email protected]

15 years. We investigate how total species

numbers change along the Himalaya and the

extent to which species composition varies

between different localities.

Although the Himalayan range is often

considered to run from east to west, particularly

from Nepal west it follows a southeast to

northwest trend, and spans c. 5 degrees lati-

tude from western Nepal to Srinagar in Kashmir

(Plate 1, Fig. A and Plate 2, Fig. B). As we will

show, total species numbers along the Himalaya

decline mostly in association with this latitudinal

gradient, rather than with longitude. The two sites

we worked at (Manali Sanctuary in Himachal

Pradesh and Overa Sanctuary in Kashmir) are

located in a region where there is a steep decline

in species numbers. The data from Overa have

been previously published (Price and Jamdar

1 990, 1991), but the data from Manali are presented

here for the first time. Therefore, a second purpose

of this paper is to document the breeding birds

(including range extensions and altitudinal

distributions) for Manali Sanctuary for which

there is no previously published ornithological

survey.

394

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003

BIRD SPECIES DIVERSITY ALONG THE HIMALAYA

Price, Trevor et a/.: Map

Plate 1

V- *

(Mm.

V/i , :%%%

mmi 98 153 238 261 ww Ai ™u B W0

7ft 4 WN CN EN WB ‘EB/jf

1 ■% . , Jf

163 149 188 192 212 209 210 224 252 256 4#^

; ; >, ■'%

'% • y/.

58 181 216 219 225 233 214 235 ,256 222 314|

139 211 255 214 229 240 246 213

L ■ | ) (:/ 4|

187 266 201 235 241 197

,// 4. W

■ 2.3.4' .//.

175 ,304 206

231 215

Bl|l

'■?/ ■

186214

Fig. A: Numbers of breeding species in 250 x 250 km squares across the Indian subcontinent,

based on Grimmett et al. ( 1 999a)

Squares with letter symbols are those subject to more detailed analysis in the text and in Fig. 1.

Symbols indicate rough locations: PK - northern Pakistan, KM - Kashmir, HP - Himachal Pradesh, UT - Uttaranchal,

WN - western Nepal (two squares are included because neither alone covers the complete altitudinal range),

CN - Central Nepal, EN - Eastern Nepal, WB - Western Bhutan, EB - Eastern Bhutan, AP - Arunachal Pradesh.

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG. -DEC. 2003

395

Price, Trevor et al. : Map Plate 2

BIRD SPECIES DIVERSITY ALONG THE HIMALAYA

fwMmm.

mw^m^assBSBBSt

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396

JOURNAL , BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG. -DEC. 2003

Fig. B: Locations of the two long-term study sites and annual precipitation estimated for half-degree coordinates along the Himalaya

(see New et al. 1 999, 2000)

BIRD SPECIES DIVERSITY ALONG THE HIMALAYA

Methods

Broadscale comparisons

Taxonomy and nomenclature throughout

this paper follow Sibley and Monroe (1990). We

used a field guide (Grimmett et al. 1999a) to

estimate the numbers of breeding bird species at

different locations in the Indian subcontinent.

We excluded species that were primarily

pelagic, leaving us with a dataset of 976 breeding

species that we considered to be depen-

dent on land or freshwater. The species used

and the data collected are available at

http://www.biology.ucsd.edu/tprice. Following

the methods of Fjeldsa (1994), we overlaid a

transparent grid of 14 squares x 14 squares on

each map in the field guide. Each square is

approximately 250 km x 250 km. Plate 1, Fig. A

shows the grid with the approximate position of

the squares (position is approximate because the

map projection used by Grimmett et al. (1999a)

differs from the one used here). Then we recorded

the presence or absence of the species in each

square. With this information, we were able to

calculate the number of breeding species in each

square, and the extent to which different squares

share species. We also used Ali and Ripley (1987)

and Grimmett et al. ( 1 999b) to assess the altitudinal

bands in which a species is found (divided into

1,000 m intervals). For many species altitudinal

distributions vary considerably along the

Himalaya (Martens and Eck 1995). We ignored

this spatial variation, simply recording minimum

and maximum elevations as far as are known.

We obtained climate data described in

New et al. (1999, 2000) and available at

http://mercurv.ornl.gov/omldaac. This dataset

provides (extrapolated) mean monthly averages

for eleven climate variables at 0.5° latitude and

longitude intervals across the whole land surface

of the world. We based our analysis on

measurements made between 1901 and 1960. We

extracted mean annual precipitation for points

along the Himalaya. We also used measurements

of climate reported for individual meteorological

stations in the Himalaya (Anon. 1967).

Field studies

The two study sites are Overa (33° 80' N,

75° 40' E) near Pahalgam in Kashmir, and Manali

(32° 20' N, 77° 20' E) in Himachal Pradesh (Plate 2,

Fig. B), 250 km to the southeast of Overa.

Overa Sanctuary (33 sq. km) forms the

watershed to a small valley running approximately

northwest to southeast (Price and Jamdar 1990).

The elevation spans from about 2,400 m to above

the treeline. We were camped at sites at

approximately 2,400 m or 3,300 m from May to July

in each of the three years from 1985-1987, as

described in Price and Jamdar ( 1 990, 1991). Either

TP or NJ returned to Overa for a period of 1-2

months (May or May-June) in each of the

subsequent four years, from 1988-1991. Our latest

observations confirmed the previous findings,

with a few additions reported in Appendix I.

Manali Sanctuary (32 sq. km) is a watershed

similar in size to Overa, but the main river runs

west to east (Singh et al. 1990). The altitudinal

range is from c. 2,000 m (just outside the town of

Manali) to above the treeline. We spent the

following periods in Manali Sanctuary, camped at

3, 1 00 m: May-June 1 994 (TP), early May 1 995 (TP),

late June 1996 (TP), late June - early July 1997

(TP), May-June 200 1 (TP, N J, KJ), and May 2002

(NJ, KJ).

We recorded all the birds observed in a daily

logbook. We estimated bird abundance using

point censuses and mistnet capture rates, but

found the most satisfactory measure to be simply

the fraction of days a species was observed, which

is reported here. This measure is weakly correlated

with mistnet captures [correlation coefficient « 0.5

in Overa (Price and Jamdar 1 990)].

We made a special effort to assess altitudinal

distributions. These are poorly known for the

Himalaya, partly because they can vary greatly

from place to place (Martens and Eck 1 995), and

partly because it has been difficult to separate

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG. -DEC. 2003

397

BIRD SPECIES DIVERSITY ALONG THE HIMALAYA

breeding from winter and migratory records (e.g.

Inskipp et al. 2000).

Altitudinal distributions must always be

approximate. Firstly, they vary even within a valley.

For example, south-facing slopes in Manali have

much more extensive kharsu oak Quercus

semecarpifolia , whereas north-facing slopes have

more extensive fir Abies pindrow, and bird species

associated with these habitats have different

altitudinal distributions on each side of the valley.

In Manali, we worked on a north-facing slope.

Secondly, individual singing males may occupy

territories far out of the normal range. This is

particularly true for estimates of lower boundaries,

as occasional males may remain singing and

unmated at low elevations until late in the season

(see for example, bimodal distribution of

Phylloscopus reguloides in Fig. 2, probably as a

result of 1 or 2 singing males at low elevations,

where the species was not generally observed), but

may also apply to upper boundaries (for example, a

lone Cettia fortipes established a territory at

c. 3,000 m in Overa). Thirdly, rare species are easily

overlooked. In Overa, we attempted to estimate lower

and upper altitudinal bounds for each species,

based on localities where it appeared that several

pairs were likely to be breeding. We based our

estimates on repeated observations of birds

throughout the 3 years of our initial study, and found

these to be upheld in four subsequent seasons. In

Manali, we conducted censuses along the entire

altitudinal gradient by halting for 2 minutes at

intervals of every 25 m altitude (measured using an

altimeter), and recording all birds seen and heard

(this was conducted over a period of 2-3 mornings

from 0600-0900 hrs in each of the three years 1 996,

1997 and 2001). We then listed the altitudinal

distributions of all species from these censuses,

and evaluated the list on a species by species basis

in the field in May 2002. Despite all the caveats

associated with estimates of altitudinal distributions,

species differ considerably in where they live on

the mountain, and our estimates are consistent

between years.

During the course of the altitudinal census

at Manali, we also recorded all tree species within

an approximate 8 m radius of the census point.

Results

Broad patterns

The number of species in each square across

the Indian subcontinent is shown in Plate 1, Fig. A.

Results confirm that the Himalaya harbours a large

number of species. Indeed, one 250 km x 250 km

square in central Nepal contains 575 species, or

60% of all the land and freshwater species

breeding in the Indian subcontinent (including

the Andaman and Nicobar Islands) and almost

twice the maximum diversity recorded from a

square in peninsular India. We combined results

from 1 1 squares running the length of the Himalaya

from Pakistan to Arunachal Pradesh (see squares

with letter symbols in Plate 1, Fig. A). These 1 1

squares contain a total of 783 species or 80% of

all the species found on the Indian subcontinent.

This partly reflects the great range of climate

regimes, from tropical to temperate, and hence

diversity of habitats. For example, 154 (20%) of

the 783 Himalayan species breed entirely above

2,000 m. An additional 249 species (32%) are found

breeding above 2,000 m somewhere in their

geographical range, as well as at lower elevations.

The other species occur at lower altitudes,

including the plains of India.

There are many more species in the east

than in the west. Across the 1 1 squares running

the length of the Himalaya, the easternmost, in

Arunachal Pradesh, contains more than twice the

number of species of the westernmost square, in

Pakistan (507 vs 233 species). The decrease in

species number is mainly from Uttaranchal to

northern Pakistan, i.e. where there is a substantial

south-north as well as east-west trend of the

Himalaya. The square including Uttaranchal

contains almost 500 species, similar to the square

in Arunachal Pradesh.

The steep decline in species numbers

across the Northwest Himalaya is associated with

398

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG. -DEC. 2003

BIRD SPECIES DIVERSITY ALONG THE HIMALAYA

a decline in precipitation (Plate 2, Fig. B). In

Pakistan, the lower reaches of the Himalaya are

very dry and semi-desert. In comparison with

Nepal, northern Pakistan has much less monsoon

rainfall, more winter precipitation and colder

winters (Stainton and Polunin 1984).

Superimposed on changes in total species

number is species turnover. 1 16 species occur in

both the westernmost and easternmost squares,

i.e. Pakistan (PK) and Arunachal Pradesh (AP),

implying that 50% of all the species in the Pakistan

Himalaya extend their range throughout the

Himalaya. There are many more species in

Arunachal, and hence a smaller fraction (23%) of

these species extend west to Pakistan. Among

species recorded breeding above 2,000 m, 63

species range throughout the Himalaya, which is

43% of the Pakistan total (26% of the Arunachal

total).

In Fig. 1, we have plotted the turnover

between adjacent squares along the Himalaya.

This plot is for all species recorded breeding above

2,000 m altitude somewhere in their geographical

range, but the patterns are similar if all species are

included. The histograms give the fraction of

species in a square that have western or eastern

range limits in that square. For example, 30% of

the species in the square KM are not found in the

square PK whereas 15% of the species in the

square KM are not found in the square HP

Typically about 1 0% of all species in a square

do not extend to the neighbouring square, and

patterns of turnover are not strikingly different at

different points along the Himalaya or to the east or

west. The main exceptions are the squares KM and

HP which have a high fraction of species (>30%)

that do not extend their ranges to the northwest.

Thus, even though some species are confined to

the northwest, the southeast to northwest

latitudinal gradient in species numbers reflects the

fact that larger numbers of southeastern species

fail to extend their ranges northwest.

c n

LU

O

LU

CL

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0

z

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Fig. 1 : Species turnover along the Himalaya.

Letter symbols refer to squares indicated in Plate 1, Fig. A, with the two WN squares combined

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BIRD SPECIES DIVERSITY ALONG THE HIMALAYA

Himachal Pradesh vs Kashmir

We now investigate the pattern of high

species turnover along the Northwestern

Himalaya by comparing Manali (Himachal

Pradesh) and Overa (Kashmir) Sanctuaries.

Vegetation and climate: The altitudinal

distributions of the major tree species for the

north-facing slope in Manali are shown in Fig. 2

(for similar data on Overa, see Price and Jamdar

1990, and Price 1991). The vegetation at Manali

differs from that at Overa in several ways. Most

strikingly, kharsu oak is common in Manali, but

absent in Overa (Price and Jamdar 1990).

Rhododendron is much more abundant in Manali

than in Overa. Juniper, at least in the area where

we worked, is much less common in Manali than

in Overa. Among tree species held in common,

maple Acer spp., fir Abies spp., pine Pinus

wallichiana, spruce Picea smithiana and birch

Betula utilis have similar altitudinal distributions

in the two sanctuaries.

We do not have comparable climate

Spruce

Cedar Chestnut

Fir

Maple

Oak

Rhododendron

Birch

Phylloscopus magnirostris

Phylloscopus occipitalis

Phylloscopus chloronotus

Phylloscopus reguloides

Phylloscopus maculipennis

Phylloscopus humei

Phylloscopus trochiloides

Phylloscopus pulcher

Seicercus xanthoschistos

Seicercus whistleri

Luscinia brunnea

Tarsiger cyanurus

Tarsiger chrysaeus

2000 2500 3000 3500

Altitude (m.)

Fig. 2: Distributions of the major tree species along the Manali transect (top) and census records for species

of four common bird genera conducted on June 3-5, 1997

Top: = 2 or more individuals recorded within an approximately 8 m radius.

the extreme records for the species as observed in the 8 m radius.

Bottom: = range over which more than one individual was recorded at each census point.

the range over which one individual was recorded.

Two minutes were spent at altitudinal intervals of 25 m along the transect, early in the morning.

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BIRD SPECIES DIVERSITY ALONG THE HIMALAYA

measurements from Overa and Manali, partly

because they were studied over different years

and the climate may have been changing: snow

disappeared from our camp in Manali

approximately one month earlier in each year of

200 1 -2002 than in each year of 1 994- 1 997 (see also

Vedwan and Rhoades 2001). Nevertheless, Manali

is probably wetter (Plate 2, Fig. B) and warmer

than Overa. To assess this indirectly, we compared

climate records for Shimla, Himachal Pradesh

(3 1°N, 77° E) with Srinagar, Kashmir (34°N, 75.5° E),

based on 30 years of data (1931-1 960; Anon. 1 967).

Although Shimla is 700 m higher, it has warmer

springs (average minimum temperature in May in

Shimla is 4 °C higher than in Srinagar) and almost

three times the annual rainfall of Srinagar.

Birds of Manali Sanctuary: We recorded a

total of 137 species in or near Manali Sanctuary.

The data are summarised in four exclusive lists. In

Table 1, we present the 81 species encountered

above about 2,800 m (higher elevations) in at least

two different years with altitudinal distribution

and estimates of abundance. Table 2 lists common

breeding species in the forests below 2,500 m, with

altitudinal distribution. These tables are separated

because we spent little time at the lower altitudes,

have less quantitative data, and may well have

overlooked some species. Appendix II refers to

additional species that have been recorded rarely

in Manali Sanctuary and Appendix III additional

species that primarily breed along the lower

boundary of Manali Sanctuary in more open

habitat.

In Fig. 2 we show altitudinal distributions

based on a single census for common species

belonging to four genera. Some species appear to

be tightly associated with particular habitats, such

as Phylloscopus humei with birch (Price and

Jamdar 1991). Two species are particularly

noteworthy. The first, Seicercus whistleri,

appears to have a disjunct altitudinal distribution

(Fig. 2). This was noted further east in the Himalaya

by Alstrom and Olsson (1999) and Martens et al.

(1999) who described the lower elevation

population as a separate species (Seicercus

burkii ); the upper population is now classified as

Seicercus whistleri\ (see also Alstrom and Olsson

2000). The possibility that both species are present

in Manali needs to be investigated. Secondly,

there is a breeding population of the Nepal wren-

Table 2: Common forest bird species in the lower part of Manali Sanctuary that do not extend to higher elevations*

Common name

Latin name

Altitudinal distributions (in m)

Low High

2Great barbet

2Ashy drongo

^lue-headed rock-thrush

2Grey-winged blackbird

^usty-tailed flycatcher

Verditer flycatcher

2Grey-headed flycatcher

^reen-backed tit

^lack bulbul

2Grey-hooded flycatcher-warbler

Nepal wren-babbler

Yellow-naped yuhina

Megalaima virens

Dicrurus leucophaeus

Monticola cinclorhynchus

Turdus boulbou /

Muscicapa ruficauda

Eumyias thalassina

Culicicapa ceylonensis

Parus monticolus

Hypsipetes leucocephalus

Seicercus xanthoschistos

Pnoepyga immaculata

Yuhina flavicollis

Present also in Overa. Muscicapa ruficauda is much more common in Overa than Manali.

2These species were not recorded in Overa, but are common elsewhere in Kashmir

and their absence may reflect the absence of forest habitat below 2,400 m in Overa.

*For additional common species that extend into Manali town, see Appendix III.

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Table 1: Common species found above 2,500 m in Manaii Sanctuary

Comparative information for Overa is from Table 4 of Price and Jamdar (1990)

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Table 1: Common species found above 2,500 m in Manali Sanctuary

Comparative information for Overa is from Table 4 of Price and Jamdar (1990) (contd.)

Common name Latin name Abundance Altitudinal distributions

(in m)

1Manali *Overa Manali Overa

1 Proportion of days observed when camped at 3,100 m. Based on n = 84 days; 34 days in June/July 1996 and 50

days in May/June 2001.

* Proportion of days observed when camped at 3,300 m (n=224 days in 1985-1987).

2Although absent from Overa the species has been recorded as breeding (rarely) in Pakistan (Roberts 1991 , 1992)

3A single singing male present at Overa in several years.

4 We did not observe this species as we paid little attention to the swifts, nevertheless it may have been present

at Overa, which is close to its range limit.

5 We did not always distinguish between Certhia himalayana and Certhia familiaris.

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babbler Pnoepyga immaculata between

c. 2,300 m and 2,600 m in the chestnut-spruce-fir

woods just above the entrance to the Sanctuary

(mostly between 2,300 m and 2,450 m). This

species was first described in 1991 (Martens and

Eck 1991) and previously only recorded from

Nepal. An individual was captured and examined

in the hand on May 22, 2001 (wing-length 55 mm,

weight 15 g).

Comparison of Manali with Overa: In our

study, Manali and Overa can be best compared in

the composition of higher elevation species

(c. 3,000 m - 3,500 m), the habitat being less

disturbed, and because we spent much more time

at these elevations. For Manali this includes all

81 species in Table 1. There are an estimated

70 common species at Overa at these elevations

(see Table 4 in Price and Jamdar 1990).

Twenty-one species are present on the

Manali higher elevation list but not in Overa, and

1 1 species are present on the Overa higher

elevation list but not in Manali. Fifty-nine species

are included in the lists from both Manali and

Overa. Thus 64% of the total number of species

in the two sanctuaries combined are found in both

sanctuaries and 36% are found in one or other

sanctuary.

High elevation Manali species missing

from Overa: Of the 21 species present at high

elevations in Manali and not in Overa, 5 have

been recorded breeding in northern Pakistan

(Roberts 1991, 1992). Apart from Tarsiger

chrysaeus, the remaining 4 (see Table 1 ) are rare

in Manali and may occasionally occur in Overa

but may have been overlooked. However, they

may also reflect a genuinely patchy distribution,

perhaps due to recent extinctions. Tragopan

melanocephalus is one such example: we never

recorded the species in Overa, but there is a small

breeding population in Pakistan (Roberts 1991).

One species, Collocalia brevirostris, has

its range limit in Kashmir, but may have been

present in Overa and been overlooked by us. A

general feature of many of the species absent or

rare in Overa is that they are very common in

Manali. Six of them were among the 15 that we

recorded on more than 90% of the days (Table 1 ).

For example, Minla strigula and Rhipidura

hypoxantha are two of the commonest species at

Manali, but they have never been recorded

breeding as far west as Overa. Causes of the

absence of these and most of the other species

are not understood. It is notable that several

ground and bush foraging insectivores that are

common in Manali are missing from Overa (Ant bus

hodgsoni, Pnoepyga immaculata, Tesia

castaneocoronata, Cettia flavolivacea,

Bradypterus thoracicus).

The absence of two species from Overa

( Turdus albocinctus and Pyrrhula

erythrocephala) is associated with the presence

of an ecologically similar species in similar habitat

there (compare Table 1 with Table 3). Turdus

albocinctus is replaced by Turdus rubrocanus in

Overa. In 1 996, a pair of T. rubrocanus was found

breeding in Manali and males of the two species

were observed chasing each other. Both these

species have similar songs and may mutually

exclude each other. Note, however, that

T. rubrocanus does occur east through Nepal (Ali

and Ripley 1987), and both T. rubrocanus and

T. albocinctus breed in similar habitat in Bhutan

(Inskipp et al. 2000). The absence of other species

may be related to the absence of their preferred

habitat. In particular, Phylloscopus maculipennis

and P. reguloides are typically found in

association with kharsu oak that is absent from

Overa and further west.

High elevation Overa species missing from

Manali: Eight of the 1 1 species present at high

elevations in Overa but not present at high

elevations in Manali are given in Table 3. Of the

other three species, Garrulax lineatus and

Emberiza cia are present and common in Manali,

but only at low elevations (Appendix III). The

third species missing from Manali, but not listed

in Table 3, is Phoenicurus caeruleocephalus ,

which is very rare in Overa.

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Table 3: Species common in Overa but absent or rare in Manali

’Abundance at the high altitude sites (c. 3,200 m). See Table 1 for explanation.

‘Common below 3,000 m in Overa (Price and Jamdar 1990) but never observed in Manali.

2 In Manali, a nest with 3 eggs found at 3,500 m on June 1 8, 1 996; an individual seen at 3,240 m on May 1 5, 2002.

3ln Manali, a pair breeding in a small patch of juniper in 1 994, and a single singing male in early May, 1 995. May be

common elsewhere in the Sanctuary where there is much juniper.

4Recorded breeding as far east as Nepal (Grimmett et al. 1999b).

5Two individuals seen near Manali town on May 8, 2002.

Eight of the 1 1 high-elevation species

absent from Manali actually have ranges

extending into Nepal, and their absence from

Manali is attributable to patchy distribution.

Causes of absence of these species from Manali

are generally unclear, but several species have

ecologically similar replacements (as noted in the

previous section). Two species that are common

in Overa, Phylloscopus affinis and Luscinia

pectoralis, occur in juniper habitat which was rare

at our study site in Manali. Both species have

ranges extending through Nepal, and the

patchiness in their distribution reflects the same

in habitat distribution.

The three species present in Overa, but

missing from Manali, which do not extend east

are Apus apus (we have an unconfirmed sight

record of this species in Manali), Phylloscopus

tytleri and Pyrrhula aurantiaca. All three species

are common in Overa. As in the case of Manali

species not present in Overa, species abundant

in one sanctuary are often absent from the other.

Altitudinal ranges: In general, and within

the limits of our ability to accurately estimate

altitudinal ranges, species that occur in both

locations occupy similar altitudinal distributions.

In particular, relative placements of congeners are

never altered. There are three examples of large

differences in altitudinal distribution between

populations of the same species. All of these

involve species in Overa extending to higher

altitudes than in Manali ( Phylloscopus

occipitalis , Garrulax lineatus and Emberiza cia).

Manali has the generally warmer climate, so we

expected species at low elevations in Overa to

have higher altitudinal distributions in Manali,

but we found no clear examples of this pattern. It

is possible that, given the lower species diversity

in Overa, the altitudinal range extensions there

are a response to reduced competition.

Similarities between sites: We calculated

similarity indices (Magurran 1988, p. 95) between

high and low elevations at Manali and Overa

(Table 4). We restricted this analysis to passerines,

which provide a more homogeneous set. The

similarity indices are designed to run from 0 to 1 .

If there were an equal number of species in each

site, all at the same density, an index of 0.5 would

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BIRD SPECIES DIVERSITY ALONG THE HIMALAYA

Table 4: Similarities between sites for passerine birds

Data from Table 4 of Price and Jamdar (1990) and

Tables 1-3 in this paper.

Above the diagonal: Jaccard’s index = (number shared

species/total number species)

Below the diagonal: Morisita-Horn index, which weights

species by their density (see Magurran 1988)

Densities were the fraction of days observed. This

data is not available for low Manali.

Low sites are centred at approximately 2,400 m, high

sites at approximately 3,200 m.

Low sites have fewer species because only forest

species were included.

imply that half the species are present in both

sites. Similarities within elevations across sites

are about 0.6. Similarities between elevations run

from 0.23 to 0.5 1 (true similarities are lower because

only genuinely forest species are included for the

lower sites; species of more open land are

excluded). A shift in approximately 1,000 m

elevation clearly results in more species’ turnover

than the shift of 250 km between Manali and Overa.

Discussion

In South America, bird diversity is

correlated with topography, precipitation and an

interaction between topography and latitude

(Rahbek and Graves 2001). The most diverse

locality is in the Andes on the equator, which

has: i) very high rainfall, probably related to

productivity, and ii) a great diversity of habitats

from tropical rainforest to Alpine tundra. A similar

association is apparent across the Indian

subcontinent: 80% of all land and freshwater bird

species can be found in the vicinity of the

Himalaya, and within the Himalaya the wetter

southeast has a greater diversity of bird species

than the drier northwest. Stainton and Polunin

(1984) note similar patterns among the flowering

plants.

As one moves from the northwest to

southeast along the Himalaya, roughly 1 0% of all

species occurring at one location are absent at a

location 250 km further on. As one moves from

southeast to northwest, the figures are more or

less comparable, except from Nepal onwards,

where about 30% of all species present at one

location are missing from a location 250 km further

northwest. The result is a steep decline in species

diversity as one moves from west Nepal to

northern Pakistan (as noted by Meinertzhagen

1928). The decline is associated with a latitudinal

change of about 5 degrees, decreased

precipitation and cooler spring weather.

The causes of turnover of species along

the Himalayan transect are far from understood,

so we undertook a comparison of the birdlife of

two small sanctuaries separated by 250 km. We

found that some species in one place are

apparently replaced one-for-one by similar

congeners in the other. Thus Phylloscopus

occipitalis has a greater altitudinal range in

Kashmir than in Himachal and this is associated

with the absence in Kashmir of the similar

Phylloscopus reguloides, occurring at high

elevations in Himachal (Manali). The absence of

P. reguloides from Kashmir is attributable to the

absence of kharsu oak. We made a special study

of the turnover of only Phylloscopus warblers

(Price 1991, Price and Jamdar 1991), but more

detailed studies of other species may result in

similar explanations. At present, many patterns

are unexplained. Several well-defined ecological

groups present in Manali are simply missing from

Kashmir, with no obvious counterparts. This

includes some of the commonest birds in Himachal

Pradesh such as a whole suite of ground foraging

insectivores, a fantai 1-flycatcher and a minla.

One finding is that rarity in one sanctuary

is no indication of absence from the other. At the

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same time, some of the commonest species in one

sanctuary are entirely missing from the other. A

possible explanation is that conditions in the non-

breeding season limit population sizes. If this is

the case, then subtle shifts in breeding

requirements may be sufficient to concentrate the

species in one location. For example, Phyllos copus

tytleri has a very small range in Kashmir and

northern Pakistan (where it is abundant) but does

not occur in Manali. Price (1999) suggested that

the population size of P. tytleri was severely

limited by its specialised wintering requirements

(the species is migratory and overwinters at

relatively high altitudes in the central Western

Ghats).

Our studies at Manali have revealed the

presence of several species previously

unrecorded in Himachal Pradesh (see Table 2 and

Appendix II). Most striking was the discovery of

Pnoepyga immaculata, previously thought to be

confined to Nepal (Martens and Eck 1991). This

discovery highlights the fact that many species

have patchy distributions. In our comparison

between the two sanctuaries, we found several

examples where absence from one or other

sanctuary is a result of a patchy distribution rather

than the true species’ range limit lying between

the sanctuaries. Six of the 10 species listed in Table

3 that are common in Overa but absent from Manali

actually have large populations elsewhere to the

east of Manali, in some cases extending through

Nepal. Patchy distributions are more striking for

species present in Overa and missing from Manali

than the converse. This may reflect the much

greater extent of the Himalaya to the east rather

than the west.

Although range maps in Grimmett et al.

(1999a) indicate several species with clearly patchy

distributions on a regional scale, patchiness on a

more local scale has scarcely been investigated,

and needs more research. For example, we do not

know how extensive the distribution of Pnoepyga

immaculata is beyond Manali. As a second

example, Gaston et al. (1993) never recorded

Luscinia brunnea in their surveys of the Great

Himalayan National Park, approximately 50 km to

the south of Manali. However, this species is one

of the commonest species at both Manali and

Overa.

At any given locality, there is high species’

turnover along the elevational gradient, and bird

communities separated by 1 ,000 m altitude are

more different than the communities in Manali

and Overa at the same altitude. Hunter and

Yonzon (1993) have recorded the presence of

many species with restricted altitudinal ranges in

Nepal and point to the need for preserves at all

elevations. Despite this, there are large differences

in species’ composition at different localities

along the Himalaya, and Arunachal shares less

than one-quarter of its species with Pakistan.

There is a tendency to treat the Himalaya as a unit

with respect to conservation issues, because the

threat of habitat destruction is universal

throughout the range. This analysis demonstrates

that the Himalaya is actually quite diverse, and

each area has its own needs. Each area must have

its own nature preserves if Himalayan biodiversity

is to be conserved.

Acknowledgements

We thank Dr. P. Khullar, Ms Vijay Laxmi

Tiwari and Mr. V.P. Pathania for logistical support

and A. Bontrager, D. Irwin, K. Marchetti and our

local assistants for help in the field. We thank

Sean Gross, Sanjay Sondhi, Tony Gaston and Pam

Rasmussen for comments or responses to queries.

Some funds for the studies at Manali were

provided by the National Science Foundation

(USA) and the National Geographical Society

(USA).

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Appendix I: Additions/corrections to the Overa Sanctuary list (in Price and Jamdar 1990, 1991)

1. Price and Jamdar (1990, 1991) reported on the birds of Overa Sanctuary based on 3 years’ fieldwork

(1985-1987). Our subsequent work at Overa for 1-2 months in each of the years 1988, 1989, 1990 and

1991, confirmed the altitudinal distributions previously reported and confirmed breeding for some species.

2. One additional species was recorded: large pied wagtail Motacilla maderaspatensis near Overa village.

3. In Price and Jamdar (1990), we stated that two species, Mycerobas icterioides and Carpodacus rhodochrous

bred throughout the entire altitudinal range that we studied (2,430-3,725 m). Although we recorded both

species throughout the range, we are not sure that they breed in all locations. In particular, it remains to be

determined that both the species breed at the highest and lowest altitude sites, and it appears unlikely that

M. icterioides breeds at the highest altitude.

4. In Price and Jamdar (1991), we noted that Phylloscopus reguloides is absent from Overa and were unsure

where P. reguloides and Phylloscopus occipitalis breed in sympatry. We now know they occur sympatrically

from the Kashmir border in the Chamba region of Himachal Pradesh to at least the Nepal border in Kumaon

(including Manali). P reguloides breeds at generally higher altitudes (e.g. Table 1 of this paper).

5. In Price and Jamdar (1 990), the generally uncommon spruce was not separated from fir in Table 1 . In Table

4, printer’s errors resulted in omission of some data and mismatching of columns. The altitudinal distributions

and densities at high elevation of all species are given in Table 1 of this paper, and the mismatched columns

in Price and Jamdar (1990) can be worked out.

Appendix II: Rare or unusual birds recorded in Manali Sanctuary

Tibetan snowcock Tetraogallus tibetanus : A flock of 10 at 3,825 m on June 1, 2001. We do not know

if this species is common, as we rarely visited higher altitudes.

Snow pigeon Columba leuconota : Eight birds seen on May 9, 2001, also seen rarely in Overa.

Black-naped green woodpecker Picus canus. Observed on May 24, 2002 at 2,430 m elevation.

Speckled wood-pigeon Columba hodgsonii. One bird seen on May 19, 2001 and two on May 8, 2002

in lower elevation forest, also seen once at Overa.

Grey treepie Dendrocitta formosae. One bird seen on May 19, 2001 at 3,100 m. Also seen in Manali

town in May, 2002.

Long-tailed thrush Zoothera dixoni. A nest with 3 nestlings discovered at c. 3,100 m on May 23, 2002.

Possibly more common in Manali and was previously overlooked by us.

Gould’s shortwing Brachypteryx stellata. A male singing on June 6, 1994, in a juniper bush at c. 3,500

m. There are only a few earlier observations of this very distinctive species west of central Nepal, and none

west of Uttaranchal. Despite searching, the species was not observed again.

Chaffinch Fringilla coelebs. A male singing on top of a cedar on May 4, 1994, at 2,000 m elevation. A

rare winter visitor to the Himalaya (Grimmett et al. 1999b).

Little forktail, Enicurus scouleri. Observed twice in the sanctuary, once at c. 3,200 m. It is probable that

the species breeds regularly but not confirmed, for watercourses were not regularly checked.

Yellow-browed tit, Sylviparus modestus. A pair feeding 4 young in a moss cup in hole in tree, June 11,

1997 at 2,750 m.

Grey-cheeked flycatcher-warbler, Seicercus poliogenys. A male singing at 2,500 m in a chestnut

grove, first seen May 17, 2001, still present June 26, 2001, and still singing loudly and presumably unmated.

Apart from possible records by Green (1986) in atypical habitat, this species has not been recorded west of

Central Nepal (Grimmett etal. 1999b).

Fire-tailed sunbird, Aethopyga ignicauda. A male was captured and present in the study area on

rhododendron (c. 3,300 m.) throughout June, 1997. It may have bred.

Two species are in Table 3 ( Turdus rubrocanus and Phylloscopus affinis) and are not repeated here.

JOURNAL , BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG. -DEC. 2003

409

BIRD SPECIES DIVERSITY ALONG THE HIMALAYA

Appendix III: Additional species of Manali

Many of the following species are found in Manali town and on agricultural land, but have been recorded along

the lower boundary of the Sanctuary (I = Infrequently observed). Note that this list includes only those species

not in Tables 1 or 2.

Brown-fronted pied woodpecker Dendrocopos auriceps (I)

Small blue kingfisher Alcedo atthis

Oriental cuckoo Cuculus saturatus

Indian cuckoo Cuculus micropterus (I)

Blue rock pigeon Columbia livia

Common sandpiper Actitis hypoleucos

Eurasian hobby Falco subbuteo

Common hoopoe Upupa epops

Rufous-backed shrike Lanius schach

Yellow-billed blue magpie Urocissa flavirostris

Eurasian golden oriole Oriolus oriolus (I)

Asian paradise-flycatcher Terpsiphone paradisi (I)

Brown dipper Cinclus paUasii

Tickell’s thrush Turdus unicolor

Oriental magpie-robin Copsychus saularis

Plumbeous redstart Rhyacomis fuliginosus

Spotted forktail Enicurus maculatus

Grey bush chat Saxicola ferrea

Common stonechat Saxicola torquata (I)

Common myna Acridotheres tristis

Jungle myna Acridotheres fuscus

Great tit Parus major

Red-headed tit Aegithalos concinnus

Himalayan bulbul Pycnonotus leucogenys

Oriental white-eye Zosterops palpebrosus

Brown-flanked bush-warbler Cettia fortipes

Streaked laughingthrush Garrulax lineatus

Rufous sibia Heterophasia capistrata (I)

House sparrow Passer domesticus

Cinnamon tree sparrow Passer rutilans

White wagtail Motacilla alba

Large Pied wagtail Motacilla maderaspatensis

Grey wagtail Motacilla cinerea

Spotted munia Lonchura punctulata (I)

Yellow-breasted greenfinch Carduelis spinoides

Eurasian goldfinch Carduelis carduelis

Common rosefinch Carpodacus erythrinus

Grey-headed bunting Emberiza fucata

Rock bunting Emberiza cia.

Note: Cettia fortipes has been recorded up to 2,225 m and Rhyacornis fuliginosus at 2,700 m. Both these

species are of interest, in that they appear to occur at much higher elevations in Overa than in Manali. Emberiza

cia has been recorded up to 3,200 m in Overa, but only 2,000 m in Manali. Garrulax lineatus has been recorded

410

JOURNAL . BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG. -DEC. 2003

FRUGIVORY, SEED DISPERSAL AND REGENERATION

BY BIRDS IN SOUTH INDIAN FORESTS

( With seven text-figures and one plate )

P. Balasubramanian1’2 and B. Maheswaran1

Key words: Tropical forest, bird-fruits, avian frugivores, seed dispersal, fruit size,

forest regeneration, Western Ghats

Plants, unlike animals, are immobile and have little control over the choice of site where they can live and grow.

Over aeons, plants have perfected the art of survival in spite of this disability by steadily interacting with the

animals in their habitat. The evolution of brilliant, attractive colours and endowment of the fruits with nutritious

supplements such as proteins, lipids and carbohydrates, have appealed to the dietary needs of birds and

mammals. Furthermore, the plant’s ability to asynchronously produce fruits, for the assemblage of vertebrates

dependent on fruits have made their dispersal possible. Plant-bird interactions play a pivotal role in maintaining

the structural and functional integrity of natural ecosystems. Also, the interactions guide the process of evolution

ofbiodiversity as well as ecological communities. On account of their unparalleled richness, tropical communities

are treasure troves of biological interactions between plants and animals. Ecological degradations, as in shrinking

habitats and fragmentation, necessitate a comprehensive understanding of the basic tenets of interacting entities,

in order to evolve effective strategies to preserve and enrich our biodiversity.

Introduction

Birds play a prominent role in pollination

and seed dispersal of plants. The flower visiting

birds, while harvesting nectar, pollinate their food

plants. Similarly, the fruit eating birds by devouring

the pulpy fruits distribute the seeds of their food

plant species. Birds, by virtue of their habits, can

play a more effective role in seed dissemination

than other animals. Compared to other vertebrates,

which disseminate the seeds in clumps, the seed

deposition pattern of birds is more efficient. The

interactions between flower-birds and bird-flowers

have been discussed (Ali 1931, Subramanya and

Radhamani 1993). A variety of birds and mammals

depend predominantly on fruit to fulfil their

nutritional requirements. Such fruit-eating birds

and mammals are termed as frugivores. Fruit traits

such as colour, size and nutritional supplements

have evolved in response to interacting

vertebrates. The evolution of fruit syndromes,

complemented by the birds’ adaptive ability to

recognise these syndromes, along with

‘Salim Ali Centre for Ornithology and Natural History

(SACON),

Coimbatore 641 108, Tamil Nadu, India.

2Email: [email protected]

physiological and anatomical modifications to

process the fruit diet have been the key in the

contemporaneous plant-bird interactive relations.

Such persistent relationships have given rise to

mutual dependence, benefiting both elements.

Though fruit-frugivore interactions have

been studied by various authors during recent

times, there is no consolidated information on the

role of birds in seed dispersal and forest

regeneration. Information is available on the fruit

types consumed by various bird species in India,

but only a few studies attempted to quantify the

role of birds in forest regeneration. A review of

those studies and reference to the various floras

indicate that several families benefit from birds

for their regeneration. Known syndromes of bird-

dispersed fruits help to identify various bird-fruits

from hitherto unexplored bird habitats.

Frugivores help seeds escape from the

deleterious effects of seed and seedling predators

(Janzen 1970). Differences in frugivore activity

have profound effects on the conditions under

which seeds and seedlings must survive, and

consequently should influence the evolution of

tree demographies. The relationship between seed

dispersal and seedling demography has profound

implications for tropical conservation policy

411

JOURNAL . BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG. -DEC. 2003

FRUGIVORY IN SOUTH INDIAN FORESTS

(Howe 1984). Seasonality in fruit production in

tropical forest ecosystems has brought to light

the roles played by ‘keystone’ or ‘pivotal’ species

during periods of scarcity.

Studies on fruit-frugivore interactions

conducted by us as part of research programmes

under the Bombay Natural History Society

(BNHS) and the Salim Ali Centre for Ornithology

and Natural History (SACON) for the past two

decades helped us to arrive at certain conclusions

and also comprehensively list the plant genera

dispersed by birds in south India. The material

for this paper has been derived mainly from the

Tropical Dry Evergreen Forests of Point Calimere,

and the Semi-evergreen, Dry and Mixed

Deciduous Forests of the Western Ghats. Though

seed dispersal is effected by exozoochorous

methods (dispersal effected by seeds clinging on

to body parts of birds and animals) also, we would

like to state that the review exclusively deals with

endozoochory (dispersal wherein fruits are

ingested, the pulp digested and the seeds

defecated or regurgitated).

Observations

Frugivory and seed dispersal in south India

The present review is based on three major

research efforts in south India - on the Tropical

Dry Evergreen Forests in Point Calimere, Dry

Mixed Deciduous Forests of Coimbatore Division

and Semi-evergreen forests of Mudumalai Wildlife

Sanctuary (MWLS), Western Ghats. The lists of

fruit-eating birds and bird-dispersed fruit species

have been compiled chiefly from these three major

studies and have been summarised in Appendix I

and II.

A total of 66 species (36 genera) of Indian

birds from 16 families have been documented for

frugivory in south India. Bird species of families

Columbidae (pigeons), Pycnonotidae (bulbuls)

and Muscicapidae (flycatchers) contributed the

maximum number of frugivorous species (7

species). While Sturnidae (mynas) hold 6

frugivorous species, Psittacidae (parrots) and

Capitonidae (barbets; Plate 1 , Fig. 1 ) hold 5 species

each. Among the 66 species of fruit-eating birds,

members of Columbidae (except the mountain

imperial-pigeon) and Psittacidae (parrots) digest

the seeds and hence are considered as seed

predators. Other frugivores could be considered

as legitimate seed dispersers.

The body size varied considerably among

frugivorous birds (range 5-2,500 g; Appendix I;

Fig. 1). Hombills (Family Bucerotidae) were the

heaviest birds, having a mean body weight of

1 ,246 g (n=3) followed by Family Corvidae (crows)

and Columbidae (pigeons) whose mean body

weight was 288 g (n=4) and 246 g (n=7)

respectively. The great pied hornbill Buceros

bicornis, a charismatic flagship species of the

Western Ghats and northeastern India, is the

heaviest among frugivores, weighing 2,500 g

followed by Malabar pied hornbill Anthracoceros

coronatus (1,000 g), jungle crow Corvus

macrorhynchos (650 g) and the mountain imperial

-pigeon Ducula badia (580 g). Hornbills and

pigeons are also voracious frugivorous bird

species in their habitat.

35

0-50 50-100 100- 150- >200

150 200

Body mass (g)

Fig 1 : Body mass of frugivorous birds in south

Indian forests (n=66)

Bird attracting flora in south India

Fruits of 219 species (141 genera) belong-

ing to 61 plant families were recorded to be used

by frugivorous birds. The members of the

families Lauraceae (21 species), Euphorbiaceae

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JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003

FRUGIVORY IN SOUTH INDIAN FORESTS

Balasubramanian, P. et al Species involved in frugivory in Western Ghats

Plate 1

Fig. 1 : Coppersmith barbet Megalaima haemacephala, a frugivore and major seed

disperser of forest trees, at Coimbatore Forest Division

Fig. 2: Ficus benghalensis, a keystone species for avian frugivores, in the Coimbatore

Forest Division

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413

P. BALASUBRAMANIAN K. MARUDHACHALAM

FRUGIVORY IN SOUTH INDIAN FORESTS

120

Berry Drupe Capsule Syconium Others

Fruit Types

Fig. 2: Morphological classification of bird-dispersed

fruits in south Indian forests (n= 219)

(13 species), Rubiaceae (12 species), Moraceae

(10 species), Meliaceae and Annonaceae (9

species each) were most represented among

fleshy-ffuited species adapted for ffugivory. The

genus Ficus (figs), known for their ability to

produce fruits asynchronously and support the

ffugivore community during periods of scarcity,

was represented by 8 species. The genera

Eugenia and Litsea were represented by 5 species

each, while Capparis , Cinnamomum , Diospyros

and Zizyphus were represented by 4 species each

(Appendix II).

The fruit types of all the 219 species were

determined. Berries were the predominant fruit

type borne by 99 species (45%) while drupes were

borne on 76 species (35%) and together

accounted for 80% of the fruits for the whole of

south India (Fig. 2). Fleshy-ffuited plants are very

common in tropical forests, with frequencies

usually over 70% (of woody plants) in the forests

and lesser in the dry forests. The frequency of

fleshy-ffuited plants in the Dry Deciduous Forests

of Coimbatore was 65%, while in Tropical Dry

Evergreen Forests of Point Calimere it was 73%.

In the Semi-evergreen Forest of Mudumalai

Wildlife Sanctuary, Western Ghats, 57% of the

species and 74% of individuals were fleshy-ffuited

species.

Colours of ripe fruits for all the 2 1 9 species

were assigned to one of eight broad colour

categories as used by Wheelwright and Janson

(1985). The analysis showed that 94 species (43%)

produced black fruit while 58 species (26%)

produced red fruit and 39 species ( 1 8%) produced

yellow fruit (Fig. 3). Black and red accounted for

70% of species.

a)

a>

o

a)

a.

</)

Colour

Fig. 3: Colours of bird-dispersed fruits in south Indian forests (n=219)

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG. -DEC. 2003

415

FRUG1VORY IN SOUTH INDIAN FORESTS

Analyses of the fruit diameter revealed that

98 species (45%) had their fruit sizes 10-20 mm,

while another 85 species (39%) bore fruits in the

size range <10 mm. Only 36 of the 219 species

( 1 6%) bore fruits of size greater than 20 mm (Fig. 4).

About 83% of fruits were less than 20 mm in

diameter. Plants of Families Annonaceae,

Burseraceae, Elaeocarpaceae, Meliaceae,

Moraceae and Myristicaceae produced large fruits

while plants of Families Cucurbitaceae,

Cordiaceae, Ebenaceae, Lauraceae,

Melastomataceae, Menispermaceae, Oleaceae

and Rubiaceae produced small fruits.

120 i

<10 10-20 20-30 >30

Size class (cm)

Fig. 4: Fruit size classification of bird-dispersed

fruits in south Indian forests (n=219)

Birds and regeneration in Tropical Dry

Evergreen Forest, south India

A study on the interrelationships between

fleshy-fruited plants and their vertebrate

consumers was conducted in the Dry Evergreen

Forest in Point Calimere, south India in the mid-

1980s (Balasubramanian 1996). During the course

of the study, 3 1 7 species of flowering plants were

recorded, of which 88 bore fleshy fruits. Sixty-

four of the 88 fleshy-fruited species were adapted

for frugivory by birds. A total of 20 bird species

of 14 genera from 1 0 families were observed to eat

fruits at Point Calimere. The major avian seed

dispersers documented were the white-browed

bulbul Pycnonotus luteolus , red-vented bulbul

Pycnonotus cafer , jungle myna Acridotheres

fuscus , brahminy starling Sturnus pagodarum ,

jungle crow Corvus macrorhynchos, house crow

Corvus splendens and Asian koel Eudynamys

scolopacea . Other seed dispersers included rosy

starling Sturnus roseus, Eurasian golden oriole

Oriolus oriolus, Indian tree pie Dendrocitta

vagabunda, jungle babbler Turdoides striatus

and Tickell’s flowerpecker Dicaeum

erythrorhynchos . Pycnonotus luteolus visited

maximum number of plants (63), followed by

Pycnonotus cafer (51), Eudynamys scolopacea

(32) and Acridotheres tristis (27). Three plant

species were visited by more than 1 0 bird species.

Salvadora persica was visited by maximum

number of bird species (15), followed by

Manilkara hexandra (12) and Ficus infectoria

01).

Even though nearly 50% of the fleshy-

fruited species in Point Calimere were consumed

by mammals, only 1 0% of the species were solely

dispersed by mammals. Most plant species are

chiefly dispersed by birds.

Birds and natural regeneration in Dry Mixed

Deciduous Forest, Western Ghats

In an effort to understand the tenets of

restoration ecology with birds as the basis of

regeneration, a project was launched by SACON

in cooperation with the Tamil Nadu Forest

Department. The studies were conducted in the

Dry Mixed Deciduous Forest, Coimbatore Forest

Division, Western Ghats (Balasubramanian et al.

1998). Of the 1 15 woody species recorded in the

study area, 52% of the species were bird-

dispersed. Figs attracted maximum number of bird

species followed by Celtis philippensis and

Syzygium cumini . Of the 35 fruit-eating bird

species, 1 8 (5 1 .42%) were major frugivores. The

red-vented, white-browed and red-whiskered

bulbuls were most frequent visitors to fruit trees

in the Dry Deciduous Forest. In the riverine forest,

barbets, bulbuls, mynas, the Asian fairy bluebird

416

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG. -DEC. 2003

FRUGJVORY IN SOUTH INDIAN FORESTS

and koel were major frugivores. The Malabar pied

hornbill, one of the largest frugivores of the Indian

subcontinent, were seen to feed on fruits of Ficus

sp., Strychnos nux-vomica and several other large-

fruited species in the riverine forest

(Balasubramanian and Saravanan 2001).

Five species, namely Ficus benghalensis ,

F. racemosa , Celtis philippensis , Cassine glauca

and Strychnos potatorum , were experimented with

to determine if the birds enhanced germination

capacity of seeds that passed through their

intestinal tracts. The germination percentage of

seeds dispersed by birds was significantly higher

than that of the seeds collected from plants for

four species. In one species Celtis philippensis ,

although a similar trend was observed, the

difference was not significant (Fig. 5).

Natural regeneration of bird-dispersed

plants was studied in the Dry Mixed Deciduous

Forests of Coimbatore Division in Tamil Nadu.

The density of seedlings and saplings of bird-

dispersed trees varied from 279 per hectare in the

highly degraded habitat to 640 in the undisturbed

habitat. Twelve out of the 1 5 species had good

regeneration potential, represented by more than

10 saplings. In the highly degraded habitat, only

3 out of 9 species had more than 10 saplings.

Hornbills and forest regeneration in Semi-

evergreen and Evergreen Forests

Malabar grey hornbill ( Ocyceros griseus ),

an endemic frugivore of restricted range in the

moist forests of Western Ghats, south India, is

dependent on fruit resources and suitable nest

trees for subsistence. As a consequence of its

specialised breeding habits and dependence on

fruits as a predominant source of food, this

hornbill is known to interact with a variety of tree

species. The Malabar grey is considered one of

the keystone species and a major seed disperser

in the Western Ghats region (Mudappa 2000).

Like most other hornbills, the Malabar grey

is known to bring a large load of fruit to the nesting

site to feed the incarcerated female and chicks

during the breeding season. Seeds of fruits

consumed are squirted out through the nest slit,

and get embedded in the litter-strewn forest floor.

These seed deposits are usually termed as midden.

60

50

Bird defecated □ Fresh

c

o

•*—*

OJ

c

40

E 30

<u

O

20

10

0

Ficus Ficus

benghalensis racemosa

Celtis

philippensis

Cassine

glauca

Strychnos

potatorum

Species

Fig. 5: Percentage germination of bird-defecated and fresh seeds, Coimbatore Forest Division

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 10Q(2&3), AUG. -DEC. 2003

417

FRUGIVORY IN SOUTH INDIAN FORESTS

Monitoring the midden deposits and analysing

the seeds therein during breeding season, helped

to detennine the variety of fruit species consumed

and dispersed by this hombill. The study was

conducted in the Semi-evergreen Forests of

Mudumalai Wildlife Sanctuary (Balasubramanian

and Maheswaran 2001). Tree species common to

this vegetation type are Olea dioica,

Actinodaphne malabarica , Per sea macrantha ,

Cinnamomum verum, Lagerstroemia microcarpa ,

and Terminalia sp.

Of the 1 ,430 individuals belonging to the 70

tree species present, 74% of individuals and 56%

of the species were adapted for frugivory. Major

disperser species in this habitat, apart from the

Malabar grey hombill ( Ocyceros griseus ), were

the fruit pigeons ( Ducula badia), barbets

( Megalaima sp.), mynas (Acridotheres sp.) and

starlings (, Sturnus sp.). During the breeding

season, an intensive nest search was conducted

in the semi-evergreen patches of Mudumalai

Wildlife Sanctuary for nesting activity of the

Malabar grey hornbill. Cavities of trees used by

hombills for nesting were located by following

breeding pairs of hombills, prior to the breeding

season. Twelve nests in the breeding season were

selected for studying midden regeneration. Two

square plots (5 sq. m each) were demarcated for

each nest site, one behind and one in front of the

nest. During the post-breeding season (May-

January), these sites were visited every week to

study regeneration from the midden deposits.

New seedlings of a total of 19 species

belonging to 1 3 families were enumerated from the

square plots. Olea dioica , one of the favoured fruits

of this birds was recorded in all the 1 2 midden sites,

while Cinnamomum verum was recorded from 1 1

midden sites. Lauraceae and Flacourtiaceae (3

species each) were the most represented families

among the regenerated seedlings. The number of

seedlings regenerated in front of the nest was

significantly higher than those behind the nest (X2

= 298.378; df = 9; P < 0.005). While 13 species

comprising 280 regenerated seedlings were recorded

behind the nests, 18 species comprising 761

seedlings had regenerated in front of the nest site,

just below the nest hole, i.e. 63% more seedlings

regenerated in front of the nest. Greater regeneration

in front of the nest was seen at all the sites except

nest site 9 (Fig, 6). Regeneration within a species

was also found to be significantly higher in front of

the nest (X2= 2 1 3.823; df = 8; P < 0.005).

120

100

80

60

40

20

0

co

O)

c

0

c n

1 2 3 4 5 6 7 8 9 10 11 12

Nest sites

Fig. 6: Regeneration of seeds behind and in front of Malabar grey hombill nests in a Semi-evergreen Forest

at Mudumalai Wildlife Sanctuary, Western Ghats

418

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG. -DEC. 2003

FRUG1V0RY IN SOUTH INDIAN FORESTS

Germination experiments were conducted

at the study site using seeds collected from the

midden, and control seeds of ripe fruits collected

directly from trees for comparison. Seeds of two

preferred fruit species, Olea dioica and Persea

macrantha, were chosen for seed germination

studies. The soil conditions in which the

experimental seeds were planted were the same

as they would be in the midden. Seeds were sown

in polythene bags under open conditions and

watered regularly.

After four months of continuous

monitoring of the seedlings, it was observed that

bird-defecated seeds had a higher regeneration

potential than control seeds collected directly

from plants. Bird-defecated seeds of Olea dioica

showed 4 1 .67% higher germination rate than fresh

seeds while for Persea macrantha , the bird-

defecated seeds showed 75% enhanced

germination over the control seeds (Fig. 7). A

statistical comparison of the bird-defecated and

control seeds for both the species showed a

significantly higher rate of germination for

bird-defecated seeds (X2 = 90.739; df =1;

P< 0.005).

Species

Fig. 7: Percentage germination of seeds defecated by

Malabar grey hornbill and control seeds, in two plant

species at Mudumalai Wildlife Sanctuary,

Western Ghats

Bird-aided regeneration in other forests of the

Western Ghats and coastal plains

Hombills are one of the dominant frugivores

and seed dispersers of the palaeotropical region.

In the Asian tropical forests, they have been well

studied for frugivory, especially in the breeding

season, and are known to disperse the seeds

undamaged (Leighton 1981). Hornbills, for most

part of the year, are scatter dispersers, but during

the nesting season large numbers of seeds are

deposited under the nests, creating conspicuous

middens. Our study in the Semi-evergreen

Forests of Mudumalai Wildlife Sanctuary, Western

Ghats has established that the Malabar grey

hornbill utilises 27 different fleshy-fruit species,

and 16 species were recorded to have regene-

rated from the midden under nest trees of

hombills. Kannan and James ( 1 999) identified 2 1

species of trees with lipid-rich fruits, utilised by

hornbills ( Buceros bicornis and Ocyceros

griseus). These included 12 Cinnamomum,

2 Myristica, 2 Beilschmedia, 2 Dysoxylum ,

1 Persea and 2 Polyalthia species.

In a study in the Moist Deciduous Forest

habitat in the Western Ghats of Kerala

(Vijayakumar and Zacharias, in press) showed that

the major frugivores were the white-cheeked

barbet Megalaima viridis , crimson-throated

barbet Megalaima rubricapilla , pompadour

green-pigeon Treron pompadora , black bulbul

Hypsipetes leucocephalus and common hill-myna

Gracula religiosa.

In the Wet Evergreen Forest of Kalakad-

Mundanthurai Tiger Reserve, southern India,

birds dispersed 59% of the 65 fleshy-fruited tree

species (Ganesh and Davidar 2001). About 33%

( 1 6) of bird-dispersed species belonged to Family

Lauraceae. While small birds dispersed 86% of

the plant species, pigeons dispersed 14% of large

fruited species. The six common frugivorous

species of that habitat were black bulbul

Hypsipetes leucocephalus , yellow-browed bulbul

Iole indica , red-whiskered bulbul Pycnonotus

jocosus, white-cheeked barbet Megalaima viridis ,

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419

FRUG1VORY IN SOUTH INDIAN FORESTS

mountain imperial-pigeon Ducula badia and

Nilgiri wood-pigeon Columba elphinstonii.

Effects of hornbills on seeds have been

documented for 34 species in Dja Reserve,

Cameroon (Whitney et al. 1998). Seeds of most

species were egested without much physical

damage. Germination trials on 24 species showed

that 23 species germinated after egestion by two

hornbills, Ceratogymna atrata and

C. cylindricus, and a major proportion of these

species did not exhibit a fall from the germination

rate in control seeds.

A study on plant-bird interaction in the

Tropical Evergreen Forest of the erstwhile

Chengalpattu district, Coromandel coast,

southern India, showed that 130 out of the 180

woody plants (72%) bore fleshy fruits

(Narasimhan et al. 1993). Major fleshy-fruited

species included Zizyphus oenoplia, Allophyllus

serratus, Ficus benghalensis, Benkara

malabarica, Hugonia mystax, Tarenna asiatica,

Memecylon umbellatum, Carmona retusa and

Dendrophthoe falcata. Though 150 birds have

been recorded from this region (Siromoney 1971),

only 20 are involved in seed dispersal. Families of

frugivorous birds recorded include Columbidae,

Cuculidae, Capitonidae, Oriolidae, Sturnidae,

Corvidae, Irenidae, Pycnonotidae and

Muscicapidae. In this forest, berries and drupes

were the major fruit types encountered, while red

(48%) followed by black (31%) were the

predominant fruit colours. The study also showed

that 72% of the fruits were less than 1 cm in

diameter, indicating the role of generalist

frugivores in dispersal.

Bulbuls have established themselves as one

of the dominant frugivores and seed dispersers

in scrub jungles or forests of secondary

vegetation in Asia, Africa and the Middle East

(Lever 1987). In the Tropical Dry Evergreen, as

well as Dry Mixed Deciduous Forests of south

India, bulbuls are the predominant frugivorous

and seed dispersing species (Vijayan 1975,

Balasubramanian 1996, Balasubramanian et al.

1998). In Point Calimere (Balasubramanian 1996),

74% of the visits by birds on fruiting plants were

by two species of bulbuls. In the Dry Deciduous

Forests of Coimbatore (Balasubramanian et al.

1998), the largest number of visits (47.5%) were

by 3 species of bulbuls followed by 3 species of

barbets (18.46%). In a study in the subtropical

shrub lands of Hong Kong, two species of

bulbuls, the red-whiskered bulbul {Pycnonotus

jocosus ) and the light-vented bulbul {Pycnonotus

sinensis ), were the most important seed dispersal

agents. These species have an unusually large

gape width for their size, and they utilise this

advantage to the maximum (Corlett 1998). The

dominant role of bulbuls in frugivory and dispersal

in tropical forest communities makes them a

valuable group of birds.

Research on plant-bird interactions at Point

Calimere showed that conspicuous colour

combined with small size of fruit enabled

frugivorous birds to access fruit resources easily

(Balasubramanian 1996). According to Terborgh

and Diamond (1970), small-fruited plant species

attract more species of birds than large-fruited

ones. Corlett (1996) observed that most of the

bird-dispersed seeds in Hong Kong had a mean

diameter of <13 mm. This trend was consistent

with findings from the Neotropics (Janson 1983,

Howe 1986), Africa (Knight and Siegfried 1983,

Gautier-Hion et al. 1985), Australasia (Beehler

1983, Green 1993) and in Indonesia (Suryadi etal.

1994).

Birds and enhancement of seed germination and

regeneration

Frugivores affect the germination success

of seeds which they either defecate or regurgitate,

as the gastrointestinal enzymes and acids within

the gut of the birds soften the hard seed coat,

thus breaking dormancy in seeds (Fleming and

Heithaus 1981). Our study has clearly established

the role played by birds in enhancement of seed

germination in Dry Mixed Deciduous and Semi-

evergreen Forests of the Western Ghats. Seeds

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FRUGIVORY IN SOUTH INDIAN FORESTS

of Olea dioica , Persea macrantha (both in Semi-

evergreen Forest), Ficus benghalensis,

F. racemosa , Celtis philippensis, Cassine glauca

and Strychnos potatorum (Dry Mixed Deciduous

Forest) showed significant enhancement in

germination after passing through the guts of

birds. Similar results have been reported by Midya

and Brahmachary ( 1 99 1 ) for Ficus benghalensis

(58% germination in bird-dispersed seeds against

34% in control seeds), and by Mishra et al. (1987)

for Azadirachta indica {16% germination in bird-

dispersed seeds against 42% in control seeds).

In South Australia, Yan (1993) observed more

than 90% germination of mistletoe (Amyema

preissii and Lysiama excocarpi) seeds dispersed

by birds.

Though ficuses occur in low abundance,

they play a very prominent role in sustaining

frugivorous populations in tropical forests. Being

aseasonal in fruit production, figs have evolved

as one of the most reliable diet resources for the

vertebrate community. The role of plant taxa that

fruit in times of scarcity, in maintaining the

population of the frugivore community was first

propounded by Terborgh (1986), followed by

Lambert and Marshall (1991). According to them,

the abundance of fruiting in figs outside the peak

fruiting season of the community, is the single

factor that supports frugivorous communities in

tropical forests. Due to this key role played by

such taxa in the maintenance of community

structure, they have been christened as ‘keystone

species.’ At least 25 species of figs in the

southern Western Ghats (Plate 1, Fig. 2) support

the frugivorous bird community there, which in

turn enhances germination and regeneration of

bird-dispersed plant species. This augurs well

for the sustenance of the forest ecosystem in the

region.

Conclusion

Birds help determine the structure of tropical

forests, as a major proportion of plant species are

consumed and dispersed by them. As seen in

this study, 73.7% of species in Tropical Dry

Evergreen Forests of Point Calimere, 50% of

species in Dry Mixed Deciduous Forests of

Coimbatore Forest Division, Western Ghats, and

55.7% of species in Semi-evergreen Forest in

Mudumalai Wildlife Sanctuary are bird-dispersed.

Dispersion of seeds by birds was also seen in 35-

70% of woody plants in Australian Wet Tropical

and Dry Tropical Forests, and 47-54% of woody

plants in North American Deciduous Forest and

49-66% in Mediterranean scrublands.

Furthermore, seeds egested by birds have

improved germination and regeneration vigour,

thus increasing the dependence of plants on birds

as a reliable and effective mode of dispersal. Thus

birds have a pivotal role in shaping the structure

of the forest. Plant communities in the present era

have survived on account of their ability to interact

mutualistically with birds.

Seed dispersal and regeneration are two

processes that are absolutely indispensable to

the maintenance of tropical forests (Galetti 2000,

Herrera 1984, 1995, Howe 1984, 1990, Stiles 1985,

1992, Willson 1992). In spite of their fundamental

importance in sustenance of tropical forests, very

little is known about the role played by dispersers

and the mechanism of dispersal. Research

initiatives have been few and far between in the

tropics and are mostly restricted to the Neotropics

(Howe 1983, 1985, 1986, Howe and Vande

Kerckhove 1981). Far greater efforts are needed

to show the effectiveness of frugivores in

dispersal and regeneration.

Though information is available for a few

species, community-wide assessments of

dispersers are negligible. Documentation of

frugivory is only the first step in understanding

the complex and manifold interactions between

birds and plants. Being mobile links in tropical

forest ecosystems, frugivores deserve greater

attention, especially on the intricacies of their role

in enhancement of germination and forest

regeneration.

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F RUG IVORY IN SOUTH INDIAN FORESTS

Acknowledgements

The senior author (P.B.) expresses his

gratitude to Mr. J.C. Daniel, Honorary Secretary,

Bombay Natural History for encouragement. He

also thanks Dr. G. Kumaravelu, Conservator of

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Appendix I: Frugivorous birds of south Indian forests

424 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, I00(2&3), AUG.-DEC. 2003

FRUGIVORY IN SOUTH INDIAN FORESTS

Appendix I: Frugivorous birds of south Indian forests ( contd .)

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FRUGIVORY IN SOUTH INDIAN FORESTS

Appendix II: Bird-dispersed plant species in south Indian forests ( contd .)

426

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FRUGIVORY IN SOUTH INDIAN FORESTS

Appendix II: Bird-dispersed plant species in south Indian forests ( contd .)

JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2G03

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FRUGIVORY IN SOUTH INDIAN FORESTS

Appendix II: Bird-dispersed plant species in south Indian forests ( contd .)

428

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FRUGIVORY IN SOUTH INDIAN FORESTS

Appendix II: Bird-dispersed plant species in south Indian forests ( contd .)

JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG. -DEC. 2003

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FRUGJVORY IN SOUTH INDIAN FORESTS

Appendix II: Bird-dispersed plant species in south Indian forests ( contd .)

430

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FRUGIVORY IN SOUTH INDIAN FORESTS

Appendix II: Bird-dispersed plant species in south Indian forests ( contd .)

Sources: Balasubramanian 1996, Balasubramanian and Maheswaran 2001, Balasubramanian et al. 1998,

Ganesh and Davidar 2001, Kannan and James 1999.

m n ■

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CROCODILE CONSERVATION, WESTERN ASIA REGION: AN UPDATE

( With three text-figures and two plates )

Romulus Whitaker1 and Harry V. Andrews2

Key words: Crocodiles, conservation, South Asia, gharial, mugger, saltwater crocodile

Crocodile conservation began in earnest in the mid 1 970s when all the three South Asian species —

the gharial (Gavialis gangeticus), mugger (Crocodylus palustris) and saltwater crocodile ( Crocodylus

porosus ) were near extinction. The Government of India assisted by FAO/UNDP eventually

established 1 3 sanctuaries for crocodile conservation in India and thousands of gharial, mugger and

saltwater crocodiles, reared from wild collected eggs, were released in the wild. Unfortunately, the

energy and enthusiasm for crocodile conservation has all but vanished. Sri Lanka has more wild

mugger than the rest of South Asia combined and a small mugger population survives in Iran.

Pakistan and Nepal have very few mugger and gharial left, while in Bhutan they are all but extinct.

The Madras Crocodile Bank houses the largest gene pool of mugger as well as breeding groups of

gharial and saltwater crocodiles.

Conservation of crocodiles in South Asia through sustainable use is seen as the strategy that could

save the three species in the long run, since protection on paper does not appear to be working.

Introduction

At the best of times in human history,

crocodiles have not been very popular. True, they

appear in myth and religion: Ma Ganga, the Ganga

river goddess, rides a mugger and the crocodile

monster is Makara, the carrier of Varuna the rain

god. Yet, then as now, the crocodile was feared,

reviled and killed when possible — and killed they

were — throughout the Indian subcontinent, the

Andaman and Nicobar Islands and Sri Lanka.

Some of us did surveys in the early 1970s and

confirmed reports from even the remotest areas

that India’s three crocodilians were indeed going

extinct, due to hunting and habitat loss (Biswas

1970, Whitaker 1974a, b, Whitaker and Daniel 1980).

In the mid 1 970s, two initiatives were made toward

saving the gharial Gavialis gangeticus , the

mugger Crocodylus palustris and the saltwater

crocodile Crocodylus porosus. One was the large-

scale GOI/FAO/UNDP Crocodile Conservation

1 P.O. Box 21, Chengalpattu 603 001,

Chennai, Tamil Nadu, India.

Email: [email protected]

2 P.O. Bag 4, Mamallapuram 603 104,

Chennai, Tamil Nadu, India.

Email: [email protected]

Project that covered all crocodilian states in India,

and the other was a private trust, the Madras

Crocodile Bank.

Thanks to the work, interest and

enthusiasm of an array of people from skilled

boatmen to unfortunate bureaucrats behind desks,

from keen young students and forest officers to

overseas consultants and Ph.D. scholars, India’s

three spectacular crocodilians were yanked back

virtually from the brink. This was one of the

world’s conservation success stories. Now it is

time to evaluate what the Project achieved and

why it came to a grinding halt.

The Indian Crocodile Conservation Project

In 1974, a consultant from the Food and

Agriculture Organization (FAO) of the United

Nations Development Programme (UNDP), H.R.

Bustard was invited by the Government of India

(GOI) to look at the crocodile situation in India

(Bustard 1974). Backed by survey reports by Indian

workers, Bustard drafted an FAO project document

for the GOI, that spelled out an egg collection, rear

and release programme with a plan to establish

protected areas at suitable locations. The project

included research, training and even involvement

432

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CROCODILE CONSERVATION

of the local people to protect their interests and to

include them in village level commercial crocodile

farming and ranching (Singh 1999a).

Since the project began more than 25 years

ago, thousands of gharial, mugger and salties have

been reared, mainly from wild collected eggs, at

special rearing centres. As a result of the GOI/

FAO/UNDP Project, 13 sanctuaries totalling over

8,000 sq. km were created for crocodile conserva-

tion in India. Twenty-one other protected areas

provide incidental protection to remnant natural

populations and restocked crocodiles. Over nine

thousand gharial, mugger and saltwater crocodiles

have been released at more than 30 sites

throughout the country.

Results of the releases have been mixed.

For example, most of the 142 mugger released in

Krishnagiri Reservoir, Tamil Nadu in 1985 were

killed off within 2 years (V.M. Narasimhan, pers.

comm.). Of the 150 mugger released in the

Anamalai Wildlife Sanctuary, only three animals

were traceable by 1992 (Choudhury 1986,

Satheesh 1992, Andrews 1999b). The 700 juvenile

gharial released in the Satkosia Gorge Sanctuary

on Mahanadi river in Orissa mostly vanished

(Singh 1991, 1999b). The saltwater crocodiles

released in Andhra Pradesh during 1978 also

vanished due to anthropogenic pressures

(Srinivas et al. 1999). On the other hand, gharial

releases in the Chambal river and saltwater

crocodile releases in Bhitarkanika have been

successful.

Gharial

This unique species, once almost on the

verge of extinction, has made a major comeback

since the 1980s. Whitaker and Basu (1979)

estimated less than a hundred gharial ( Gavialis

gangeticus) in the wild. Since then, of the 4,330

gharial reared at Kukrail centres in Uttar Pradesh

state, 3,495 were released (Singh et al. 1999). In

Madhya Pradesh, 250 have been released since

1 985 into the Chambal, Ken and Son river systems.

During the 1995-1997 surveys of the National

Chambal Sanctuary (Rajasthan, Madhya Pradesh

and Uttar Pradesh) 1,214 gharial were counted,

with an adult (Plate 1 , Fig. 1 ) male to female ratio

of 1:3.6. A total of 75 nests were also located, all

of which indicates that the gharial population is

recovering (Rao et al. 1995, Rao 1999, Sharma

1999). The current gharial status is shown in

Table 1. Next to the Chambal Sanctuary,

Katemiaghat Sanctuary, also in Uttar Pradesh,

has the second largest wild gharial population.

However, this population is currently facing the

threat of extinction due to habitat pressure and

inadequate recruitment. The remaining gharial

sites in India have 50 or fewer animals.

In particular, the new Indian state of

Uttaranchal, along with Rajasthan and Madhya

Pradesh hold the world responsibility of the

welfare of the largest wild gharial population. The

Chambal river is the last bastion of the gharial

and a wide array of river life. Any designs to

Table 1: Total wild gharial in India

Source: ENVIS (Wildlife & Protected Areas) (2)1, 1999.

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tamper with this river system for so-called

“development” and “betterment of the people”

must be thwarted.

In northeast India, gharial are near

extinction, with an estimated population of 10

individuals (Choudhury 1 997). The same author

has reported breeding in isolated areas, besides

individual specimen sightings in Siang, Subansiri

and the Noa-Dihing rivers, tributaries of the

Brahmaputra. There is an urgent need for

conservation and management action in this

state. Surplus gharial in captivity from the states

of Uttar Pradesh or Orissa can be transported

and released in protected areas such as D’Ering

Memorial Sanctuary and Namdapha National

Park, and an efficient monitoring system set in

motion.

The West Bengal Forest Department has

reintroduced gharial into Thorsa and Teesta rivers

on an experimental basis, encouraged by historical

accounts of the presence of this species in the

state (Srinivas et al. 1999). The historical and

current distributional range of the gharial is shown

in Fig. 1.

Gharial outside India

In Nepal, gharial were once distributed

throughout the width of the country in all the

major river systems and their tributaries. Currently

there are only small, isolated populations in the

Kamali, Babai, and the Narayani river systems, all

of which are in, or adjacent to, protected areas

(Maskey and Mishra 1981, Andrews and

McEachem 1994, Maskey 1999). Since 1981,432

gharial have been released and 70 more in 1993

(Andrews and McEachem 1994, Maskey 1999).

The current estimated population in the wild is

105 individuals distributed in seven rivers

(Maskey 1999). Conservation efforts started in

1 978 are still under way through an egg collection,

rear and release programme.

The gharial populations in Pakistan,

Bhutan, and Bangladesh are near extinct and

probably not viable. Estimated populations are

Fig. 1: Historical range and current distribution of the

gharial within the Indian subcontinent

1 0- 1 5 in Pakistan, around 5 in Bhutan and 5- 1 0 in

Bangladesh. The gharial is extinct in Myanmar;

one was shot on the Maingtha river in ‘Upper

Burma’ in 1929 (Smith 1931).

Mugger

The mugger (Plate 1, Fig. 2) Crocodylus

palustris is the most adaptable of the three Indian

crocodilians and has been encountered up to

400 m in crystal clear hill streams, sewage treatment

ponds, cold deep rivers in the Himalayan foothills,

oxbow lakes in Nepal and saltpans in southern Sri

Lanka. The current and historical range of the

mugger is shown in Fig. 2. Surveys in the early

1970s identified Gujarat and Tamil Nadu (two of

India’s drier states) as having most of India’s mugger

(Whitaker 1 987). The present estimated wild mugger

population in Tamil Nadu is 465, of which 52% are

adults (Andrews 1999b). The reported population

for Gujarat is 429 individuals with 88% adults (Vijaya

Kumar etal. 1999).

Tamil Nadu and Gujarat are the most

completely surveyed states for mugger and a look

at the comparative data on croc numbers for the

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CROCODILE CONSERVATION

Whitaker, Romulus and Harry V. Andrews: South Asian crocodiles

Plate 1

Fig. 1 : Male gharial Gavialis gangeticus showing the ‘ghara' of a mature animal

Fig. 2: Mugger Crocodylus palustris hatching

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CROCODILE CONSERVATION

Whitaker, Romulus and Harry V. Andrews: South Asian crocodiles

Plate 2

Fig. 3: Saltwater crocodile Crocodylus porosus nesting

.art Vf’di

Fig. 4: In India, most crocodiles are in captivity

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Fig. 2: Historical range and current distribution of the

mugger within the Indian subcontinent

1 970s and the 1 990s reveals some definite trends.

In general, mugger numbers have increased, due

to restocking and protection, and in cases like

the Moyar river in Tamil Nadu, simply by doing

more complete surveys (Vijaya Kumar and Vyas

1997, Andrews 1999b).

There are fairly stable small mugger

populations in the state of Karnataka along the

Cauvery and Kabini rivers and in the

Ranganathittu Bird Sanctuary. Ranganathittu has

approximately 17-20 adults and 10-15 subadults;

juveniles and hatchlings are also seen in this

Sanctuary (Venugopal 2000). Along remote

stretches of the Cauvery river, adult mugger are

sporadically seen and the estimated total

population along this river is 50 adults. The Kabini

river (which originates in Kerala) flows through

the Nagarhole National Park and has a small stable

population within the protected area. During a

preliminary survey, 22 adults were counted by

direct sighting. The habitat is pristine, several

mugger tunnels and good nesting banks were

also observed within the Park (Andrews 1995).

Other areas which reportedly have small

mugger populations include the Tungabhadra

Reservoir (Bhadra Sanctuary) and Shiva

Samudram.

The state of Andhra Pradesh has been

involved in crocodile conservation since 1976

and has released over 320 mugger in seven

different localities (Vijaya Kumar and Choudhury

1990, Vijaya Kumar 1993, Srinivas et al. 1999).

Isolated small populations occur in Manjira,

Ethypothala and Siwaram Wildlife Sanctuaries,

but their present status is unknown.

The estimated mugger population in

Rajasthan is around 470, distributed in 1 1

protected areas (Sharma et al. 1999).

Ranthambore National Park has the highest with

200 crocs and the National Chambal Wildlife

Sanctuary has around 100. In the other protected

areas numbers vary from 10-50 individuals. There

are increasing reports of human - croc conflict

from the Chambal area which calls for immediate

education programmes and a Flying Squad to

capture and translocate problem animals.

Mugger in Kerala are found in the Neyyar

Reservoir, Parambikulam Reservoir and the Nugu

and Kabini rivers. Conservation work was taken

up during 1977 when the wild population in the

State was less than 60, and releases have been

done in Neyyar and in Parambikulam. The current

estimated population is 259, based on the surveys

of 1997 (Pillai 1999). The highest densities are in

Neyyar and Parambikulam. The population in

Parambikulam is stable and the habitat is still

pristine (Andrews, pers. observ.). Currently Kerala

has 192 mugger in captivity and has to deal with

constant human - croc conflicts in Neyyar

Reservoir. The Forest Department has started an

education programme and the capture of large

problem animals.

In Maharashtra, C. palustris were once

abundant in all the major river systems and lakes,

but were depleted by the 1 960s (Gogate 1 999).

The first author remembers that, at Powai Lake in

the late 1950s, a reward of Rs. 50 was offered for

killing a crocodile and Rs. 10 for destroying a

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CROCODILE CONSERVATION

crocodile egg, probably by the Fisheries

Department. A conservation programme was

started in 1 977 and reintroduction was carried out

up to 1 992 in and around protected areas, however

there have been no surveys or monitoring, so no

information on the current wild population is

available.

In Goa, small breeding populations of

mugger still exist in several parts, including

Selaulim Dam, Opa river and small lakes in the

Ciba-Geigy and IOAR compounds. The largest

concentrations are found in the mangrove-lined

brackish creeks of the Zuari and Cumbarjua rivers.

While there has never been any formal release

programme, 38 rescued mugger have been

released here in the past few years.

Unfortunately, crocs are killed by local fishermen

and illegal clearance of mangroves has

devastated the habitat of the mugger and a myriad

other taxa (Alvares and D’ Sousa, in press).

Perhaps the most remarkable feature of the

mugger situation in India is that while there are

less than 2,500 left in the wild (Table 2), there are

over 3,000 in captivity at the various State

Government rearing centres and at the Madras

Crocodile Bank. Captive breeding and wild egg

collection for rearing has been stopped since 1 985,

because suitable release sites are scarce and so

are funds. The entire croc conservation

programme came to a halt mainly because

sustainable use of the mugger as a resource was

not taken up, although it was one of the goals of

the GOI/FAO/UNDP project.

Mugger outside India

Sri Lanka: Sri Lanka has a large mugger

population probably numbering several thousand

throughout the island nation (Whitaker and

Whitaker 1979, Santiapillai et al. 2000). While the

largest numbers occur in two national parks, Yala

and Wilpattu, mugger can be seen in large tanks,

in rice fields, waterways and small and large rivers

in most sparsely populated areas. The mugger is

strictly protected in Sri Lanka and a crocodile

management and conservation project is being

formulated (Whitaker, in prep.). A recent film on

the mugger in Sri Lanka reveals unique underwater

footage of breeding behaviour and parental care

Table 2: Total wild mugger in India

Source: ENVIS (Wildlife & Protected Areas) (2)1, 1999; Vijaya Kumar and Vyas 1997; Alvares and D’Sousa, in

press.

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CROCODILE CONSERVATION

by the male. Sri Lanka holds the responsibility for

the long-term survival of the mugger.

Iran: In Iran, a small number of muggers are

hanging on by the skin of their teeth in the

southeastern, Baluchestan region. Frazer (2000)

has reported areas of riverine and estuarine

wetlands, such as the Govater Bay and Hur-e-

Bahu (which is a 75,000 ha Ramsar site), as

important for Crocodylus palustris. Other areas

include the Sarbaz and Kajou rivers, where

muggers are protected by local folk beliefs (Kami

andSaghari 1993). During 1992 surveys, 1 6 gando

(= crocodile in Farsi) were counted in the Kajou

river and 30 in the 16 adjacent ponds. During the

same survey, 72 muggers were recorded in the

Sarbaz river and adjacent ponds along it (Kami

and Saghari 1994). Mobaraki (1998a) located nests

in the Bahukalat Protected Region. Here muggers

inhabit small streams and pools, and during the

dry months, migrate across the border into

Pakistan (Kami and Saghari 1993, Mobaraki

1998b). There is some Government interest in

mugger conservation, and select areas are now

protected. However, more recent information on

current status is lacking.

Pakistan: The mugger of Pakistan was

thought to be near extinct, with a few small isolated

populations scattered in various areas (Khan 1993,

1994). A captive-breeding programme was started

during 1993 and the Government of India supplied

gharial and mugger for this programme. However,

during 1999 surveys, an estimated population of

500- 1 ,000 mugger has been reported for the Deh-

Akro Wildlife Sanctuary in Sindh Province, which

consists of 25 lakes formed by the seepage from

irrigation channels. There are also reports of crocs

in Hingol National Park, the largest park in Pakistan

with an area of 619,043 ha (Chaudhury 2000).

Currently all the major wetlands in Pakistan are

threatened, thanks to the World Bank-aided

‘National Drainage Programme’ which involves

construction of dams and channels, diversion of

natural flows, and funnelling agricultural runoffs

into river systems (Rose 1998, Chaudhury 2000).

Nepal: In Nepal, mugger have been reduced

to small isolated populations mainly in protected

areas, such as the Royal Sukla Phanta Wildlife

Reserve, Bardia Reserve and the Royal Chitawan

National Park. A small number have been reported

from the Sapta Koshi area and the Lumbini district.

The Mahakali and Bahuni rivers adjacent to Sukla

Phanta represent excellent habitat and are

contiguous with similar areas in Uttar Pradesh.

Surveys by IUCN Nepal revealed depressed

numbers in this area. The estimated wild population

of mugger is around 200 (Andrews and McEachem

1 994). Loss of habitat and construction of dams

and barrages has affected C. palustris populations

and recruitment in the wild. Each year, juveniles

and hatchlings, flushed below the barrages (low

dams), cannot return during the post monsoon

season and usually perish. Conservation efforts in

Nepal started in 1 978 with wild gharial egg collection

for rearing and release programmes. Releases started

in 1981 and IUCN Nepal started a programme for

mugger in 1992.

Bhutan: In Bhutan, mugger became extinct

by the late 1 960s and during 1981a captive breeding

programme was started at a centre at Phuentsholing.

Mugger from Uttar Pradesh, India, were obtained

for this purpose. In May 2000, there was a small

group of 17 mugger breeding successfully, and 4

adult female gharial (Whitaker, pers. observ.).

Mugger have been released in the Manas river.

There is, however, no information on the fate of

these animals (Dorji, 1997 and unpubl. data).

Extensive surveys are suggested for the Bado,

Manas, Sunkosh Torsa, Raidak and the Puna

Tsongchu river systems, from where there are

sporadic reports of both mugger and gharial.

Saltwater Crocodile

The saltwater crocodile Crocodylus

porosus{ Plate 2, Fig. 3) is happiest in undisturbed

mangroves, now one of India’s most threatened

habitats. It is no surprise that the ‘saltie’ is now a

rare reptile in South Asia, and in India restricted

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CROCODILE CONSERVATION

to the Bhitarkanika Sanctuary in Orissa, the

Sunderbans in West Bengal and the Andaman

and Nicobar Islands, although historically this

species occurred all along the west and southeast

coast of India (Fig. 3). Table 3 gives the numbers

of wild salties in India.

Orissa: In 1976-77, prior to releases, the

density of salties in the Bhitarkanika Sanctuary

was 0.87 individuals per km of river (excluding

hatchlings), there were 29 adults, six subadults

and 61 juveniles (Kar and Patnaik 1999).

Following this, about 2,000 young crocodiles

were released in Bhitarkanika over the next 24

years. Along with the successful restocking of

gharial on the Chambal river, the Bhitarkanika

saltie project was a dramatic success. In 1998, a

total of 672 crocs were counted (including 150

hatchlings, 146 yearlings, 160 juveniles, 144

subadults and 72 adults) for a density of 5.0 crocs

per km (Kar and Patnaik 1 999). In 200 1 , a census

conducted by the Rajnagar Mangrove and Forest

Division came up with a figure of 1 ,285 salties in

Bhitarkanika Sanctuary (Kalpavriksh 2002). The

carrying capacity of the Sanctuary is limited by

the absence of any buffer zone surrounding it.

Any croc straying out of the protected area has

little hope of survival. The rearing centre at

Dangmal in the heart of the Sanctuary now has

about 300 captive salties. Egg collection from wild

nests has been stopped there.

West Bengal: The Sunderbans in West

Bengal is a vast tidal swamp Tiger Reserve and

part of one of the world’s largest mangrove

ecosystems. The sad part is that there are very

few salties in what is ideal habitat for the species.

Fig. 3: Historical range and current distribution of the

saltwater crocodile around the Indian subcontinent

The Bhagabatpur croc project of the West Bengal

Forest Department has released 332 young salties

in the 1 .2-1.5 m size class in the Sunderbans since

1 979, but there has been no monitoring or recent

census (Chowdhary and Choudhury 1999). A lot

of the best nesting habitat (swampy areas where

freshwater meets the tidal zone) has been

converted to paddy cultivation long ago. The

entry of thousands of permit holders into the

Reserve to fish, and to collect honey and leaves

for roofing creates continuous disturbance.

Andaman and Nicobar Islands: In the

Andaman and Nicobar Islands, the mangroves

are still largely intact on the seaward side, but the

inland portions of North Andaman Island were

destroyed for firewood, and saltie nesting habitats

used for growing rice (Whitaker and Whitaker

1978, Andrews and Whitaker 1994). Early

Table 3: Total wild saltwater crocodiles

Source: ENVIS (Wildlife & Protected Areas), Vol. 2, No. 1, 1999.

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estimates of the population in North Andaman

were 100-200 animals (Whitaker and Whitaker

1978). Choudhury and Bustard (1979) and

Choudhury (1980) estimated that there were 50

breeding females for North Andaman Island.

Andrews and Whitaker (1994) recorded the adult

population of North Andaman as 95 animals, but

a subsequent survey of a single island, Landfall,

by Andrews ( 1 999a) turned up 3 8 adults. Ten nests

were located but North Andaman settlers had

raided all. There are an estimated 19 adults in

Middle Andaman, 19 adults in South Andaman

and 27 adults in Little Andaman Island. The total

estimate for adults in the Andaman Islands is thus

198 (Andrews 1999a).

Surveys conducted during 2000-200 1 in the

Nicobar Islands indicate that most of the adults

and subadults have been taken by Thai poachers,

except in the Galathea creek on the southeast of

Great Nicobar island where a stable population still

exists. The west coasts of Great Nicobar and Little

Nicobar Islands have pristine habitats but

population densities comprise very few adults and

mostly hatchlings, yearlings, juveniles and a few

subadults (Andrews, in prep.). A small population,

of all size classes, occurs in the Middle Nicobar

group, however the mangrove habitat in this region

is very scanty and cannot sustain a large population.

Saltwater crocodiles outside India

Myanmar still has some very small

populations and individual Crocodylus porosus

scattered around the region. Surveys conducted

by Thorbjamarson et al. (1999) and Platt (2000)

indicate that there are still small, fragmented

populations and there is reproduction.

Thorbjamarson et al. (1999) reported 10 adults

and 100 juveniles plus hatchlings and nests within

the Meinmahia Kyun Wildlife Sanctuary, a 136

sq. km island. Outside the protected area, salties

are near extinct, with only a few isolated

individuals remaining.

Sri Lanka also has isolated populations of

C. porosus , mainly on the southwest coast which

still has some suitable habitat left. Detailed

surveys are required to establish the status and

survival potential of this, now rare, species.

The Madras Crocodile Bank Trust

The Madras Crocodile Bank was set up in

1975, for the breeding and rearing of South Asia’s

three crocodilians, for restocking wild habitats. It

is also a research and public education centre.

The Bank has achieved these goals with the

breeding of thousands of mugger and also gharial

and saltwater crocodiles. Close to a thousand

crocs, besides eggs, have been supplied for

restocking and for breeding programmes

throughout India and in Bangladesh. The Bank

has generated a number of documentary films,

several books and over 600 scientific and popular

articles. Over the past quarter century, more than

8 million people have visited the Bank and learned

about crocodilians. The Crocodile Bank presently

houses over 2,500 crocs of 14 species, snakes,

lizards and turtles and is recognised as India’s

Centre for Herpetology. It is also the office of the

Vice-Chairman, West Asia, Crocodile Specialist

Group (CSG) of the IUCN Species Survival

Commission (SSC). Current ongoing studies

include crocodile surveys and habitat assessment,

temperature sex determination, growth, behavioral,

breeding biology and hormonal research.

Discussion

Up to the 1970s, the three crocodilians of

South Asia were in big trouble, due to killing for

skins and meat, eating of their eggs and the loss of

most of their habitat. In India the GOI/FAO/UNDP

Crocodile Conservation Project, the Madras

Crocodile Bank Trust, State Forest Departments,

and like-minded individuals reversed the decline.

About 9,800 gharial, mugger and saltwater

crocodiles were released into established as well

as newly protected areas in India, and for the first

time positive publicity was given to these long

feared and maligned reptiles. Data is difficult to

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CROCODILE CONSERVATION

extrapolate from the hodgepodge of non-

standardised survey reports from some states

and no data at all from others. However, it appears

that there is little room for complacency with less

than 1,500 wild gharial, 2,500 wild mugger and

under 1 ,200 wild saltwater crocs left in India. There

was a certain amount of spin-off from the Indian

efforts in Nepal, where the Frankfurt Zoo helped

start an egg collection, rear and release

programme with advice from the FAO project. A

similar start was made in Bhutan. Unfortunately

Pakistan, Bhutan, Myanmar, Sri Lanka and

Bangladesh have made no substantial

conservation effort towards protecting the crocs

they had left in the wild.

Action Plan

In India, the IUCN/SSC/CSG Western Asia

convened a regional meeting at the Madras

Crocodile Bank Trust in 1993 (CSG-India 1993),

and in Gwalior, Madhya Pradesh in 1997 (CSG-

India 1997) resulting in an Action Plan. Similarly,

the three crocodilians are given a place in the

South Asian Amphibian and Reptile Specialist

Group Action Plan.

Executive summary of the IUCN/SSC/Crocodiie

Specialist Group Western Asia region -

Crocodilian Action Plan

1 . Liaison with the Ministry of Environment and

Forests, mainly to reconstitute the Technical

Committee for crocodile conservation.

2. Research: The action plan identified several

priority areas, including a database for wild

and captive crocs, management and use.

Main research priorities, however, will be

continued surveys, monitoring, and

identifying potential restocking sites.

3. Publicity and awareness, especially in areas

of potential croc - human conflict.

4. Skill development and training.

5. Regional interaction and coordination.

6. Funding.

The outcome of these meetings has been

circulated to the Government of India, all State

Forest Departments, all regional members, IUCN/

SSG/CSG Chairmen and Steering Committee

Members.

The Future of India’s Crocodilians

After such an ambitious and successful

beginning to the rehabilitation of India’s three

endangered crocodilians, it is very disappointing

that few of the states involved have sustained

some level of interest and action.

Currently there is very little monitoring,

management or conservation efforts and studies

on crocodiles within India and in the region, and

there is an urgent need to revive this interest. It is

also vitally important to implement a feasibility

study for farming and ranching.

Crocodile conservation becomes necessary

because, aside from the obvious value of the skin,

meat and other by-products, the crocodile plays

a vital ecological role as a master predator in the

aquatic habitat where it lives. By preying on weak

and diseased fish and animals, it maintains genetic

quality; by its habit of selective feeding, it controls

predatory fish. Its presence thus actually helps

to increase yields of edible fish for man.

While many developing countries have

found to their dismay that crocodile populations

are remarkably easy to exterminate, crocodiles

have responded well to protective management

initiative, wherever adopted. Crocodile (and

alligator) ranching, farming and rehabilitation

programmes have been underway in several

countries for a number of years. Most of these

have been successful in maintaining wild

crocodile populations and protecting millions of

acres of wetland habitat, besides generating

income for local people.

The programmes vary from country to

country in dramatically different scenarios, from

outright licensed hunting of adult alligators (as in

Louisiana), to closed cycle captive breeding

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(South Africa) and collection of eggs in the wild

(Australia and Zimbabwe). Two things are

common to all of these wildlife management

operations: (a) local people (often tribal people)

are making a good economic return, and (b) wild

crocodile populations are doing well.

The Indian experience in crocodile

conservation and rehabilitation in the last 25

years has been very encouraging. However,

persisting with a simplistic policy of bans and

attempts to preserve wildlife for its own sake,

has again put crocodiles in the region under grave

threat. Here, conservation is anti-people and the

alarming decline in all our major wildlife species

is the tragic result. No single conservation

strategy can solve the problems faced by wildlife

in India and it is vital that we continue to test

new and innovative conservation methods —

even if it means upsetting some people. Most

opponents of sustainable use of wildlife are more

interested in protecting the principle of

“preservation” rather than trying to solve our

problems of dwindling wildlife. Fundamentalist

belief in the animal welfare movement and lethargy

on the part of some key government bureaucrats

are two reasons why conservation cannot

achieve in India what so many other countries

have accomplished.

In many countries, conservation through

sustainable use is a strategy that has proved

remarkably effective in saving wildlife and

involving the people in it. It is essential that India

too looks at all the various conservation options,

including sustainable use of wildlife if we are to

effectively conserve crocodiles and their

dwindling habitat. Crocodile farming has done

wonders for the crocodilians in many developing

countries and India is lagging way behind.

Most captive crocodiles in India are now

several generations removed from the wild (Plate

2, Fig. 4). They are no longer wildlife, they are

domestic reptiles in the same way chickens, sheep,

cows and pigs were once wild and have been

domesticated by humans. Wildlife utilisation is

already being practised on a massive scale by

India’s fishermen, tribals who collect minor forest

produce and by the Irula Snake Catchers’

Cooperative, whose members catch snakes to

produce life saving antivenin. Crocodile farming

can now point the way to a new and dynamic

approach to managing wildlife — before we have

lost everything.

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Kalpavriksh (2002): Crocodile population up in

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Rao, R.J. (1999): Status and Conservation of Crocodiles

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GROWTH OF KNOWLEDGE ON THE REPTILES OF INDIA, WITH AN

INTRODUCTION TO SYSTEMATICS, TAXONOMY AND NOMENCLATURE

Indraneil Das1

Key words: Reptiles, taxonomy, systematics, history of herpetology, India, Fauna of British India

The progress in our understanding of the contents of the reptile fauna of India is reviewed. The

early classification of the fauna was undertaken during Vedic times, and included groupings based

on form, medium occupied, sensory powers and whether wild or domesticated. The Linnean

system of binomial nomenclature, adopted since 1758, is the most widely used scheme of

classification, but has its limitations, chiefly in being incompatible with the principle of common

descent. Phylogenetic and other classifications, based on natural groups, have lead to a proliferation

of studies on systematics, from the traditional studies of morphology and anatomy, to the use of

more modem molecular techniques. Aspects of systematics, taxonomy and nomenclature are discussed

for non-systematists. The last stocktaking of the reptile fauna of India, in the three volume Fauna

of British India series by Malcolm Smith is compared with the fauna now known from the country.

A general conclusion reached is that there is much to be learnt of the country’s highly diverse and

endemic reptile fauna, but efforts need to be accelerated in the face of loss of habitats as a result of

deforestation and of systematic expertise itself, with cuts in research funding and realignment of

policies on basic research.

Introduction

“ Cross-cultural evidence indicates that

people everywhere spontaneously organize

living kinds into rigidly ranked taxonomic types

despite wide morphological variation among

those exemplars presumed to have the nature of

their type .”

(S.Atran, 1990: 70)

The earliest known attempt to classify the

reptiles of India was made during the Vedic period,

based on form, reproductive mode, medium

occupied (earth, air or water), the presumed

number of sensory powers possessed, and

whether wild or domesticated (Rao 1957, Ghildial-

Sharma and Sharma 1989). For instance, the work

‘Manu Smriti’, compiled between 200 bc and 200

ad, classified animals on the basis of their

reproductive modes, and snakes, crocodiles and

tortoises, therefore, were grouped along with birds

and fish, for being (primarily) oviparous. The

umasvati, a Jain work written between 135-219 ad,

’Institute of Biodiversity and Environmental

Conservation, Universiti Malaysia Sarawak,

94300, Kota Samarahan, Sarawak, Malaysia.

Email: [email protected]

classified animals according to their sensory

powers. Within this system, reptiles were grouped

with humans, for possessing the senses of sight,

hearing, taste, smell and touch. The classification

systems of ancient Indian physicians, Charak and

Susruta, made use of differences based on

habitats, and the eight categories identified

include aquatic species that live in water, those in

dry hills, amphibious species and animals living

in marshy or water-logged areas. The work

susruta nagarjuna classified snakes into 5 groups

(including both venomous and non-venomous).

So why did our ancestors consider it

important to classify animals? There are two

fundamentally opposing schools of thought.

Diamond ( 1 966) and Gould ( 1 979) considered the

primary purpose of indigenous names to be

utilitarian. On the other hand, the complex

classification and naming process (= folk taxonomy

and nomenclature) support Berlin’s (1992) theory

that humans are innately curious about the natural

world, and that names are supplied to species that

may not always have a direct utilitarian value.

In taxonomy, the modern science of

classification of animals and plants, it is said that

stability of combinations of nomen is an indication

of lack of progress! This generalisation is true for

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TAXONOMY OF REPTILES OF INDIA

the herpetofauna of India and surrounding

countries for the latter half of the last century,

when workers had to depend on the three-volume

(1931-43) monographic review of the herpetofauna

of southern and parts of southeast Asia by

Malcolm Arthur Smith. These volumes, which are

understandably dated, did not cover the

amphibians, despite giving their titles, though this

group was covered in its entirety in an even earlier

work, that of Boulenger ( 1 890). This essay briefly

summarises the history and direction of the

modem science(s) of systematics and taxonomy,

and thereafter reviews the status of taxonomy,

systematics and nomenclature of reptiles

inhabiting the Republic of India. It also analyses

changes in the current understanding of the group

compared with that which existed at the time of

Malcolm Smith’s monographs. The essay

concludes with comments on the future of

systematic research in the country. A glossary of

technical terms used in systematics, taxonomy,

nomenclature and in biogeography is appended.

Linnean Classification

“ For his descriptions of living nature

Linnaeus drew on his daily readings in Roman

poetry , particularly Ovid and Virgil.... Read

through the prism of later Romantic nature

poetry, Linnaeus ’ nature writings are beautiful

(L. Koemer, 1 990: 23)

Classification is defined as the grouping of

objects that share properties/attributes into classes,

although contemporary practise of systematics and

taxonomy places emphasis on common ancestry,

rather than commonality of character states (see

‘Phylogenetic classification’).

The system of binomial nomenclature owes

its existence to the Swedish physician, Carolus

Linnaeus or Carl von Linne (1707-78). Linnaeus

arranged ail species of plants and animals known

to him in a hierarchical scheme, arranging species

within a genus, and genera within a family, and

families within a class. Nearly two million scientific

names have been proposed for animal taxa since

1758 (Hammond 1 992). Many were named in the

last century, and most of these currently refer to

taxa valid at present. Some are no longer in use,

as they were subsequently shown to be a less

recent name of an existing taxa (= junior synonyms)

or same name for a different species (= homonym).

More rarely, a name is not available because it

has been suppressed by the International

Commission of Zoological Nomenclature, for the

purpose of achieving stability in names. However,

names placed in synonymy do not disappear from

use, they are listed as such and especially when

groups are revised, some of these may be revived

to accommodate distinct populations warranting

a distinct taxonomic status, especially from within

a constellation of closely related species.

To become available for any nomenclatural

act in zoology, nomina (namely, names of taxa with

a nomenclatural status) need to have several

properties. They must be published intentionally

as new, following the binomial system of

nomenclature, as outlined in the Fourth Edition

of the International Code of Zoological

Nomenclature (International Commission of

Zoological Nomenclature 1999). The earliest

nomina available are those mentioned in the 10th

edition of Linnaeus’s systema naturae (published

in 1 758), which was taken as the starting point for

biological names. New names being proposed are

to be followed by the words ‘new species’ or ‘sp.

nov.’; in the past, the word(s) or letters ‘Mihi’ or

‘M’ (meaning ‘from me’ or ‘to me’), ‘Nobis’, ‘Nob.’

or ‘N’ (‘from us’ or ‘to us’) were used to indicate

that a new name was being proposed. However,

in older works, these were also used to designate

new' name combinations and spellings.

A few words about the origin and evolution

of modern concepts of systematics and taxonomy

may be in order. Contemporary classification of

the living world is based on the Darwinian theory

of evolution (or more accurately, the premise of

common descent), and uses the Linnean system

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of binomial nomenclature (after January 1 , 1 758),

a ‘downward’ classification system. Hierarchies

of the Linnean system, therefore, do not

necessarily reflect natural (that is, geneological)

relationships as understood at present (see

Stevens 1997).

Categories in the Linnean system include:

kingdom, phylum, class, order, family, genus

(plural: genera), and species (singular and plural),

and are used to convey information on relative

positions of taxa in the taxonomic hierarchy.

Standardised termination (suffix) of name indicates

a specific category:

-ini is a Tribe name (e.g. Ranini, to which

the common frogs belong)

-inae is a Subfamily name (e.g.

Lygosominae, to which the slender-bodied scincid

lizards belong)

-idae is a Family name (e.g. Varanidae, the

family including monitor lizards and Bataguridae,

the family to which hardshelled turtles of Asia are

assigned)

-oidea is a Superfamily name (e.g.

Gekkonoidea, to which gekkonids and

eublepharids belong)

Scientific names of organisms are typically

coined in ancient Greek or perhaps more commonly,

in Latin, a language no longer in use (and therefore

not subject to change, as those that are in use are).

Latin and Greek were the languages of learned

people, such as scientists and other scholars.

Scientific names of organisms can also be derived

from indigenous names, such as the nomen for the

crocodilian Gavialis Oppel, 1811, derived from the

north Indian vernacular name, for this sole living

member ofthe Family, ‘Gharial’ (from ghara, meaning

earthen-pot-like), an allusion to the narial

excrescence in adult males. The ‘r’ was apparently

misspelt V in the original description.

Phylogenetic Classification

“ The conventional Linnaean hierarchy

will not be able to survive alone: it will have to

coexist with the ideas and terminology of

phylogenetic ( cladistic ) systematics...

...One should always keep in mind that an

important function of classifications is

information retrieval .”

(Preface by A. Minelli & O. Kraus to the

International Code of Zoological Nomenclature,

Fourth Edition, 1999: XVII)

The number of species concepts in theory

and practice continues to proliferate. The infertility

criteria has been traditionally used as a marker for

species limit in the Biological Species Concept, that

defines species as reproductive ly isolated groups

of interbreeding organisms, as opposed to the

Recognition Species Concept (most inclusive

population of individual biparental organisms which

share a common fertilisation system) or the

Phylogenetic Species Concept (clusters of

organisms possessing uniquely shared characters).

See also the section ‘Species Concepts’ for

definitions of these and other concepts.

Some shortcomings of the Linnean system:

• Sometimes, the Linnean system is thought

unnecessary for phylogenetic taxonomy;

• There is no information on common ancestry;

• The categories themselves are insufficient to

specify relationships.

These problems make the adoption of the

phylogenetic system of classification more

attractive.

Emil Hans Willi Hennig (191 3-76), in the mid-

1900s, presented a system of classification based

on phylogeny, underlining common descent as a

criterion for relatedness and excludes reliance on

mere similarity. Originally published in German in

1950 and translated into English in 1 966, it argues

that biological classification should be guided by

the evolutionary theory [“The best general

classification is one that exactly reflects the

geneological (or phylogenetic) relationships”]. A

phylogenetic tree constructed shows a branching

diagram of entities with hypothesised genealogical

relationships and sequence of historical events.

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The states for a particular character are identified

in each ingroup and outgroup, the states polarised

to distinguish ancestral (plesiomorphic) and

derived (apomorphic) states, and the shared,

derived (synapomorphic) states plotted onto the

cladogram of the ingroup. The tree thus

constructed shows the historical pathway

followed within the ingroup in the evolution of a

certain character. Modern phylogenetic

systematics (also referred to as ‘cladistics’)

therefore dates from the work of Hennig. Among

the anomalies in the old vs. new systems of

classification is the discovery that some groups

(including Reptilia) are paraphyletic (i.e., their

members do not share a common ancestor). The

squamates form a natural group, lizards and

snakes sharing sister relationships, and

crocodilians are in fact closer to birds than to

other living reptiles, and turtles are not related to

either of these three groups, and have a somewhat

unresolved relationship (see for instance, Rowe

1986). Thus, ‘Reptilia’ as a clade is an unnatural

entity and comparisons made on their biology

that exclude other groups, especially birds, are, at

best, incomplete. Indeed, these findings challenge

the naturalness of the group of organisms dealt

with in the science of herpetology, a group that

includes the amphibians (of uncertain

relationships, although the group itself appears

monophyletic; Zardoya and Meyer 2001).

Nonetheless, two textbooks of herpetology have

appeared in recent years (Pough et al. 1998, Zug

et al. 200 1 ), and there is a general proliferation of

societies, journals and symposia dedicated to the

science of herpetology.

Phylogenies provide important insights into

taxonomy, including the evolution of characters and

long-term patterns and dynamics of adaptation and

divergence. The three overlapping charges of

modem systematics include:

• The naming of species, according to the rules

of nomenclature

• Their classification, and

• The reconstruction of phylogenies.

Thus, the purpose of classification of

organisms has witnessed a shift away from the

mere groupings of species at the time of Linnaeus,

to the arrangement of taxa to show natural (or

evolutionary) relationships. Recent advances in

molecular techniques provide new characters for

refining classification, that are particularly useful

for recognition of cryptic species, recognition of

large, unresolved groups and for resolving

phylogenies at higher levels.

Despite its widespread acceptance for the

past two and a half centuries, the Linnaean

binomial nomenclature is now known to be

logically incompatible with the phylogenetic

nomenclature ( sensu de Queiroz and Gauthier

1992). The concept of genus, in particular, while

mandatory for binomial nomenclature, cannot be

incorporated into a phylogenetic system of

nomenclature, which requires abandonment of

mandatory ranks. While no resolution to

nomenclature under the system is at hand yet,

Cantino et al. (1999) presented as many as 13

methods for naming species in the context of

phylogenetic nomenclature. The systems take into

account the stability, uniqueness and ease of

pronunciation of species nomens; their capacity

to convey phylogenetic information, and

distinguish nomen governed by a Code

of Phylogenetic Nomenclature (now under

preparation) both from clade nomens and

from species nomens governed by the current

Codes of Botanical or Zoological Nomenclature.

Species Concepts

“In the consideration of competing species

concepts it is necessary that systematists examine

the precise implications of each, in terms of

theory as well as practice .”

(J.I. Davies, 1995: 556)

“In the field of nomenclature there is a set

of rules to guide us, but when it comes to species

concepts, it is ‘everyone for himself ' ...”

(J.W. Amtzen& A.M. Bauer, 1996: 321)

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Malcolm Smith’s tomes, upon which the

contemporary classification of Indian reptiles (see

Table 1 ) is largely based, follows the typological

species concept. Type specimens are designated

as an objective and non-ambiguous reference to

a natural population of a specific taxa (Dubois

and Ohler 1996). The type locality refers to the

geographic area from where the sample was

extracted. A single specimen that forms the basis

for the description of a new species is referred to

as ‘holotype’. A series of specimens form

syntypes ( 4 cotypes ’ in the older literature). From

the syntype series, a subsequent author (such as

a reviser) can designate a lectotype, which

becomes the nomen-bearing type. As a result, the

rest of the syntypes become paralectotypes, and

lose their primary type status. If types from the

original description are lost, destroyed or

otherwise not available for study, a neotype can

be designated to bring stability to the group.

Ideally, a neotype is obtained from the type locality

or a locality close to it. Typological thinking of

essentialism, which dates back to Aristotle, is non-

evolutionary in nature, and one might therefore

argue for the abandonment for such a non-

evolutionary scheme of classification. Particularly,

one of several lineage-based species concepts

may be more appropriate, as these are consistent

with the evolutionary history.

We assume here that species are real entities

and not concepts for the purpose of understanding

the natural world (see Hull 1976, for a discussion).

Species concepts abound. Opposing camps offer

the following (for more comprehensive reviews, see

Ereshefsky 1992, Lee 1995):

The Biological Species Concept is

associated with the idea that species are

populations of interbreeding organisms, that are

reproductively isolated from other such groups

(see Mayr 1942, 1963).

The Evolutionary Species Concept

emphasises the extension of species through time,

defining species as a lineage of ancestral-

descendant sequence of populations that evolve

separate from other such lineages, with its own

unitary evolutionary role and tendencies

(Simpson 1961, Wiley 1978). Frost and Hillis (1990)

discussed the application of the Evolutionary

Species Concept to herpetofaunal groups.

In the Recognition Species Concept,

species definitions are associated with the

‘recognition concept’ emphasising the unification

of species rather than their separation from one

another It emphasises common fertilisation and

specific mate-recognition systems shared by

conspecifics, rather than reproductive isolation

between heterospecific organisms (Patterson

1980, 1985).

The Isolation Species Concept, an

alternative to the Recognition Species Concept,

emphasise reproductive isolation between

organisms (Patterson 1985, Dobzhansky 1970).

The Ecological Species Concept, a

modification of the Evolutionary Species Concept,

emphasises the importance of ecologically based

natural selection in maintaining species, which

defines species as a lineage or a closely-related

set of lineages that occupies an adaptive zone

minimally different from that of any other lineage

in its range and which evolves separately from all

lineages outside its range (Van Valen 1 976).

In the Phenotypic/Phenetic Species

Concept, species definitions that emphasise the

evidence and operations used to recognise

species in taxonomic practice, especially those

within phenetic or numerical taxonomy, and

perceive species as discrete clusters of

morphologically similar organisms, isolated from

other such clusters (Sokal and Crovello 1970).

The Phylogenetic Species Concept,

associated with phylogenetics or cladistics, is

used for four classes of species definitions (Davies

1995). The Hennigian Species Concept recognises

species as lineages that occupy the internodes of

a phylogeny and go extinct when a speciation

event takes place. The Autapomorphic (or

Monophyletic) Species Concept recognises

species as minimal monophyletic groupings, as

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TAXONOMY OF REPTILES OF INDIA

evidenced by autapomorphies. The Phylogenetic

Species Concept recognises species as divergent

population systems (with fixed or diagnostic

characters combinations that need not be

autapomorphic). Finally, the Geneological Species

Concept is similar to the Autapomorphic Species

Concept but replaces the emphasis on monophyly

of organisms with monophyly of genes carried

by organisms.

The General Lineage Concept of de Queiroz

( 1 998) concludes, after reviewing the various species

concepts, that these refer to all aspects or properties

of a single common entity, and that differences

between these definitions are not as disparate as it

would appear (see also de Queiroz 1 999).

Adoption of either Simpson’s (1961)

Evolutionary Species Concept or de Queiroz’s

(1998) General Lineage Concept, for instance, has

led to dramatically altered faunal lists for

Continental North America (Collins 1990), Mexico

(Grismer 1999) and the Philippines Archipelago

(Brown and Diesmos 200 1 ). As might be expected,

difficulties in standardising lists (for biotic

inventories and for a variety of other needs), as a

result of proliferation of checklists based on

different species concepts, exist not just for

reptiles, but also for well-studied groups such as

birds (see Helbig et al. 2002, McKitrick and Zink

1988).

The Code of Zoological Nomenclature

“ Ordinary languages grow spontaneously

in innumerable directions; but biological

nomenclature has to be an exact tool that will

convey a precise meaning for persons in all

generations .”

(Preface by J.C. Bradley to the International

Code of Zoological Nomenclature, First Edition,

1961)

The word ‘nomenclature’ is derived from

the Latin nomen (= name) + calare (= to call), and

the function of scientific nomenclature is to

facilitate communication between biologists (who

may speak different languages). In this essay,

‘nomen’ refers to scientific names, especially

those coined according to the rules of binomial

nomenclature. Scientific nomen are typically

coined in Latin (a dead language) or ancient

Greek. Article 36 of the Code of Botanical

Nomenclature even makes it mandatory to

provide descriptions or diagnoses of new spec ies

in Latin, although zoologists have been exempted

from this practice. Nonetheless, the technical

vocabulary of practising systematists rests on

an extensive glossary of Latin, derived largely

from the early works of Pliny the Elder (23-79 ad),

with significant Greek influence (Steams 1992).

The International Code of Zoological

Nomenclature (ICZN) is the law-book for

zoological systematists, and applies to both living

and fossil animals (botanists and bacteriologists

have their own Code of Nomenclature). It is based

on, and defines the Linnean nomenclatural

system of classification (ranks of taxa). The Code

of Zoological Nomenclature appeared in 1 886,

and early authors are therefore sometimes

inconsistent in their nomenclatoral practices. The

current version is the Fourth Edition of 3 999,

which came into force from January 1 , 2000, and

replaces the Third Edition of 1 985, and takes into

account over 500 written comments by zoologists

and groups of zoologists, made in response to a

Discussion Draft that was distributed widely in

1995.

The Code adopted by the International

Union of Biological Sciences, is in dual English-

French texts, and published by the International

Trust for Zoological Nomenclature, London. The

publication comprises 90 Articles, a Glossary and

two Appendices (Code of Ethics and General

Recommendations). In its own words, the Code

“provides the maximum universality and

continuity in the scientific names of animals

compatible with the freedom of scientists to

classify all animals according to taxonomic

judgement.”

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Some properties of scientific nomen include:

• uniqueness (no two species should have the

same nomen),

• universality (one nomen applied to the same

species worldwide), and

• stability (in case of old, unused nomen

threatening relatively recent well-used

nomen, the Commission has the powers to

set aside the former).

The binomial system of nomenclature

prescribes how nomen are to be written: every

species has a generic nomen and specific nomen

(e.g. Homo sapiens , for humans), and both generic

and specific nomen are in italics. Subspecies

nomen are also rendered in italics, but not

categories above the genera, such as Family and

Order nomen. ICZN makes it compulsory for

nomen not to be hyphenated (e.g. novae-guineae)

and special font types representing diacritic marks

(e.g. accent, cedilla, tilde and umlat) are not

permitted. The name of authority and year follow

the scientific nomen (species/subspecies nomen)

of taxon, without parentheses in case an original

name suggested at the time of description is valid.

If the species nomen has been allocated to a genus

different from the one it was originally allocated

to, the name(s) of author(s) are placed in

parentheses. An invalid nomen (such as a junior

synonym) may be written without the use of

parentheses. Finally, nomen are available for use

only if they are validly published (in journals,

monographs, books and Faunas). Publication of

new nomen in electronic media, meeting notices

and conference abstracts do not constitute a

formal publication from a nomenclatural

standpoint. As for the nomen itself, every author

is entitled to provide a nomen for a valid taxon,

regardless of how long, unpronounceable, or

grammatically incorrect it might be. Exceptions

where the International Commission of Zoological

Nomenclature have intervened are rare, and

include the case of Dybowski’s (1926) rather

lengthy nomen such as Siemieniewicziechino-

gammarus , Axelboeckiakytodermogammarus,

Garjajewiakytodermogammarus and Cancelloi-

dokytodermogammarus, which were suppressed

(Opinion 105, International Commission of

Zoological Nomenclature 1929). Nomina with the

potential to offend or hurt a community can also be

suppressed.

Estimates of global species richness of

animals varies between 5-15 million, of which

slightly over a million species bear a scientific

nomen (Erwin 1997, Stork and Gaston 1990). The

geographical trend in the distribution of species

is higher species richness in tropical, rather than

temperate regions and high richness values for

such environments as rainforests and coral reefs.

Nonetheless, many species nomen are synonyms

(duplicate names for the same biological species)

or nomina dubia (names without reference to a

biological species), making estimates of total

species counts difficult (Alroy 2002). The most

current inventory of the world’s described species

can be found at http://www.sp200Q.org.

Old Techniques and New

“ The development of time-shared, multi-

access computing systems utilising standard

teletype machines and the availability of

extremely simple programming languages are

recent events that provide an opportunity for

some major changes in the work of sy sterna t is ts.”

(J.A. Peters, 1969)

“In the 1950s, it was cyto-

taxonomy ...Counting chromosomes and

analysing karyotypes was the ‘ in-thing

claiming to solve all taxonomic problems. Nearly

concurrent with fun with numbers, was

experimental taxonomy: determining species

relationships by testing infertilities via

hybridisation...

Then in the mid-1960s another fad

dominated the systematics scene — numerical

taxonomy ...The next holy grail was the fertile

family of flavonoid compounds that took the

spotlight during the era of chemotaxonomy. Note

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that each of these fads faded, if not to oblivion,

at least to second class citizenship...

And so now in the midst of the molecular

taxonomy craze, I must ask: How long will it

last? What is the life expectancy for this fun with

isozymes and nucleic acids for the taxonomist?

If the current preoccupation is anything like its

predecessors, it will be replaced. By what? I

predict that the next ‘in thing’ will be to

determine the genetic basis of character

differences .”

(A.R. Kruckeberg, 1997)

Classifications of organisms have been

traditionally based on morphology and anatomy,

that is, structures that can be perceived by the

naked human eye or through optical instruments

(such as microscopes). Recent advances in

physics and chemistry make it possible to utilise

techniques that were unthinkable a few decades

ago. These include bioacoustics (as it increasingly

becomes evident how species-specific animal

calls are, because of their connection with

reproductive activity), and especially bio-chemical

techniques, including protein electrophoresis.

Molecular systematics makes use of

proteins and genes, in drawing systematic and

biogeographic conclusions, including nuclear

DNA (deoxyribonucleic acid), mitochondrial DNA

and ribosomal nucleic acids (the basic assumption

is that hereditary information is carried by

molecules of DNA). Recent advances in molecular

techniques provide new characters for refining

classification. A variety of molecular techniques

are available, including allozyme/isozyme analyses

(comparisons in net electric charge); mitochondrial

DNA (mtDNA, which is useful in reconstructing

phylogenetic history), through PCR amplification

and sequencing (allowing rapid amplification of

sequences from a large number of samples).

Phylogenetic reconstruction in molecular

systematics is commonly done using

mitochondrial gene sequences and less commonly,

sequences of nuclear ribosomal RNA genes (see

reviews in Hillis et al. 1 996).

Molecular techniques are also crucial for

studies in phylogeography, described as “the field

of study concerned with the principles and

processes governing the geographical

distribution of geneological lineages, especially

those within and among closely-related species”

(Avise 2000). Phylogeography is a subdiscipline

of biogeography that seeks to interpret how

historical processes have affected geographical

distributions of gene-based organismal traits.

Genetics provide answers to questions regarding

measures of migration (gene flow) between

populations and separating human-induced

factors from natural cyclicity of population

contraction and expansion. Two types of

conservation units may be discernable, using tools

in molecular phylogenies: management units

(MUs), which represent sets of populations that

are currently demographically independent and

evolutionarily significant units (ESUs) that

together encompass the evolutionary diversity

of a taxon (Moritz 1996). Both are significant, the

former for short-term management, the latter for

long-term issues.

An archive of biological data and

software to promote public access to freely

available information can be found at

http://www.iubio.bio.indiana.edu.

Why Systematics?

“ Among many others, butchers and artists,

farmers and showmen all deployed distinctive

taxonomies in their work... ”

(H. Ritvo, 1997: xii)

“ For several of us, the priority is to

document biodiversity and to provide

identification tools: both are needed to promote

awareness and conservation. This means

goodbye to time-consuming, low > output methods.

Methods have to be selected, case by case,

according to the material being investigated,

the threats facing a habitat, the urgency of the

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TAXONOMY OF REPTILES OF INDIA

threats, the needed output, cost efficiency and

context. In the real (non-academic) world, this

is just called efficiency /”

(M. Kottelat, 1995)

Although systematic biology has been

known to be inextricably linked to conservation

biology, and indeed to human welfare itself, the

principles of systematics have not entirely been

integrated into the principles and practice of

conservation science (Dimmick et al. 1999). For

instance, classification and naming of organisms,

especially endangered species, is the first step

towards their recognition, and subsequently, their

conservation and management. Poor knowledge

of systematics and taxonomy have been

considered responsible for the inability to

recognise severe threats to the survival of many

species of European freshwater fishes, a group

under study for over 450 years (Kottelat 1997)

and even for extinction of taxa (Daugherty et al.

1 990). It is now generally acknowledged that many

of the most important organisms in agriculture

are little understood in the context of systematics,

with serious implications for pest management

and control, and sustainable agriculture (Miller

and Rossman 1 995). And it was systematists who

drew global attention to the biodiversity crisis.

Systematic Agenda 2000, a global initiative

to discover, describe and classify the world’s

species, is guided by a consortium of botanists

and zoologists. It outlines a programme to

discover, describe and inventory the Earth’s

biological diversity, seeks answers to four

fundamental questions:

• What are the Earth’s species?

• What are their properties?

® Where do they occur?

• How are they related?

Most of the world’s systematists work

through or with museums that have collections

of preserved materials belonging to their particular

group of specialisation. Natural history collections

themselves have a variety of uses: apart from

having aesthetic and cultural values, they have

proved invaluable in documenting the

distribution, variation and seasonality of activity

of species (Earl of Cranbrook 1996), and

supporting studies of time series to investigate

changes in ecology (Brooke 2000) and declines

of extant species (Shaffer et al. 1 998).

Notwithstanding the fact that a large number

of museums, regional and national, exhibit natural

history objects, only a few of these are genuinely

research museums, in that they can hardly be

considered caretakers of systematic resources and

support a staff dedicated to undertaking full time

research. Among the more well-known ones in

India are the Zoological Survey of India,

headquartered in Kolkata, with regional stations

throughout the country, and the Natural History

Museum of the Bombay Natural History Society

in Mumbai. Both institutions have collections that

date back to the middle or early part of the

19th Century, including many herpetological

types. The staff of both institutions have

traditionally played an important role in promoting

systematic zoology in the country and their

influence in the region has been significant,

particularly in view of the fact that few tropical

Asian countries have natural history museums

(see Ng 2001). A research museum is thus the

caretaker of not only biological specimens

presented in life-like dioramas for the visiting

public, but also a repository for skins, moults,

casts, skeletons, cleared-and-stained as well as

whole specimens in fluids, and their tissues, as

well as associated information, such as

georeferenced localities and ecological data, for

scientists. The databases can be made publicly

available either in hardcopy (such as the

museum’s official journals or occasional

publications), softcopy on CDs and diskettes or

on the Web, such as Mexico’s 1 ,069 bird species,

listed at http://chipotle.nhm.ukans.edu/nabin.

A large database, Atta (after the genus of leaf-

cutting ants from the New World) of the Instituto

Nacional de Biodiversidad (INBio), a scientific

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TAXONOMY OF REPTILES OF INDIA

institution, with ‘social orientation’ from Costa

Rica, maintains a relational database comprising

over 2 million records. Besides basic data,

Atta also stores taxonomic information,

photographs and illustrations, and is viewable at

http://www.inbio.ac.cr/en/default.html.

Many of the species awaiting formal

discovery by modern science in the future are

predicted to be taxonomically cryptic, or similar

to known species, hence simply not recognised

until a thorough revision of the group is

undertaken, sometimes utilising contemporary

laboratory (including gene sequencing) and field

(ecological and behavioural) techniques. Cryptic

species are frequently localised, some restricted

to patches of forests a few dozen hectares in extent

or to one or two adjacent hill streams, making

their discovery difficult, unless a concerted effort

is made to conduct an exhaustive inventory. Other

species may show populations with disjunctions,

structured into well defined phylogenetic

assemblages or metapopulations, some with

significant genetic variants, all requiring careful

consideration for identification and conservation

(see Sites and Crandall 1997). Supplying names

to these “hidden” species, thus, is the first step

towards their universal recognition and protection

(Longino 1993, Wheeler 1995). True, the

recognition of cryptic species is increasing the

conservation burden; it also emphasises the

importance of moving away from taxon-based

conservation to protection of the environment at

the level of landscapes and ecosystems (Lovich

and Gibbons 1997).

One of the goals of these studies is also to

develop stable taxonomies of the fauna. A product

for the public and often, for industry, as a result

of these researches is comprehensive (i.e.,

covering all nominal taxa) monographs, revisions,

field guides and other identification devices. Field

guides are important in promoting conservation

awareness and action and assisting capacity

building. They are also resources supporting

environmental assessments (such as monitoring

and evaluation) of development projects,

encouraging ecotourism, building biodiversity

databases, land-use planning through GIS

applications and the production of regional and

international Red Data Books of Threatened

Species (Whitten 1996).

Comparisons with Smith’s Fauna

‘'‘‘Systematic collecting has been carried

out in nearly all parts of the Indian Empire, and,

except in the more inaccessible mountain

districts, the herpetological fauna of the country

is now pretty well known.”

(M.A. Smith, 1931: 1)

Malcolm Smith’s three volume Fauna of

British India on ‘Amphibia and Reptilia’ (although

the first group was not covered), published in

1931, 1935 and 1943, was a vastly revised edition

of the 1890 volume bearing the same title by

George Boulenger. The latter series was to set

high standards for monographs that have been

emulated by many others for different countries

and regions of the world. Indeed, nearly half a

century after the three volumes were published,

they still remain the single most important source

of information for practising reptile systematists

and others who wish to obtain information on

the region’s reptile fauna. A large number of

species have been described since the days of

Malcolm Smith (who, incidentally, never

conducted field work in India, but had material

from Indian and other museums mailed to him in

London), several species were omitted and a

number have been synonymised under other

species, or revived from synonymy. Several new

genera have been created to house species

known at the time of the Fauna, and a few have

been relegated to the synonymy of existing

genera. In this section of the essay, a comparison

is made with the systematic knowledge of Indian

reptiles at the time of Smith’s Fauna and that

which exists at present.

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Table 1 presents a summary of species

nomen of Indian reptiles that are considered valid.

Column 1 lists the original nomen, retaining the

original orthography; Column 2 lists the nomen

as used in the Fauna of British India volumes;

Column 3 provides the current nomen of the taxon;

Column 4 provides the citation for the preceding

nomen combination, if different from that in

Column 3; and Column 5 provides additional

nomenclatural remarks, as thought appropriate.

Nomens of families, genera and species are

arranged alphabetically, and not in any inferred

evolutionary order. Finally, a glossary of technical

terms is provided for non-systematists (Appendix

1). The cut-off date for the checklist was January

31,2003.

The geographical coverage of Smith’s

Fauna was southern, and perhaps because Smith

was more familiar with the southeast Asian

region, being the Physician to the Royal Court of

Bangkok (see Smith 1957 for an account),

extending to mainland southeast Asia, up to

southern Thailand. Among species now known

to occur within the political boundaries of the

Republic of India, 450 species were listed in Smith.

The current checklist counts 506 species. Species

concept employed by Smith, different

approaches, not to mention paucity of material,

are factors which are partially responsible for

differences in the allocation of species nomen in

the two columns. For instance, Smith’s (1943)

volume on snakes recognised a single species of

Indian Naja , even treating the Andamans

endemic, sagittifera as a synonym of N.

kaouthia. More recent studies by WUster (1998a)

and Wiister and Thorpe (1992) have shown that

a specific status is more appropriate for these

populations. More importantly, the typological

classification employed by systematists at the

time of M.A. Smith has the potential to

misrepresent actual biodiversity values in terms

of species richness. Included in the problem is

the more frequent usage at the time of Smith of

the subspecies category [frowned upon by

proponents of the Evolutionary Species Concept

(and other related concepts). However, the

subspecies concept, synonymous with ‘island

races’ (of Inger 1954, 1961) and pattern classes

(e.g. Grismer et al. 1994, Shaffer and McKnight

1996), continues to have its proponents (see, for

instance, Smith et al. 1997).

The two lists differ in other ways too. As

many as 139 species are now assigned to genera

different from that in Smith. The usage of names

of species is made not only by systematists and

taxonomists, but also by practising physiologists,

geneticists, ecologists, conservationists, and

policy makers, to name just a few end-users of

taxonomic lists. Frequent name changes can be

exasperating to many, some of whom perhaps

unrealistically wish to have stability of names for

a fauna that is poorly understood. However, as

many as 308 species of Indian origin listed in

Smith have been allocated to genera different from

that originally described in, suggesting that we

are on the path towards reaching a more natural

system of classification, if the binomial system

of nomenclature continues to survive. A number

of species whose nomen were made available in

older works were inexplicably unlisted in Smith’s

Fauna, even as synonyms, including

Asymblepharus tragbulensis (Alcock, 1898),

Barkudia melanosticta (Schneider, 1801),

Coluber vittacaudatus Blyth, 1 854 and Typhlops

exiguus Jan in: Jan and Sordelli, 1 864. It is possible

that the present list too omits some valid species

described in obscure works. Indeed, the non-

availability of some publications has been a

serious impediment to studies, and several critical

works dealing with systematics and nomenclature

of the region’s reptiles are not easily accessible

to biologists. These include unpublished theses

(e.g. that of Moody 1980, which suggested the

overhauling of the generic-level systematics of

the Agamidae) and works in languages other than

English, and frequently of limited circulation

(that are far too numerous to mention in this

essay).

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Table 1: Synopsis of names of reptile species known from India. Nomenclature in M.A. Smith’s Fauna compared

with that followed at present. For original names, the original orthography has been retained (including upper case

first letters for species nomen, accents, etc.)

Original name Name in Smith’s Current name References Remarks

Fauna

CROCODYLIDAE

Crocodylus palustris

Lesson, 1831

Crocodylus porosus

Schneider, 1801

GAVIALIIDAE

Lacerta gangetica

Gmelin, 1879

BATAGURIDAE

Emys baska

Gray, 1830

Testudo amboinensis

Daudin, 1801

Cyclemys oldhamii

Gray, 1863

Emys Hamiltonii

Gray, 1831

Geoemyda silvatica

Henderson, 1912

Emys Thurjii

Gray, 1831

Emys dhongoka

Gray, 1832

Emys kachuga

Gray, 1831

Geomyda tricarinata

Blyth, “1855” 1856

Emys trijuga

Schweigger, 1812

Batagur ( Morenia ) petersi

Anderson, “1878” 1879

Batagur smithii

Gray, 1863

Crocodylus palustris

Crocodylus porosus

Gavialis gangeticus

Batagur baska

Cuora amboinensis

Cyclemys dentata

Geoclemys hamiltoni

Geoemyda silvatica

Hardella thurgi

Kachuga dhongoka

Kachuga kachuga

Geomyda tricarinata

Geoemyda trijuga

Morenia petersi

Kachuga smithi

Kachuga Sylhetensis Kachuga sylhetensis

Jerdon, 1870

Crocodylus palustris

Crocodylus porosus

Gavialis gangeticus

Batagur baska

Cuora amboinensis

Cyclemys oldhamii Fritz et al.

(1996, 1997)

Geoclemys hamiltonii

Geoemyda silvatica

Hardella thurjii

Kachuga dhongoka

Kachuga kachuga

Melanochelys tricarinata McDowell (1964) -

Melanochelys trijuga McDowell (1964) -

Morenia petersi

Pangshura smithii Das (2001);

Iverson (in prep.);