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Society

April 1987

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Journal of the Bombay

Natural History Society.

VOLUME 84(1) : APRIL 1987

Date of Publication: 15-10-1987

CONTENTS

Page

Changes in the soft part coloration of the Indian Reef Heron, Egretta gularis

(Bose) related to age and breeding status. By B. M. Parasharya and R.

M. Naik .. 1

Survey of the Freshwater Turtles of India Part II: The Genus Kachuga.

By Edward O. Moll. (With two colour plates and four text-figures ) 7

The Butterflies of the Nilgiri mountains of Southern India (Lepidoptera:

Rhopalocera) . By Torben B. Larsen .. 26

Notes on comparative body size, reproductive effort and areas of manage-

ment priority for three species of Kachuga (Reptilia, Chelonia) in the

National Chambal Sanctuary. By R. J. Rao and L. A. K. Singh. ( With a

plate and three text-figures ) . . 55

The genus Piper Linn, in Karnataka, India. By B. A. Rahiman and M. K. Nair.

( With eight text- figures ) . . 66

Polychaeta of the Pulicat Lake (Tamil Nadu). By S. K. Sunder Raj and P. J.

Sanjeeva Raj. (With four plates) . . 84

A Catalogue of the Birds in the Collection of Bombay Natural History

Society — 32. By Humayun Abdulali . . 105

Geographical distributional list of Ichthyofauna of the Garhwal Himalaya

with some new records. By H. R. Singh, S. P. Badola and A. K. Dobriyal.

(With a text- figure) . . 126

Sundarbans honey and the mangrove swamps. By Kalyan Chakrabarti . . 133

Annual reproductive cycle of the male field rat, Rattus rattus brunneusculus

(Hodgson) in hilly terrain of Mizoram. By N. S. Chauhan and R. N.

Saxena. (With a plate) . . 138

Studies on Anopheles (Cellia) maculatus Theobald, 1901 in Bastar District,

India (Diptera: C’ulicidae). By Zakir Husain Husainy. (With five text-figures) 145

On some insect galls on Tectona grandis Linn, from India. By P. Jayaraman.

(With seven text- figures) . . 157

Two UNUSUAL CASES OF HOMONYMY IN ORTHOPTERA WITH NEW NAMES FOR SPECIES

from India. By M. S. Shishodia and R. K. Varshney . . 167

New Descriptions:

A new species of Asynapta Loew (Diptera: Cecidomyiidae: Porricondylinae)

from Aurangabad, India. By R. M. Sharma. (With twelve text- figures) . . 170

On a new Cyprinid fish of the Genus Danio Hamilton (Pisces: Cyprinidae)

from Manipur, India. By R. P. Barman. (With a text- figure) . . 172

Ophiopinotus pinotus gen. et. nov. (Hymenoptera: Torymidae). By Tasawwer

Husain and Prem Prakash Kudesia. (With seven text-figures) . . 175

Taxonomic studies on the Marine Ostracoda from the east coast of India.

By C. Annapurna and D. V. Rama Sarma. ( With four plates) . . 177

Mecistocerus monubumensis sp. nov. (Coleoptera: Curculionidae: Cryptorhyn-

chinae) from India. By Lehna Singh Arya and H. R. Pajni. (With three text-

figures ) .. 181

Contribution to the knowledge of Desmids of India — some new taxa from

Karnataka State. By G. R. Hegde. (With five text-figures ) . . 183

Dasineura psorafeae (Diptera: Cecidomyiidae) — a new Gall-Midge, infesting

inflorescences of Psoralea corylifolia Linn. By R. M. Sharma. (With thirteen

text-figures ) . . 186

Obituary

S. P. Shahi (1917-1986) . . 190

Reviews

1. Annotated checklist of the birds of Hong Kong. (H. Abdulali) .. 191

2. Threatened animals of India. (R. Reuben) . . 191

3. The useful plants of India. (M. R. Almeida) . . 192

Miscellaneous Notes:

Mammals: 1. Group number and composition of Hanuman Langur (Presbytis entellus )

in Jaipur, India. By Reena Mathur and B. Ram Manohar (p. 193); 2. Occurrence of the

Bicoloured Leaf-nosed Bat (Hipposideros fulvus ) in Rajasthan. By S. Bhupathy (p. 199);

3. Sighting of a Rusty Spotted Cat (Felis rubiginosa ). By Digveerendrasinh (p. 200);

4. Note on the sighting of a Caracal (Felis caracal ) at the Sariska National Park. By

Divyabhanusinh (p. 201); 5. Occurrence of Large Indian Civet (Viverra zibetha) in Orissa.

(With a photograph). By L. N. Acharjyo and S. K. Patnaik (p. 201); 6. A note on the

food of the Small Indian Civet (Viverricula indica) at Point Calimere Wildlife Sanctuary,

Tamil Nadu. By M. Ayyadurai, V. Natarajan, P. Balasubramanian and S. Alagar Rajan

(p. 203); 7. Unusual coloration of Nilgai (Boselaphus tragocamelus) . By M. K. Ranjitsinh

(p. 203).

Birds: 8. Occurrence of the Barheaded Goose (Anser indicus ) in South India. By M.

Krishnan (p. 204); 9. The Osprey (Pandion haliaetus haliaetus ) preying on a gull. By

M. C. A. Jackson (p. 205); 10. Occurrence of Greyheaded Lapwing. Vanellus cinereus

(Blyth) in Bangalore. By S. Subramanya (p. 205); 11. Addition to the birds of Point

Calimere, S. India. By Nitin Jamdar (p. 206); 12. Further additions to the avifauna of

Point Calimere, Ruddy Shelduck (Tadorna ferruginea). By R. Sugathan, David S. Melville

and S. Alagar Rajan (p. 206); 13. The Whitewinged Black Tern, Chlidonias leucopterus

(Temminck) in Saurashtra, Gujarat. By Taej Mundkur (p. 208); 14. Occurrence of Pied

Crested Cuckoo (Clamator jacobinus ) in Suru valley, Ladakh. By Nitin Jamdar (p. 208);

15. Sighting of Blacknaped Oriole. By D. P. Banerjee (p. 209); 16. Nest of the Pied

Myna Sturnus contra Linnaeus. By Vasant R. Naik (p. 210); 17. Kaliveli Tank and

Yedayanthittu Estuary — A little known wetland habitat in Tamil Nadu. By Pieter (p 210);

18. The Redfronted Babbler Stachyris rufifrons and Redheaded Babbler S. ruficeps in

northern Thailand. By C. J. O. Harrison (p. 214); 19. The Indian Grey Tit (Parus major )

on an abandoned honey comb. By Debi Goenka and Heta Pandit (p. 218); 20. Host

plants used by Baya Weaver bird (Ploceus philippinus Linn.) for nesting in eastern

Rajasthan. By Satish Kumar Sharma (p. 218).

Reptiles: 21. Distribution of the keeled box turtle Pyxidea mouhotii (Gray). By Indraneil

Das (p. 221); Breeding the king cobra (Ophiophagus hannah ) in captivity. (With a photo-

graph). By Shekar Dattatri (p. 222); 23. A list of the snakes of the Bhavnagar District,

Gujarat State, By Raju Vyas (p. 227).

Amphibia: 24. A note on the breeding habits of Jerdon’s Ramanella, Ramanella montana.

By A. G. Sekar (p. 231);

Fishes: 25. On the synonymy of Danio feegraclei Hora, 1937 from Burma (Pisces:

Cyprinidae). ( With a text-figure). By R. P. Barman (p. 232).

Insects: 26. Observations on Indian Trabutinini Silvestri and Phenacoccini Sulc (Pseudo-

coccinae: Pseudococcidae : Homoptera). By R. K. Avasthi and S. Adam Shafee (p. 235);

27. Studies on the biology of Parnara naso Fabr. (Lepidoptera : Hesperiidae) . By V. N.

Rao and K. S. Behera (p. 238); 28. Distribution of Drosophila species inhabiting the

tropical rain forests of Sampaje Ghats (Coorg District), Karnataka, South India. By N.

Muniyappa and G. Sreerama Reddy (p. 240); 29. Polymorphism in Callosobruchus chin-

ensis (Linn.) (Coleoptera: Bruchidae). By H. R. Pajni, Sadhana Sahnan and Ruchira

Sharma (p. 245); 30. Occurrence and life history of Cassida circumdata Herbst (Coleoptera:

Chrysomelidae) in Keoladeo National Park, Bharatpur, India. ( With four text-figures ).

By M. John George and K. Venkataraman (p. 248).

Other Invertebrates: 31. A note on the food of the spider Argiope arcuata Simon

(Family: Araneidae). By V. Natarajan (p. 253).

Botany: 32. New distributional records of Dicots for Punjab State (India). By M. Sharma

(p. 254); 33. Caldesia parnassiifolia (Linn.) Perl. (Alismataceae) — A New Record for

Western Himalaya. By P. C. Pande and Y. P. S, Pangtey (p. 257); 34. Notes on Cypera-

ceae of Maharashtra. By S. M. Almeida and M. R. Almeida (p. 257); 35. Oianthus

disciflorus Hook. f. — A little known taxon from Peninsular India. (With a text-figure).

By R. R. Venkata Raju and T. Pullaiah (p. 258); 36. On the identity and synonymy of

Hedyotis congesta R. Br. (Rubiaceae). By D. B. Deb and Ratna Datta (p. 261); 37. A

note on the Brachystelma glabrum Hook. f. (Aselepiadaceae) from South India. By E.

Vajravelu (p. 262); 38. Notes on the distribution of rare and little known Carex rostrata

Stocks from north-west Himalaya. (With two text-figures) . By R. D. Gaur, K. S. Negi and

J. K. Tiwari (p. 263); 39. Two new records of grasses from Andhra Pradesh. (With two

text-figures). By N. Yesoda, P. Venkateswara Prasamia and R. R. Venkata Raju and T.

Pullaiah (p. 265); 40. Rectification of the position of Anthraxon microphyllus (Trin.)

Hochst. var. hindustanicus (Jain & Deshpande) Almeida and Almeida. By P. C. van

Welzen (p. 268).

©

JOURNAL

OF THE

BOMBAY NATURAL HISTORY

SOCIETY

1987 APRIL Vol. 84 No. 1

CHANGES IN THE SOFT PART COLORATION OF THE

INDIAN REEF HERON, EGRETTA GULARIS (BOSC)

RELATED TO AGE AND BREEDING STATUS1

B. M. Parasharya2 3 and R. M. Naik:'

Age dependent changes in the soft part coloration of the Indian Reef Heron,

Egretta gutaris (Bose) from the day of hatching to adulthood are described. The

nuptial colour changes in the soft parts are correlated with the nesting status.

Introduction

Coloration of the soft parts of the Indian

Reef Heron, Egretta gularis (Bose) changes

with the age and breeding status, so that the

coloration when viewed with other evidences

gives a clue to the bird’s age and breeding

status; a knowledge of the soft part colour

changes has been useful in some of our field

studies on the heron. Moreover, in certain

areas where the reef heron is sympatric with

the Little Egret, Egretta garzetta, they tend

to interbreed, and since a striking difference

1 Accepted August 1985.

2 Present address : B. A. College of Agriculture,

Gujarat Agricultural University, Anand 388 110,

India.

3 Department of Biosciences, Saurashtra Univer-

sity, Rajkot 360 005, India.

between the two species is coloration of their

soft parts, we feel that a detailed description

of the soft part colours at different stages of

life in both the species should prove useful

in further studies on the inter-relationship

between the two herons.

Plumage colour variations and sequence of

plumage changes in the reef heron are des-

cribed earlier (Naik and Parasharya 1983).

However, a brief review is necessary here. In

the white as well as grey phases the natal down

is white in most of the feather tracts, but it is

subjected to some colour variation ranging

from light grey to grey in the dorsal tract.

The pin juvenile feathers start appearing,

pushing out the natal down at the age of 6-8

days. The juvenile white plumage ranges

from an almost pure white to one heavily

dappled with grey. The juvenile grey is grey

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

dorsally and light grey or white ventrally. The

full adult plumage is attained at the age of

20-21 months. The adult white plumage is pure

white and the adult grey is grey all over except

the chin and throat which are white. Both the

juvenile and adult grey plumages may, or

may not, have wing patch of variable size on

one or both the wings.

Before the breeding season a drooping

nuchal crest of two narrow plumes develops in

both the sexes and phases. In addition to the

crest, filamentous ornamental feathers deve-

lop on the scapular region and long, narrow,

lanceolate plumes on the lower parts of neck

and upper breast. Nuptial feathers are shed

just after the nesting is over, but in some

cases it happens in April when the birds are

still feeding their young, so that in the begin-

ning of the breeding season (February) 90%

of the birds may be with nuchal crest but

during the second peak of breeding (July-

August) only 40% may have it.

Materials and Methods

The present report is based on our field

notes made during the studies on the distri-

bution (Naik and Parasharya in press), breed-

ing biology (Parasharya and Naik in press',

Naik and Parasharya in prep.) and ecology of

the reef heron (unpublished) and also on our

observations of some captive birds maintained

in our University aviary for studying the

sequence of plumage changes and polymor-

phism (Naik and Parasharya 1983).

Observations

Colours of the soft parts described below

are applicable to both the grey and white

phases of the Reef Heron, unless mentioned

otherwise. There were always a small number

of E. garzetta and/or garzetta-gularis hybrids

inter-breeding with E. gularis in the breeding

colonies, so that some of the variations in the

coloration of soft parts may possibly be attri-

buted to the hybridization. The age dependent

changes in the plumage and coloration of soft

parts are summarized in Table 1.

Downy nestlings

Skin. The freshly hatched chicks have pink-

yellow (flesh coloured) skin all over. The skin

appears greyish green at the age of 3 days

when the feather follicles deep within the skin

start generating the juvenile plumage.

Eye. Iris colour of the freshly hatched chick

is brown, and it remains so until the natal

down is pushed out by the juvenile feathers.

Beak. The beak of freshly hatched chick

is pinkish yellow with some brown at the

tip of both the mandibles and base of the

upper mandible. By the time the juvenile

feathers start appearing as pin, the brown on

the upper mandible has increased in density

to be prominent. Though the brown on the

lower mandible also tends to spread a little,

the lower mandible continues to remain almost

yellow throughout the downy plumage stage.

Legs. At the age of about 3 days, a slight

green tinge appears on the tibia and upper

tarsus and the feet start turning light yellow.

Gradually the tibia and tarsus become light

greyish green and the feet become light

yellow, tinged with green.

Lore. Lore (facial skin) which is pink-

yellow at the time of hatching gradually turns

dark brown to grey.

Chicks in juvenile plumage and subadults

Skin. The skin which appears greyish green

during the initial stage of the juvenile plum-

age growth, starts turning yellow after the

age of 10 days, and it is conspicuously yellow

2

SUMMARY OF AGE-DEPENDENT CHANGES IN THE TYPE OF PLUMAGE AND COLORATION OF SOFT PARTS IN

SOFT PART COLORATION OF REEF HERON

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JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

by about the 16th day. Later, the skin starts

turning grey but the yellow tinge may conti-

nue to persist for a long period. However, all

the birds acquire uniformly grey skin before

reaching the age of 6 months.

Eye. Slight white tinge appears in the

brown iris at the age of 11 days, and the

brown progressively becomes lighter. The iris

turns cream coloured by the age of 4 weeks

and yellow by the age of 6 months.

Beak. The upper mandible is largely dark

brown or grey and the lower one yellow

when the juvenile feathers start appearing. By

the age of 20 days, threefourths of the upper

mandible becomes yellow, the brown being

restricted to the base and tip. An overall

appearance of the beak is yellow at the age

of 4 weeks but the brown at the base and tip

continues to persist and does not disappear

completely even at the age of 6 months.

Legs and feet. When the pins of juvenile

feathers appear, the tibia and tarsus are grey-

ish green and the feet light yellow. The tibia

and tarsus gradually turn yellowish green, but

at the age of 15 days, a tinge of grey appears

on the front face of the tarsus. At the end

of 4 weeks, the tibia and tarsus are greyish

green, and the frontal surface of the tarsus

is marked with black bands. At the end of

6 months a slight dull green colour still

persists on both the tibia and tarsus of some

birds, though they have turned completely

black in the others. The feet remain light

yellowish green at least up to the age of 6

months, but thereafter they gradually turn

light yellow.

Lore. The lore remains dark till 4 weeks of

age. Thereafter, light greenish yellow colour

appears among brown and gradually the yellow

tinge increases to make the facial skin yellow

with only a little tinge of green by the end of

6 months.

Adult

Eye. The iris of the adult bird is yellow.

Beak. The beak colour of the adult bird is

variable. Variations in the beak colour are

categorized into two basic types, yellow and

black, which occur in the grey as well as

white morphs in the non-breeding as well as

breeding seasons. It may be mentioned here

that the birds feeding in certain types of

habitat often get their beak smeared with

mud which on drying obscures the true beak

colour (Naik et al. unpublished).

Yellow : This may range from pure yellow

to pinkish, (a) Pure yellow: Both the man-

dibles are pure yellow. This is the most fre-

quent beak type, (b) Pinkish yellow: Though

the basic colour of both the mandibles is

yellow, there is a pinkish tinge all over.

Though infrequent, the pinkish yellow beak

was recorded during the non-breeding season

also.

Black: Both the mandibles are completely

black without any yellow tinge.

Lore. The lore of the adult birds is yellow.

Occasionally we came across birds that had

small patches of blue on the yellow lore and

also those with bluish green or blue lore.

Legs. In non-breeding adults, the tibia and

tarsus are black and the feet are bright

yellow. The bright yellow of feet often ex-

tends up to the distal end of tarsus.

Nuptial colour changes

The beak, lore and iris are usually yellow,

but they acquire the nuptial colours during

the courtship (pre-pairing and pair formation)

stage of the nesting cycle. In both the grey

and white phases, the beak usually remains

yellow, but in some it may acquire a red tinge.

If the bird happens to have a black beak, no

recognisable change in the beak colour occurs.

4

SOFT PART COLORATION OF REEF HERON

The iris which is normally pale yellow turns

to bright yellow and may even acquire a red

rim in some birds. The lore turns to bright

pink red or crimson. The feet and lower

tarsus which are normally yellow turn to

pink-red. The upper tarsus and thigh region

which is dull grey normally do not change

colour, but in some birds even these parts

also acquire a red tinge. Pink-red in the feet

and lore of the individuals perching alone

(unpaired) is brighter than that of paired

birds or nest-building birds. The pink colour

tends to fade during the nest-building phase,

and it almost completely fades out when the

first egg is laid. The toes turn to pale orange-

yellow and ultimately resume their original

colour. The lore first turns to pale pink, then

to bluish white and thereafter to pale yellow

and ultimately to its original yellow colour.

If a pair loses its clutch during the incuba-

tion period and if it is to re-build the nest,

the nuptial colours return to the soft parts

once again. The birds breeding in juvenile

plumage also acquire nuptial colour when

they start nesting.

We recorded the beak colour of 181 white-

phase reef herons in different stages of their

nesting cycle in breeding colony and the re-

cord is summarized in Table 2. These data

illustrate some of the points we have made

hereinbefore for both the phases, namely the

yellow is the most frequent beak colour for

the birds in every stage of the nesting cycle,

the 3^ellow beak may turn pinkish in some

(but not in all) birds during the nesting

season and a small proportion of birds have

their beak black in colour which remains

apparently unchanged through the nesting

season.

Discussion

The age-dependent changes in the colora-

tion of soft parts of Indian Reef Heron, des-

cribed in this paper and summarized in Table

1 for the convenience of the readers, should

be useful in ageing the chicks and distinguish-

ing the older from younger juveniles in the

field.

Table 2

Frequency of different beak colours of the

WHITE-PHASE REEF HERONS IN RELATION TO THEIR

NESTING STATUS, AS RECORDED IN THE NEW PORT

Colony, Bhavnagar

In the present state of our knowledge it is

not possible to say as to how much variation

in the soft part coloration described here is

attributed to hybridization between the Little

Egret and reef heron that has been occurring

since long in our area (Naik and Parasharya

1985) However, it is certain that the appear-

ance of blue lore and black beak (which are

characteristic features of the Little Egret) in

our population of the reef heron is a result

of gene flow from the Little Egret populations.

Nuptial colour changes in soft parts during

early stages of the nesting cycle are now

known to be widespread in the family

Ardeidae (Blaker 1969). Ali and Ripley

(1968) have mentioned colour changes asso-

5

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

dated with breeding in many Indian Ardeidae,

but not for the Indian Reef Heron. In certain

populations of the reef heron in Africa, the

beak turns from yellow to black as a part of

nuptial colour changes (Hancock and Kush-

lan 1984). In our population of the reef

heron, however, only noticeable change in the

beak colour is for the yellow to turn pink-

yellow in some birds. A few birds that we

observed having pink-yellow beak in the non-

breeding season were probably exceptional;

an increased blood flow in the subcutaneous

network of blood capillaries, either because of

some emotional stress or premature hormonal

changes, may have turned the beak pinkish.

Recently, Hancock and Kushlan (1984)

have merged the reef heron with the Little

Egret. These authors consider the Little Egret,

Egretta garzetta (Linnaeus) as a polytypic

species consisting of six subspecies, namely

garzetta — an all white race with a range in

the temperate, subtropical and tropical zones

of the Old World, nigriceps — also an all-

white race occurring on the islands of south-

eastern Asia and the southwest Pacific, im-

maculata — also an all-white race of northern

R EFE

Ali, S. & Ripley, S. D. (1968): Handbook of

the birds of India and Pakistan. Vol. I. Oxford

University Press, Bombay.

Blaker, D. (1969) : Behaviour of the Cattle

Egret Ardeola ibis. Ostrich 40(3) : 75-129.

Hancock, J. A. & Kushlan, J. A. (1984) : The

Herons Handbook. Croom Helm, London.

Naik, R. M. & Parasharya, B. M. (1983):

Sequence of plumage changes and polymorphism,

in the Indian Reef Heron, Egretta gularis. Sand-

grouse 5: 75-81.

— — — (in press ) :

and eastern Australia, gularis — a polymor-

phic race on the west African coast from

Mauritania to Gabon, schistacea — another

polymorphic race distributed along the coast-

line extending from the west coast of Indian

subcontinent to Persian Gulf and the Red

Sea, and south along the African coast up

to Tanzania, and dimorpha — also a poly-

morphic race found on Madagascar and

other islands in and around Mozambique

Channel. While available information on the

soft part coloration of the above mentioned

assemblage of herons are illustrated by Han-

cock and Kushlan (1984), detailed informa-

tion of the changes in soft part coloration

dependent on age and nesting status in these

birds are urgently needed to understand their

evolution.

Acknowledgement

One of us (B.M.P.) is grateful to the

Council of Scientific and Industrial Research,

New Delhi, for an award of Senior Research

Fellowship.

E N C E S

Impact of the food availability, natural-habitat des-

truction and regional cultural variations of human

settlements on the nesting distribution of a coastal

bird. Egretta gularis, in western India. J. Bombay

nat. Hist. Soc.

(in prep.) :

Inter-breeding and gene flow within Egretta guTaris

and E. garzetta complex, along the Gulf of Kham-

bhat in western India.

Parasharya, B. M. & Naik, R. M. (in press) :

Breeding biology of the Indian Reef Heron, Egretta

gularis (Bose). J. Bombay nat. Hist. Soc.

6

SURVEY OF THE FRESHWATER TURTLES OF INDIA

PART II: THE GENUS KACHUGA 1

Edward O. Moll2

{With two colour plates & four text-figures)

[Continued from Vol 83(3): 552]

Subgenus Pangshura

Contains four species — smith'd, sylhelensis,

tecta and tentoria. The subgenus is diagnosed

by a suite of apomorphic or derived charac-

teristics summarized in Table 1 and Figures

2-6. Members are small to moderate-sized

species (< 30 cm CL) with pronounced

sexual dimorphism which inhabit a variety of

lentic and lotic habitats through much of

India. The name is derived from “panshura>”

a Bengalese word for chelonian.

Kachuga smithii (Gray 1863)

Brown Roofed Terrapin — Plate III, A-C

Identification : A small species (to 23 cm

CL) identifiable from other Pangshura by a

relatively low, vaulted shell (H/CL < 44%)

having only a weak, horizontal spine (or

none) on the third vertebral scute.

Description : For coloration see descriptions

of subspecies. Head moderate in size with

short, tapering snout (less than length of

orbit) projecting beyond lower jaw; skin at

back of head divided into large irregular

scales; upper jaw serrate, lacking medial

notch or projection; alveolar surface broad,

decked by a finely-serrate ridge on each side,

1 Accepted December 1986.

x Dept, of Zoology, Eastern Illinois University,

Charleston, Illinois, U.S.A.

converging but not meeting at midline. Lower

jaw serrate with single, projecting tooth

anteriorly, alveolar surface concave except for

a median symphyseal ridge and a serrate ridge

along lingual surface meeting symphyseal ridge

at midline; coronoid process prominent.

Hyoid moderately developed; ossified portions

include a single-element body with a shallow,

rounded notch posteriorly and a shallow “V”

shaped notch anteriorly, a pair of small,

rounded elements attached at either side of

anterior notch (ceratohyals?), a narrow,

elongate, outwardly-bowed pair of first cera-

tobranchial horns and a pair of short (2-3

times longer than wide) second ceratobran-

chial horns.

Shell oval, widest across a plane through

seventh marginals; posterior margin of cara-

pace slightly serrate; median keel low; raised

areas at posterior of scutes not pronounced,

obtuse; Vertebrals 1, 3 and 4 usually longer

than wide, 2 and 5 usually wider than long;

seam contact formula — 1M 4> 6M 8<

10>. Bridge long, exceeding length of either

anterior (shorter) or posterior (longer) lobe

of plastron; axillary somewhat smaller than

inguinal scute; plastron truncate anteriorly;

notched posteriorly; plastral formula — Ab>

F> P> A> H> G.

Distribution : Brown roofed terrapins occur

in the Indus, Ganges and Brahmaputra Drain-

ages of Pakistan, Nepal, India, and Bangla-

7

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

desh. Figure 9 map the distribution in India

as verified by this survey.

Geographic Variation : Two subspecies (one

new) are recognized herein.

Kaclsuga smithii pallidipes subsp. nov.

Pale-footed Roofed Terrapin — PI. Ill, B&C

Holotype : Field Museum Natural Flistory

224177, adult male in alcohol; collected in the

Gandak River, Bherihari Wildlife Sanctuary,

Bettiah (West Champaran) District, Bihar on

3 June 1983; original number 2827 Edward

O. Moll.

Paratypes : IJSNM 257779, adult female

in alcohol; Kamali River, Royal Bardia Wild-

life Reserve, 2 km N. Thakurdara, Nepal,

23 April 1985 (obtained by Joseph Mitchell);

FMNH 224186, adult female skeleton, Ghagra

River, near Kailaspuri at Girija Barage,

Bahraich District, Uttar Pradesh, 12 January

1983.

Identification'. A subspecies of Kachuga

smithii differing from the nominate form by

the absence of a plastral pattern and a re-

duction of pigment on head, limbs, feet and

penis.

Description of holotype : An adult male

measuring 8.6 CL, 6.2 CW, 7.8 PL, 3.1 FI

and weighing 88 g. Coloration in life — cara-

pace light grayish olive to brownish olive

(older scutes) with pale yellow rim around

the periphery; single mid-saggital black stripe

with cinnamon-rufous center on Vertebrals 2

and 3. running length of the shell but becom-

ing obscure on V5 and 6; plastron straw

yellow, immaculate, having no dark pattern;

vague dark blotches present on ventral side

of marginals.

Head brownish olive, lightly mottled with

smoke gray dorsally; skin creamy white be-

hind eyes; iris pale gray; throat immaculate,

colorless; a pair of narrow dark stripes ex-

tend anteriorly from eyes converging to meet

at light gray snout; mandibles bright spectrum

yellow; neck smoke gray dorsally, colorless

ventrally, unstriped.

Limbs — ground color on leading face of

front limb smoke gray above elbow and late-

ral half of foreleg; large triangular scales on

lateral border of foreleg, narrow bandlike

scales on anterior aspect; toes and webbing

yellow; posterior aspect of limb, feet and

medial half of foreleg colorless.

Penis colorless lacking the dark pigment

characteristic of most members of the order.

Description of Paratypes: USNM 257779,

adult female measuring 15.6 CL 11.6 CW

15.5 PL 5.65 FI and weighing .453 kg. Colo-

ration of preserved specimen — carapace

brownish gray with a black, middorsal stripe;

plastron light, largely devoid of pattern but

tiny smudges of pigment present near poste-

rior lateral margins of scutes; bridge and

ventral side of Marginals 3-9 with heavy con-

centrations of dark pigment.

Head and neck dark grayish brown dorsally

with lighter cinnamon brown band running

posteriorly from eye over tympanum; throat

light, colorless.

Limbs — anterior face of forelimb and feet

generally grayish brown, becoming lighter on

medial aspect; lateral border of limb light

edged; hind feet immaculate, devoid of pig-

mentation; skin of leading face of hind leg

grayish brown.

FMNH 224186, female measuring 16.2 CL

11.6 CW 15.5 PL 6.0 H and weighing 0.63

kg. Coloration in life — carapace buff with

dark brown middorsal stripe; plastron straw

yellow, unpatterned.

Head and neck drab dorsally with lighter,

cinnamon-colored band running posteriorly

from eye over tympanum; tip of snout and iris

light grayish blue; mandibles light orange

yellow; neck dirty gray above, light cream

ventrally.

8

FRESHWATER TURTLES OF INDIA

Fig. 9. Distribution map of Kachuga smithii in India (see legend of Fig. 7). Solid

circles indicate localities of the nominate race, K. s. smithii. Open circles indicate

localities of K. s. pallidipes. The half open circle indicates Allahabad, a possible

intergrade site between the two races (see text).

9

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

Limbs with toes and skin flap at

lateral edge of hind foot immaculate, devoid

of pigment, otherwise outer leading surface of

fore and hind legs gray; trailing surface of

legs creamy white.

Distribution : The three members of the

type series of this subspecies come from north-

ern tributaries of the Ganges. See Figure 9

and comments on intergradation under

remarks.

Natural History : Little is known about the

natural history of this form. All three records

were associated with riverine environments.

The holotype was collected during the day in

a hoop trap baited with fish set near a sand

bank in the Gandak River. The Nepal speci-

men was collected by hand under a pile of

sticks and debris in a pool on the floodplain

of the Karnali River. The other female was

taken by a fisherman in the Ghagra River but

details are lacking. Remains of a freshwater

prawn were found in the gut of the latter

specimen.

Remarks : Presently it seems best to regard

this taxon as a subspecies of K. smithii rather

than as a distinct species for the following

reasons. 1) The male from the Gandak River

has been selected as the holotype or best re-

presentative of this race. The two female

paratypes from more westerly drainages show

some characteristics of the nominate form

(more dark pigment on the head, limbs and

ventral part of shell) suggesting intergradation.

One of the original types used by Gray

(1863) in the species description from the

Chenab River in northwestern India (BMNH

63.2.21.87) lacks pigmentation on the pecto-

ral scutes and only small amounts are present

on the abdominals. This too could represent

intergradation. More collections are needed to

confirm whether the observed variation has

resulted from intergradation rather than some

other factor such as sexual dichromatism. 2)

A precedent exists for this type of variation

in a related species. In what appears to be a

case of unusual parallelism, populations of

Kachuga tentoria from northern tributaries of

the Ganges also exhibit a reduction of dark

pigment and have no plastral pattern. More

collections are available for this species and

intergradation is evident in the more western

drainages of the Ganges (see K. tentoria).

A peculiar situation exists with a series of

specimens from Allahabad in Uttar Pradesh.

The Zoological Survey of India collection has

five specimens of K. smithii from this locality.

Three reportedly have the dark plastral

pattern (ZSI 457, 471, 472) and two do not

[ZSI 451, 200 (1912)]. Another specimen in

the British Museum (BMNH 1908.12.28.2)

from this locality has dark blotches only on

the anal scutes. Whether these specimens were

actually obtained in Allahabad or whether

they were obtained elsewhere and shipped to

markets there is not known. Neither is there

any information on the coloration of the rest

of the body in this collection. The entire

problem requires additional study.

Kachuga smithii smithii (Gray 1863)

Brown Roofed Terrapin — Plate III, A

Identification'. A subspecies of K. smithii

having a plastral pattern of large dark brown

to black blotches on each scute narrowly

bordered with yellow; sides of head, leading

surface of limbs, feet and penis dark pig-

mented.

Description : Female (FMNH 224143);

carapace brownish olive bearing a middorsal

dark brown stripe; a small dark triangle decks

areolar portion of Pleurals 2 & 3 and vertical

dark bars border seams between the more

posterior marginal scutes and the posterior edge

of Pleural 4; plastron, bridge and ventral side

of marginal chiefly dark but narrowly bor-

10

FRESHWATER TURTLES OF INDIA

dered with light yellow; head and neck olive

dorsally; a tawny blotch present behind eye;

vague striping evident on lateral portion of

neck; iris pale blue-gray; mandibles deep buff

yellow; skin on outer surface of limbs olive

with bandlike scales on forelegs appreciably

lighter than ground color; vague striping pre-

sent on hind legs and rump.

Size and Sexual Dimorphism : Minton

(1966) reported that eight females and three

males from Pakistan ranged from 15.3-22.7

CL and 10.1-10.8 CL respectively. Smith

(1931) recorded the largest specimen as 23.0

CL 15.5 CW and 8.5 H. This race was rarely

encountered on our survey. Six shells found

in garbage dumps near Rajmahal, Bihar rang-

ed from 13.2 to 18.3 (mean 15.7) CL. A

subadult female obtained from fishermen at

Kahalgaon, Bihar measured 14.3 CL 10.3 CW

13.7 PL 5.9 H and weighed 0.315 kg.

Males differ from females by being consi-

derably smaller and by having a longer tail

which is heavier at the base. Minton (1966)

states that the tail of males projects free about

10 per cent of the carapace length whereas

that of the female is about 5 per cent.

Natural History : Brown roofed terrapins

are typically associated with rivers and occur

in current as well as more lentic habitats such

as backwaters. Minton (1966) found them to

be a social basking species on the Indus

where they undergo a period of quiescence

from early December to early March. He

reported finding females with eggs in early

October; a clutch of seven laid by a captive

contained eggs 43 to 45 mm long and 22 to

24 mm wide. Chaudhuri (1912) reported that

five to eight eggs are buried in sand nests but

gave no season. Ewert (1979) reported the

mean size of four hatchlings to be 3.92 CL

and 3.67 PL. The species is generally report-

ed to be omnivorous with a carnivorous bias

(Das 1985, Minton 1966. Smith 1931). Gut

contents of the subadult female from Kahal-

gaon contained only plant material.

Distribution : The brown roofed terrapin

has been reported from the Indus, Ganges and

Brahmaputra Drainages of Pakistan, India and

Bangladesh. The Museum d’Histoire Natu-

relle in Geneve, Switzerland has specimens

catalogued as K. smithii from Assam but I

have not examined these. Minton (1966)

found the turtle to be common in the Indus

Drainage while Smith (1931) considered it to

be much rarer in the Ganges Drainage. Reza

Khan (1982) also reported that the species is

uncommon in Bangladesh. The species was

rarely encountered on our survey. Figure 9

maps localities verified for India.

Specimens were collected from the following

sites :

FMNH 224143 — Ganges River, Kahalgaon, c. 50

km W. Sahibgani. Bhagalpur District, Bihar.

EOM 2720-2725 — Ganges River, Rajmahal, Dumka

(Santhal Parghana) District, Bihar.

Type locality of this race is “North-

western India: Punjab; “River Chenab...”

Syntypes are BMNH 1947.3.4.69-70. Much

of the former Punjab is now part of Pakistan

and most of the Chenab River now found in

India is in Jammu and Kashmir rather than

the state of Punjab. Hence the type locality

could well be Pakistan. The dot on figure 9

is the western-most point of the Chenab in

India. Other preserved specimens verified in-

clude :

MCZ 3233 — Ludhiana, Punjab.

ZSI 17606 — Firozp-ur, Punjab.

Kachuga sylhetensis (Jerdon 1870)

Assam Roofed Terrapin

Identification'. A small terrapin (20 cm

CL) differing from other Pangshura by typi-

cally having 13 pairs of marginal scutes and

11

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

a fifth vertebral scute that is wider in the

anterior half than in the posterior.

Description : Head medium-sized with

slightly projecting snout; upper jaw slightly

hooked with serrations along the tomium be-

ing fine or absent; head patterned with a pair

of narrow yellow stripes running posteriorly

from eye to meet at mid-line on back of head

forming a chevron; an additional light stripe

runs along underside of the mandible curving

upward to meet tympanum; neck with light

longitudinal stripes.

Shell steeply peaked as in K. tecta and

tentoria but more serrate posteriorly; oval in

shape being widest across plane through sixth

marginals; median keel relatively narrow on

first through third vertebrals; sharp pointed

spines present on the third (largest) and

fourth vertebrals; Vertebrals 3 and 4 longer

than broad, V2 and 5 broader than long and

VI either as wide or somewhat wider than

long; scute contact formula: 1M 4> 6> 8M

11 <; coloration of preserved specimens olive-

brown with a lighter vertebral keel. Plastron

oval with slight notch or no notch between

anals; plastral formula F >< Ab > P >

H > A > G; inguinal and axillary scutes

well developed; pattern a large dark blotch

on each plastral scute, bridge, and ventral side

of marginals.

Size and Sexual Dimorphism : As typical for

the genus, males are much smaller than the

females. The largest specimen recorded is a

female 19.7 cm CL (Jerdon 1870). Measure-

ments of a typical male and female are:

BMNH 1929.11.21.1 M — 8.5 CL 6.8 CW

8.0 PL 4.7 H

BMNH 1947.3.4.22 F — 18.3 CL 14.3 CW

17.8 PL 8.4 H

In addition to being smaller, males have

longer tails which are thicker at the base than

those of females.

Natural History : Most specimens have come

from hill streams; nothing else seems to be

known of the natural history.

Distribution : The Assam roofed terrapin

has been reported from the Khasi, Garo and

Naga Hill areas of Bangladesh and Assam.

As these areas were off limits to our survey,

no living specimens of this species were seen

or collected. I have examined museutn speci-

mens from the following localities:

BMNH 1947.3.4.22 (type) — Khasi Hills, Sylhet

District, Bangladesh.

BMNH 1929.11.21.1 — Khasi Hills, Cherrapunji,

Meghalaya, India.

ZSI 110 — Cachar District, Assam, India.

ZSI 3923 — Garo Hills, Assam, India.

Kachuga tecta (Gray 1831b)

Indian Roofed Terrapin — Plate III, D-F

Identification'. A small Pangshura (23 cm

CL) with a high vaulted shell (height/ length

> 45%) most easily distinguished from its

closest relative, K. tentoria, by its pattern and

brighter coloration. Plastral pattern of small

dark blotches or streaks (1-4) on most scutes;

head with a large red to orange crescent-

shaped blotch behind eye; neck with bright

yellow stripes and limbs bearing bright yellow

spots. Smith (1931) used as a key character

that the second vertebral is longer than the

third in tecta but not in tentoria. Although

useful, this character is variable being correct

in only 76 percent of the 21 K. tecta and only

63 percent of the 45 K. tentoria examined.

Description : Sexes colored similarly (live

female, Lucknow, U.P.); carapace raw umber

(brown) with somewhat lighter middorsal

stripe (can also be red) bordered in black and

a narrow yellow border on marginals; plastron

buff-yellow with 2-4 small, round to elongate,

black markings on scutes other than gulars

and/or anals which may have only one; two

dark spots on bridge, one each on inguinal

and axillary scutes and on underside of each

12

FRESHWATER TURTLES OF INDIA

Fig. 10. Distribution of Kachuga sylhetensis (see legend of Fig. 7).

13

JOURNAL, BOMBAY NATURAL HIST. SOCIETY , Vol. 84

marginal; head dark with large orange yellow

to spectrum orange crescent-shaped, post-

ocular blotch curving upward from below eye

and extending obliquely across head to meet

and form chevron with blotch from opposite

side; iris dark; mandible orange yellow be-

coming orange beneath snout; neck dark,

patterned with yellow stripes being brightest

laterally and ventrally; outer surface of limbs

dark, studded with bright orange yellow spots

on scales; rump with vertical orange yellow

stripes on lighter background.

Shell oval, steeply pitched, widest at plane

through sixth or seventh marginals, posterior

edge slightly serrate; middorsal keel widest on

first three vertebrals, a prominent pointed

spine present on Vertebral 3; VI widest in

anterior half, usually as long or slightly longer

than wide; V4 flask-shaped, much longer than

wide; V5 widest in posterior half and wider

than long; V2 and 3 vary in length-width

relationships; scute contact formula — 1>

4M 6M 8M 10M. Plastron truncate anteriorly

and notched posteriorly; anterior lobe shorter

than posterior, both shorter than bridge; plas-

tral formula-Ab >< F > H> A> P> G;

axillary and inguinal scutes large, subequal

in length.

Head moderate; snout shorter than orbit;

skin at back of head divided into a series of

irregular scales; secondary palate broad with

single denticulate ridge; edge of tomium coar-

sely serrate and lacking medial notch. Lower

mandible ending in prominent medial tooth;

alveolar surface concave except for a serrate

ridge along lingual border which joins a

symphyseal ridge at midline; moderate coro-

noid process present. Hyoid moderately deve-

loped; ossified portion comprising a single-unit,

elongated body with a broad shallow notch

posteriorly and a deeper, narrower notch

anteriorly; a narrow, curving, elongated pair

of first branchial horns and a pair of small

kidney-shaped second branchial horns.

Size and Sexual Dimorphism ; Females are

much larger than males. The largest specimen

recorded is 23 cm CL (Smith 1931). Minton

(1966) reported two adult females and a male

from Pakistan measured 16.4, 17.3 and 8.4

cm CL. Measurements of two females and

one male examined in this survey are:

Live F — 15.3 CL 11.5 CW 14.7 PL 7.3 H

weight 0.51 kg.

EOM 2784 (BNHS uncataloged) F — 18.3

CL 14.2 CW 17.2 PL 7.3 H weight 0.96 kg.

Live M (mature?) - 6.6 CL 5.4 CW 6.3 PL

3 . 7 H weight 0 . 054 kg.

In addition to size males differ from

females by having a longer, thicker tail in

which the vent opens beyond the carapacial

rim.

Hatchlings-. Four hatchlings from eggs laid

by a female at Lucknow, Uttar Pradesh averag-

ed 3.15 CL 2.8 CW 2.8 PL 1.8 H and 7 grams

weight. Young K. tecta are strikingly attractive

turtles. The carapace is bright lime green rimm-

ed with a yellow orange border. There is a

broken middorsal stripe of flame scarlet bor-

dered in black and each pleural has a tiny

black spot at the posterior dorsal edge where

a lateral keel would be. The plastron is chrome

orange patterned with small, irregular, black

blotches on each scute including the axillary,

inguinal and underside of the marginals. The

head and neck are dark olive in ground color

with the latter decked with bright yellow

stripes. A large crescent-shaped flame scarlet

blotch begins under the eye, curves upward

behind the eye and meets its counter part from

the opposite side at the back of the head

forming a “V” shaped figure. A small flame

scarlet spot marks the posterior of each eye

lid. The scales of the limbs as well as the

webbing between the toes are bright yellow

on a dark olive background.

14

FRESHWATER TURTLES OF INDIA

Natural History : Indian roofed terrapins

chiefly inhabit lentic habitats (tanks, nullahs

and backwaters) in the Ganges and Indus

drainages. Slow moving or quiet vegetation-

choked waters appear optimal. We observed

this species in a weedy backwater of the

Ghagra River, a small impoundment pond

near Lucknow, U.P. and in a slow moving

nullah crammed with aquatic vegetation which

flowed into a nearby ox bow lake in the

Udaipur Forest Area of Bihar. Khan (1982)

reported K. tecta occupying flowing and stag-

nant waters in Bangladesh. In the Narmada

River where K. tentoria and K. smithii are

seemingly absent, we found K. tecta to be

moderately common.

Like its close relative the Indian tent terra-

pin, the Indian roofed terrapin is commonly

seen basking on logs or the river bank.

Parshad reported that it is herbivorous (Smith

1931). However, we caught one specimen in a

hoop trap baited with chicken entrails.

A specimen from Lucknow laid a clutch of

eight eggs on January 13. The eggs averaged

37 x 21 mm and 10.75 grams. Relative to

the female reproductive efforts, calculations

are: RCM — 0.21, EMI — 2.1, ELI — 24.2

and EW1 — 14.

Distribution : The Indian roofed terrapin

definitely occurs in the Indus, Narmada,

Ganges and Brahmaputra River Systems of

Pakistan, India and Bangladesh. Based on the

Indian distribution, it probably occurs in

Nepal as well. A series of specimens in the

Museum of Comparative Zoology (MCZ

3459, 3460, and 3462) labelled Rangoon, if

verified would extend the range much farther

East. Figure 11 depicts the distribution in

India as verified by the survey:

Live F — Kukrail, nr. Lucknow, Lucknow District,

India.

Live M — Harhi Nala, 15 km W Bettiah. Bettiah

Dist,, Bihar.

EOM 2658 — 6 km S Katarnia Ghat, nr. Girija

Barage, Bahraich District, U.P.

EOM 2868 — Bedaulia, Manika, Muzaffarpur Dis-

trict, Bihar.

EOM 2784 (uncataloged specimen BNHS) — Nar-

mada River, Dhavdi Ghat, nr. Punasa, East Nimar

District, M.P.

In addition the following preserved speci-

mens have been examined and verified :

BNHS 1290-1291 — Chandola Lake, nr. Ahmedabad,

Ahmedabad District, Gujarat.

ZSI 17609 — Makhu, Firozpur District, Punjab.

ZSI 21672 — Magwall Village, Jammu District,

Jammu-Kashmir.

Uncatalogued specimens BNHS — Hindon River,

Mohen Nagar, nr. Ghaziabad, Meerut District, U.P.

ZSI 18015 — Baradighi Tea Estate, Jalpaiguri Dis-

trict, West Bengal.

ZSI 19236 — Cherrapunji, Meghalaya, India.

Remarks : The Indian roofed terrapin is the

only Kachuga to be listed on CITES (Appen-

dix I) or the Indian Wildlife (Protection) Act

(Schedule I). Reasons for listing are obscure;

we found the species to be relatively common

in the aforementioned sites and it did not

appear to be a popular market species. Khan

(1982) judged it to be the most common

turtle in Bangladesh. Nevertheless because of

the turtle’s status, the aforementioned pre-

served specimens were either taken as shells

or in the case of EOM 2784, a turtle which

drowned in a fisherman’s net. All specimens

from the survey were left within the country.

Kadiuga tentoria (Gray 1834)

Indian Tent Terrapin — Plate II

Identification : A moderate-sized Pangshura

(27.1 cm CL) with a high, vaulted shell

(height /length > 45%); differing from K.

tecta by having one or two small reddish to

brownish spots behind the eye instead of a

broad crescentic band and by having a plastron

with a single large dark blotch per scute or

Jacking in dark markings.

Description : Shell oval being widest at a

15

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

Fig. 11. Distribution of Kachuga tecta in India (see legend of Fig. 7).

E F

(A) Kachuga tentoria tentoria — Juvenile (8.1 cm CL) from the Godavari River near Manthani, A.P.

(B) Kachuga t. tentoria — Ventral view of A. (C) K. t. flaviventer — Male (8.3 cm CL) from the Gandak

River, Bettiah District, Bihar. (D) K. t. flaviventer — Ventral view of C. (E) K. t. circumdata — Male

(8.5 cm CL) from the Yamuna River nr. Etawah. (F) K. t. circumdata — 1 Male (8.1 cm CL) and female

(23.5 cm CL) from the Yamuna River near Etawah.

Plate II

J. Bombay nat. Hist. Soc. 84(1)

Moll: Freshwater Turtles

E F

(A) Kachuga smithi smithi — Female (14.2 cm„CL) from the Ganges near Kahalgaon, Bihar. (B) K . smithi

subsp. nov. — Male (8.6 cm CL) from the Gandak River, Bettiah District, Bihar. (C) K. smithi subsp,

nov. — Ventral view of B. (D) K. tecta — Immature female (6.4 cm CL) from near Bettiah, Bihar.

(E) K. tecta — Hatchling (3.0 cm CL) from Lucknow, U.P. (F) K. tecta — Ventral view of E.

Plate III

J. Bombay nat. Hist. Soc. 84( 1)

Moll: Freshwater Turtles

FRESHWATER TURTLES OF INDIA

plane passing through the seventh marginals;

a prominent middorsal keel runs the length

of the carapace breaking at the end of each

vertebral to form a knob or spine, the most

pronounced being a sharp upward projecting

spine on V3; in adults Vertebrals 3 and 4

typically longer than wide with 5 being wider

than long; VI and 2 variable, with 1 often

being hour glass or bell shaped (pinched in

the middle) in large individuals; seam contact

formula — 1> 4> 6M 8M 10M. Plastron

truncate anteriorly, notched posteriorly; plas-

tral formula — AB > F > P > H > A > G;

bridge long exceeding lengths of both the

shorter fore lobe and longer hind lobe of

plastron; axillary somewhat smaller than in-

guinal scute. Cloacal bursae present with pro-

nounced villous lining.

Head medium-sized with short, pointed,

projecting snout; skin at back of head divided

to form a series of irregular-shaped scales;

upper jaw serrate lacking median notch or

prominent projections; alveolar surface broad,

bearing single “V” shaped denticulate ridge.

Lower jaw similarly serrate with single, pro-

minent, projecting tooth at apex; lower alveo-

lar surface concave bordered by serrate ridge

along lingual surface meeting a short symphy-

seal ridge at midline. Hyoid moderately deve-

loped with ossified portions including a single-

element body having a prominent rounded

notch posteriorly and a smaller “V” shaped

notch anteriorly, a pair of thin, elongate, out-

wardly bowed first ceratobranchial horns and

a pair of small, rounded second ceratobran-

chial horns.

Distribution : The Indian tent terrapin is

restricted to drainages of rivers flowing into

the Bay of Bengal in India, Nepal and Bangla-

desh. Figure 12 depicts the distribution in India

as verified by the survey.

Remarks: Until recently K. tentoria has

been considered a subspecies of K. tecta.

However, as both appear to be sympatric

over a broad geographic area including Bangla-

desh (Khan 1982), Bihar, and Uttar Pradesh

(this paper); herein I follow Pritchard (1979)

in regarding them as separate species pending

additional study. Smith (1931) lists the type

locality for K. tentoria as Dhond (Krishna

River Drainage), Poona District of Maha-

rashtra. See Mertens (1969) for a history of

the nomenclature of these two species. -

Three subspecies of K. tentoria (one re-

surrected) are recognized herein. — Kachuga

t. tentoria in the rivers of peninsular India;

K. t. circumdata in the western and central

drainage of the Ganges and K. t. flaviventer

in the eastern Ganges and its northern tribu-

taries.

Kachuga tentoria tentoria (Gray 1834)

Indian Tent Terrapin — Plate II, A+B

Identification : A race with a dark plastral

pattern, no pleuro-marginal ring and reddish

head markings.

Description : Sexes colored similarly (FMNH

224163 juv., Godavari River); carapace anti-

que brown, unicolor except for hazel to amber

stripe along middorsal keel from Vertebrals

1-3; plastron yellow with large dark blotches

on each scute, bridge, axillary, inguinal and

underside of marginals; ground color of head

olive to brownish olive; a poorly defined clay

band present behind eye; red markings in-

clude a small red postocular spot in clay band,

a smaller red mark located at dorsal posterior

edge of eye and a thin poorly defined red line

in occipital region; iris gray olive; mandibles

straw yellow; neck with dull straw yellow

stripes on lateral and ventral portions; limbs

olive with edges of scutes cream; rump mark-

ed with vertical black and cream stripes.

Size and Sexual Dimorphism : Males are

much smaller than females. Three males from

the Mahanadi River measured 8.1, 9.7 and

17

2

JOURNAL , BOMBAY NATURAL HIST. SOCIETY, Vo I. 84

Fig. 12. Distribution of Kaehuga tentoria in India (see legend of Fig. 7). Starred

circles indicate localities of the nominate race, K. t. tentoria. Solid circles indicate

localities for K. t. circumdata. Open circles indicate localities for K. t. flaviventer.

FRESHWATER TURTLES OF INDIA

10.9 cm CL. Another 7.5 cm CL with no

sperm in the epididymides was judged sub-

adult. Six females from this locality ranged

from 19.2 to 23.0 (mean 20) cm CL. Two

other females 18.5 and 18.0 cm CL with no

enlarged follicles on their ovary and relatively

small oviducts were judged subadult.

Proportions of a typical male and female

of this race follow:

BNHS 1329 F — 19.5 CL 14.2 CW 18.9 PL

8.9 H weight 0.86 kg.

FMNH 224141 M — 9.7 CL 7.5 CW 9.2 PL

4.6 H weighing 0.105 kg.

In addition to size, males differ from females

by having a longer (preanal > postanal) and

thicker preanal portion of the tail with a vent

opening beyond the edge of the carapace.

Natural History. Indian tent terrapins occur

in small to large rivers of peninsular India.

They are frequently seen on logs and rocks

basking. Females appear completely herbi-

vorous. They never entered hoop traps baited

with chicken entrails or fish and other than a

small feather, the guts of four individuals from

the Mahanadi River contained only leaves and

stems of vascular plants. Males and juveniles

appear more omnivorous. They did enter bait-

ed hoop traps and the stomach of one male

examined contained 75 percent vegetation and

a fresh water prawn. Ovaries of two females

collected on the Mahanadi River in mid-

February appeared post-reproductive having

few enlarged follicles and several small, old

corpora lutea.

Distribution : Kachuga t. tentoria ranges

from at least the Mahanadi River drainage

southward to the Krishna drainage. The

turtle was taken at the following localities on

the survey:

FMNH 224163 — Godavari River, Manthani,

Karimnagar Dist., A.P.

Live Juvenile — Godavari River, Polavaram, West

Godavari Dist., A.P.

FMNH 224141, BNHS 1329 — Mahanadi River,

Tikarpura, Dhenkanal, Orissa.

The following additional records have been

verified from preserved collections:

ZSI (Type Kachuga t. intermedia Blanford 1870)

— Hasdo River, Bilaspur Dist., M.P.

ZSI 17775 — Seonath River, Bilaspur, Bilaspur Dist.,

M.P.

ZSI 16767 & 68 — Mahanadi River, Cuttack, Cuttack

Dist., Orissa.

Kachuga tentoria flaviventer (Gunther 1864)

Plain-bellied Tent Terrapin — Plate II, C & D

Identification: A small tent terrapin (20.3

cm CL) with reduced pigmentation, an un-

patterned plastron and little or no striping on

neck and rump.

Description : Sexes colored similarly. Male

(FMNH 224178) — carapace brownish olive;

light middorsal stripe with pale orange wash

on Vertebrals 1 and 2, cream on V3, becoming

faint on V4 and 5; pleuro-marginal juncture

and border of shell also cream; plastron,

bridge and underside of marginals cream and

unpatterned; head pale, mottled with brownish

olive; a near colorless patch washed with pale

salmon extending from behind eye back over

mastication musculature to meet patch from

opposite side; immediately posterior a dark

horizontal line marks juncture of head and

neck; neck colorless to pale cream; a small

irregular splotch of cinnamon rufous occurs in

colorless area immediately behind eye; iris

light smoke gray; limbs almost colorless ex-

cept for a scattering of dark pigment along

leading face; webbing and underside of feet

creamy.

Female (FMNH 224132) colored as above

with the following exceptions: carapace light

buff or cinnamon ground color; the central

stripe being a darker, tawny coloration with

a lighter center; plastron unpatterned but dark

blotches present on underside of marginals;

head smoke gray dorsally, cream laterally and

on mandibles; a light cinnamon -brown spot

19

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vo 1. 84

behind eye and another at posterior dorsal

edge of eye; three additional spots at posterior

of head (one medial flanked by two lateral);

webbing of feet pale yellow.

Size and Sexual Dimorphism : Sexes widely

disparate in size. A female and two shells pre-

sumed to be female from Kahalgaon measured

16.5, 16.9 and 18.8 CL. Five males from the

Gandak River in northwestern Bihar showing

well developed secondary sex characters rang-

ed from 6.6 to 8.8 (mean 8.0) CL. Measure-

ments of two typical specimens are:

FMNH 224132 F — 16.5 CL 12.4 CW 16.1

PL 7.9 H weight 0.64 kg.

FMNH 224178 M — 8.2 CL 6.3 CW 7.7 PL

4 . 1 H weighing 0 . 09 kg.

In addition to size males differ from females

by having a longer tail (preanal > postanal

portion) which is relatively thicker at the base.

Hatchlings’. Vijaya (1982e) provided mean

measurements for hatchlings from six clutches

of the pale-bellied tent terrapin (see com-

ments under geographic variation) laid in nests

along the Rapti River near Gorakhpur, U.P.

Largest and smallest of these means were:

Clutch I — 2.7 CL 1.7 CW 2.3 PL 1.6 H

and 6.5 g weight. Clutch V — 3.35 CL 2.7

CW 3.1 PL 1.8 H and 10 g weight. Shell

coloration — carapace mottled light and dark

olive with a light yellow stripe along the

middorsal keel; pleuro-marginal ring light

geranium pink at hatching fading in the first

few months to a pale olive; plastron buff

yellow and unmarked; underside of marginals

with a pepper like dusting of dark pigment;

head ground color pale olive; two flesh pink

spots located posterior to eye and at posterior

of eyelid respectively and a narrow, flesh

pink bar, broken in the middle, at back of

head; vague striping discernible on neck and

rump.

Twelve hatchlings obtained from three

clutches of eggs collected at the Katarniaghat

Gharial Sanctuary in U.P. had the following

mean dimensions: 3.55 CL 3.23 CW 3.2 PL

2.0 H and 9.0 g weight. Their shells were

slightly serrate posteriorly with carapacial

spines being very small except for a prominent

projection on V3. VI was broadest anteriorly

lacking the pinched bell-shape of large adults.

None had a plastral pattern but varied as to

the presence of dark pigment on the under-

side of the marginals. Amounts varied from

none to having a dark blotch on each scute.

Another variable feature was the amount of

red in the pattern. Some had an extensive

amount including a geranium pink pleuro-

marginal ring, middorsal stripe and head

pattern. Concerning the latter at one extreme

some individuals had both a transverse pink

line across the back of the head, two pink

spots at the snout and a postocular, pinkish

crescent behind the eye. The crescent (similar

to that of K. tecta but less extensive) was

formed by an elongation of the postocular

spot to meet the elongated one at the rear of

the eyelid (this was evident in other indivi-

duals in which the merger was incomplete).

At the other extreme were individuals with

no pink whatsoever; the pink areas of other

individuals were colorless.

Natural History : Like the other races of

this species the pale-bellied tent terrapin ap-

pears to be chiefly a river turtle. We found

this race in Katarniaghat Gharial Sanctuary,

an impoundment of the Ghagra River, but

otherwise all were seen or collected in areas

of flowing water. Many were observed bask-

ing near the bank in a sandy area of the

Gandak River in May. Gut contents of one

male and one female examined contained only

leaves and stems of aquatic vegetation. How-

ever, five males were taken in traps baited

with chicken entrails and fish suggesting that

they may be somewhat omnivorous.

Vijaya (1982e) found nests of this turtle

20

FRESHWATER TURTLES OF INDIA

in “soft, clayey river bank soil” along the

Rapti River near Gorakhpur from 6-8 Dec-

ember. Nests varied from 15 to 26 cm in depth

and were located 3 to 14 metres from the

water. Six nests contained from 4 to 8 (mean

6) eggs per nest. She provided mean egg sizes

for each nest ranging from 41 x 29 mm and

1 1 . 7 g to 45 x 27 mm and 18.5 g. Incubation

times (time to emergence from substrate?) in

artificial nests ranged from 125 to 134 days at

nest temperatures varying between 27° and

28°C.

On this survey we found three clutches

comprising 6, 7, and 10 eggs in nests made by

this turtle in sand banks along the Ghagra

River in the Katarniaghat Gharial Sanctuary

on December 5. Mean size of the eggs was

42 x 25 mm and 15.3 g weight. Mean in-

cubation time for 12 hatchlings (to emergence

from egg) was 95 days from eggs kept on

moist cotton in plastic boxes at ambient tem-

peratures ranging from 24° to 33 °C.

Distribution : The pale-bellied tent terrapin

inhabits the northern tributaries of the Ganges

and possibly the Ganges proper from Bihar

eastward. So far I have examined no speci-

mens from West Bengal or Bangladesh. Speci-

mens collected on the survey are from the

following localities.

FMNH 224142 — Katarniaghat Gharial Sanctuary,

Ghagra River, Bahraich Dist., U.P.

Hatchlings (Vijaya 1982e) — Rapti River, nr.

Gorakhpur, Gorakhpur Dist., U.P.

BNHS 1339 & FMNH 224178 — Gandak River,

Bherihari Wildlife Sanctuary, Bettiah (West Cham-

naran) Dist., Bihar.

FMNH 224132 — Confluence of Kosi and Ganges

River, Khalgaon. c. 50 km W Sahibgani, Bhagal-

pur Dist., Bihar.

Remarks : Gunther (1864) recognized this

taxon as distinct and described it as a new

species Pangs hura flaviv enter. The description

was based on a single specimen presumed to

be from India but lacking in precise locality

data. It had been collected by a Mr. Mc-

Clelland who had also sent several other speci-

mens of Bengal species. Subsequent authors

(e.g. Boulenger 1889, Smith 1931) considered

it a variant of Kachuga tecta. Now that more

specimens have been found, it is evident that

this is a legitimate taxon. However, it appears

to be a subspecies of Kachuga tentoria rather

than a separate species. Evidence for this

comes from the aforementioned hatchlings

examined from the Rapti and Ghagra Rivers.

Characteristics of these specimens particularly

those from the Ghagra River appear to be

intergrading with those of K. tentoria circum-

data. The presence of circumdata characters

(i.e. the pink pleuro-marginal ring and pink

head markings) on some but not others of this

group is typical of an intergrading popula-

tion. The subject requires more study but for

now I believe a subspecies designation best

fits the evidence.

Kachuga tentoria circimndata (Mertens 1969)

Pink-ringed Tent Terrapin — Plate II, E+F

Identification : A moderate-sized tent terra-

pin (to 27.1 CL) having a single large dark

blotch on all plastral scutes, a reddish ring

at pleuro-marginal juncture of carapace, and

reddish head markings.

Description : Males appear somewhat darker

than females otherwise sexes colored similarly

(FMNH 224162 M) — carapace olive-green

with a geranium pink ring around pleuro-

marginal juncture; middorsal stripe comprised

of streaks of geranium pink bordered by

black; plastron straw yellow with large dark

blotch covering over half of each scute; bridge

and underside of marginals black bordered in

yellow; inguinal with black spot but not

axillary; seams of plastral scutes and margi-

nals washed with pink; head olive green with

a geranium pink circular spot behind eye and

a pair of short, oblique, geranium pink bars at

posterior of head; iris smoke gray; mandibles

21

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

light straw yellow with orange wash becoming

olive near snout; neck olive gray with dull,

cream colored stripes on sides and venter;

rump and base of tail also striped (stripes

more pronounced than in other races).

A female (BNHS 1340) was similar but

differed as follows: Carapace antique brown

with a cinnamon-rufous rather than pink

pleuro -marginal ring; middorsal stripe vague,

almost nonexistent; head markings less red

and more cinnamon rufous; two irregular

spots (instead of bars) present at back of

head.

Size and Sexual Dimorphism : Females great-

ly exceed males in size. A collection of thirteen

females from the Chambal River in the

Morena District of M.P. ranged from 18.1 to

27.1 (mean 22.1) cm CL. A sample of 11

males from the Yamuna River in the Etawah

District of U.P. ranged from 7.7 to 8.8 (mean

8.3) cm CL. Measurements of a typical male

and female are:

FMNH 224162 M — 8.4 CL 6.7 CW 7.8 PL

4.6 Hand 0.0825 kg.

Live F — 23.3 CL 17.2 CW 22.5 PL 11.2

H and 1.6 kg.

In addition to size males differ from females

by having a longer tail with a proportionately

thicker base.

Hatchlings : A single hatchling from an egg

obtained at Deogarth Ghat, on the Chambal

River measured 3.7 CL 3.5 CW 3.4 PL 2.1

H and weighed 11.5 g. Coloration — carapace

smoke gray with vague dark mottling on most

scutes with a cinnamon-rufous pleuro-marginal

ring; anterior portion of middorsal stripe also

cinnamon-rufous bordered with black; plastron

straw yellow with large, black blotches cover-

ing most of the scutes and light cinnamon-

rufous along scute seams; head olive with

bright geranium pink markings including a

spot at posterior edge of upper eyelid and

adjacent skin, a larger postocular spot and a

transverse bar across back of head; chin with

three additional pink spots along lateral edge

of mandible; iris light gray; neck with cream

stripes on an olive gray background; limbs

gray with cream-colored scutes, webbing and

lateral skin flaps; rump with dark and cream

stripes. See Moll (1985) for a colored photo-

graph.

Natural History : The pink-ringed tent ter-

rapin is a riverine form which readily basks

at any opportunity. We observed hundreds of

these turtles basking on logs, islands and

along the banks of the Yamuna and Chambal

Rivers. In January when the larger Kachuga

were scarce and presumably dormant, this

species was still basking in some number along

the Chambal. There is some indication of habi-

tat separation between the sexes and age

groups — small males and juveniles were rare-

ly seen in the river proper where females

were common. However, in a backwater

behind a sandbar projecting out from the

shore of the Yamuna River, we once collected

11 males, 6 juveniles and 3 females. Three

yearlings were also captured in a small (10'

wide) weedy tributary of the Chambal hiding

in and around vegetation masses.

Digestive tracts of one male, one female

and two immature females were examined.

The male contained a beetle and an equiva-

lent amount of aquatic vegetation whereas the

female guts were packed with vegetation alone.

This supports findings from the other races

indicating that females tend to be chiefly herbi-

vorous while males are more omnivorous.

Nesting was occurring along the sandbanks

of the Chambal River when we arrived there

on 19 January but all nests found had been

destroyed by predators. Jackal tracks and

occasionally hyena tracks were associated with

these nests. Two eggs, one cracked and one

entire remained in one of the open nests. The

cracked egg measured 47 x 28 mm while the

FRESHWATER TURTLES OF INDIA

entire egg was 47 x 27 mm and weighed 19.5 g.

Rao and Singh (1985) reported that nesting

occurs from October through January in the

National Chambal River Gharial Sanctuary.

They calculated the turtles’ average clutch size

as 6 (3-12) eggs and thought two clutches

to be typical. Mean egg size was 48.6 x 27.6

mm and 21 g. Using data from eight of their

females, I caculated the mean female repro-

ductive effort as: RCM — 0.08, ELI — 22,

EWI — 12, and EMI — 1.4.

Distribution : The pink-ringed tent terrapin

occurs in the upper and central Ganges and

such tributaries as the Hindon, Yamuna,

Chambal and Gomati. Mertens (1969) named

the type locality for this race as Meerut,

Meerut District, U.P. He was not confident

about the localities of his paratypes which

were listed as Calcutta or vicinity of Calcutta.

Based on our observations from this survey,

it seems likely that Merten’s paratypes were

market specimens shipped in from more

western states. Because of the heavy market

trade in turtles of West Bengal, distribution

records from this state are often unreliable.

More likely the race of tent turtles which

occurs naturally in West Bengal is K. t. flavi-

venter or some as yet undescribed popula-

tion. Specimens of this race were collected at

the following localities on our survey:

FMNH 224105 & 224109 — Hindon River, nr.

Ghaziabad, Meerut Dist., U.P.

BNHS 1335 & FMNH 224162 — Yamuna River,

5 km S Etawah, Etawah Dist., U.P.

Refe

Acharji, M. N. (1950): Edible chelonians and

their products. J. Bombay nat Hist. Soc. 49: 529-

532.

(1955): A collection of chelo-

nians and snakes from Chota Nagpur, Bihar. Rec.

Indian Mus. 53: 383-392.

Anderson, J. (1876): On the cloacal bladders

and on the peritoneal canals in Chelonia. J. Linn.

Soc. London. 12: 434-444.

BNHS 1 340 — Chambal River, Deogarth, 30 km NE

Morena, Morena Dist., M.P.

FMNH 224185 — Gomati River, nr. Lucknow,

Lucknow Dist., U.P.

Acknowledgements

Many officials and individuals aided in the

surveys from which these observations were

made. I am particularly indepted to Shri Samar

Singh, Joint Secretary for Wildlife, and his

staff for providing introductions and for help-

ing me to obtain the necessary permits in the

states visited. My colleagues and companions

on the survey, Ms J. Vijaya and Satish

Bhaskar provided invaluable assistance on all

aspects of the study. Mr. J. C. Daniel, Shri P.

Kannan and Shri S. Biswas provided consi-

derable useful advice based on their extensive

knowledge of the Indian chelonian fauna. My

thanks to Joseph Mitchell who kindly pro-

vided information on the Kachuga smithii

specimen used as a paratype in this paper and

to the U.S. National Museum for loaning me

the specimen. A special thanks to Rom and

Zai Whitaker who provided living and labo-

ratory space for me at the Madras Crocodile

Bank.

The project was funded through an Indo-

American Fulbright Fellowship and by a grant

from the New York Zoological Society. Bom-

bay Natural History Society kindly served as

my host institution in India.

EN CES

(1878-79): Anatomical and zoolo-

gical researches and zoological results of the Yunnan

Expeditions. Reptilia and Amphibia. Calcutta.

Boulenger. G. A. (1889): Catalogue of the

chelonians, rhvnchocephalians and crocodiles in the

British Museum (Natural History). Taylor and

Francis, London.

Carr, J. L. (1981): Phylogenetic implications of

chromosomal variation in the Batagurinae (Testu-

23

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demic Press, London.

25

THE BUTTERFLIES OF THE NILGIRI MOUNTAINS OF

SOUTHERN INDIA

(LEPIDOPTERA: RHOPALOCERA ) 1

Torben B. Larsen2

This paper is an account of the three hundred or so butterflies to be found in

the Nilgiri Mountains of southern India. Special emphasis is placed on placing the

butterflies in perspective in relation to a number of defined ecological zones. Follow-

ing a description on the climate, topography and vegetation zones in the Nilgiris is

a section on the history of entomological exploration. The major part of the paper

is devoted to a summary account of each species known to occur in the Nilgiris.

Every attempt has been made to update the nomenclature which is cross-referenced

to that of Wynter-Blyth (1957) whose nomenclature actually dates back to the

1930ies. Since this nomenclature is the same for most of peninsular India and since

the Nilgiris contain populations of virtually all species known to occur in southern

India, the paper should also be useful out of the specific Nilgiri context. The data

included will be the basis for a later analysis of the ecological and zoogeographical

nature of the Nilgiri butterflies, but some

discussed.

Introduction

The Nilgiri Mountains

The Nilgiri Mountains are situated in South

India with their centre at 11°25'N and 76°45'

E. The name means Blue Mountains and was

bestowed upon them by the plains people at

least 700 years ago when the Nilgiris were

only seen rising in the distance from the

steaming and insalubrious jungles that sur-

rounded them.

The Nilgiris are a well-defined massif that

forms the southern limit of the main Western

Ghats system that stretches unbroken from

Bombay in the north. To the immediate south

of the Nilgiris is the Palghat Gap, a stretch

of dry lowlands separating the South Sahyadri

from the main Ghats system. The distance is

not great and on a good day the nearest of the

1 Accepted January 1987.

2 Snoghoj alle 29C, 2770 Kastrup, Denmark.

initial non-quantitative conclusions are

South Sahyadri ranges, the Annamalais, is

clearly visible from the Nilgiri plateau.

To the northeast the precipitous Moyar

Gorge creates a narrow boundary between the

Nilgiris and the Biligiriranga Mountains. The

latter can be looked at as the southwestern

link, though at best a tenuous one, with the

Eastern Ghats system.

The Nilgiris may be described as a right-

angled triangle with the right angle placed in

the northwestern extremity. The western

slopes of the main plateau rise abrupt-

ly from levels of 100 to 300 m through a

steep escarpment to 1800 m or so. At this level

the Nilgiri Plateau commences. Though less

precipitous the same situation pertains to the

southern slopes. The northern slopes rise from

the Mysore Plateau from levels of 700-900 m

in a less precipitous manner still, and they are

hence rather less imposing. However, the

Nilgiris most certainly are a sharply defined

geographical feature.

26

BUTTERFLIES OF THE NILGIRI MOUNTAINS

The plateau of the Nilgiris is hardly a

plateau in the strict sense of the word. Valleys

drop to 1600 m and peaks rise to more than

2500 m, but most of the area is an undulating

landscape confined within the 1800 m contour

of the scarp and technically the expression

plateau is correct (though slightly disconcert-

ing to an author from a country in which the

highest point is some 180 m and lovingly

known as the Mountain of Heaven).

Just to the north of the Nilgiris proper lie

the Ouchterlony Valley and the Wynaad with

an average elevation of 1200-1300 m, provid-

ing level country at an altitude not found

elsewhere in the range. They form an integral

part of the Nilgiris as well as the link with

the Western Ghats proper and will be includ-

ed in the systematic part of the paper.

When making ones way through the Nilgiris,

whether by car or by foot, it is easy to forget

how small a geographical feature they actually

are. The western slopes extend some 40 km

due south, the southern and the northern

slopes are both some 60 km. As will be seen

there can be few areas of similar size with

as much ecological variation within its boun-

daries.

The area covered by this paper is the Nilgiris

as a geographical feature, from the surround-

ing lowlands to the highest peaks. Most of this

falls into the administrative Nilgiri District

(PIN 643 000) of Tamil Nadu State, but much

of the lower parts of the western slopes fall

into the Mallapuram and Palghat districts of

Kerala State, while portions of the southern

slopes fall into Coimbatore District of Tamil

Nadu.

The northern borders of the area covered

are clearly defined by the Moyar river, and

most of the southern border is defined by the

Bhavani river. Elsewhere the limit is simply

taken as the foot of the mountains and the

immediate surrounding plains.

Climate

Situated at 11° north the Nilgiris are well

into the tropical zone with the result that tem-

perature variations during the year are rela-

tively modest. Mean monthly averages between

the coldest month (December or January) are

normally no more than 5°C below that of the

warmest summer month (usually May). How-

ever, the altitudinal temperature differences

are highly significant. The annual mean tem-

perature is 28° at the foot of the southern

slopes and only about 15° in Ooty at 2200 m.

These altitudinal temperature differences have

profound ecological effects. Average monthly

temperatures for selected localities are given

in table 1 below.

Rainfall patterns are dominated by the mon-

soon regime that affects all of India, but diffe-

rent parts of the Nilgiris are affected in

different ways. The SW monsoon normally

commences in the latter part of May or in

early June, continuing till some time in Sept-

ember with the occasional pauses. This is

India’s life sustaining main monsoon, but the

positioning and topography of the Nilgiris are

such that the main effects of the SW monsoon

are felt on the western and northwestern

slopes. The southern and northeastern slopes

are partly in a rain-shadow and receive as

much of their total rainfall from the retreating

NE monsoon as they do from the main mon-

soon. The plateau receives rain in an inter-

mittent fashion from both the monsoons and

from more localised thunder showers at other

times of the year.

The first four months of the year (January

to April, and much of May) are everywhere

rather dry, though the southern slopes get

some rain even then. Interestingly this dry

season is most marked on those parts of the

western slopes that otherwise receive more

precipitation than any other areas of the

Nilgiris. But for the very pronounced drought

27

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

Table 1

Source: von Lengerke (1977)

the wetter parts of the Nilgiris would doubt-

less be even more rich in flora and fauna than

they already are. I suspect that the exceptional

richness of the flora and fauna at Kallar, a

locality with a relatively modest 1500 mm of

rain a year, is due to the fact that it is ex-

ceptionally well distributed in time, avoiding

periods of pronounced drought. Some illustra-

tive examples of rainfall patterns are given in

table 2 below.

The volume of rainfall varies by a factor of

more than twelve in the Nilgiris. Both low-

land and highland areas of the western slopes

that are fully exposed to the SW monsoon

receive more than 6000 mm a year, and more

than 3500 mm is normal in this area. The

driest parts of the Moyar Gorge, an area in

permanent rain shadow, receive less than 500

mm. Some examples are given in table 3 be-

low, compressing data from table 2 above.

Ecology

Rainfall patterns, temperatures and to a

lesser extent topography combine to produce

in the small area covered by the Nilgiris a

network of very complex ecological conditions

with a degree of variation that is not usually

found in so small an area. Given the statistics

this is not surprising: Elevation ranges from

100 m to well over 2500 m; rainfall from less

than 500 mm a year to more than 6000 mm,

perhaps in some cases much more; the lowest

temperature ever recorded at the foot of

Nadgani Ghat is about 18°, while frost is a

regular feature at Ooty and elsewhere on the

plateau.

While broad ecological zones of considera-

ble diversity can be identified the net result

is not always a very tidy pattern because of

topographical quirks, rain-shadows and local

peculiarities. To give one example. The

28

Annual rainfall in selected Nilgiri localities (by month and year total in

BUTTERFLIES OF THE NILGIRI MOUNTAINS

29

Source: von Lengerke (1977).

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vot. 84

Nadgani site where many of the species re-

corded in the systematic part were observed

probably had a rainfall of about 6000 mm a

year, but just two kilometres further west,

where I usually camped, rainfall was down to

3000 mm. On many occasions I was repeated-

ly drenched during a day’s collecting only to

come back and find that there had been no

rain at the camp site.

Table 3

Total rainfall in four selected Nilgiri localities

AND ITS DISTRIBUTION BETWEEN SW MONSOON

MONTHS, NE MONSOON MONTHS AND THE DRY

SEASON IN PERCENT

Source: Calculated from table 2.

Such differences make the development of

a broad ecological framework difficult, though

such a framework will be essential for a sub-

sequent interpretation of the total Nilgiri

butterfly fauna in ecological biogeographical

terms. I have however, attempted to make

such a framework based on the work of pre-

vious authors, leaning heavily on a simplified

version of the forest classification by Cham-

pion & Seth (1968), as well as on my own

observations and conclusions.

Ecological framework for the Nilgiris

For the purposes of analysing the butterfly

fauna I have arrived at a classification frame-

work covering twelve main ecological zones.

The first, and rather crucial division is one

based on altitude, the attendant climatic diffe-

rences, and their joint effect on the floristic

and faunistic composition. On this basis the

Nilgiris may be divided into three major divi-

sions as follows:

tropical/lowlands 100-1500 m

subtropical/middle levels 1500-1900 m

MONTANE /plateau 1900 m+

The exact altitudes of change from one divi-

sion to the next is subject to local variation

and there is often a transition zone. Where

the subtropical division merges directly with

evergreen tropical forest it may be difficult to

define when the transition between the two

has actually been effected.

The following paragraphs briefly characterise

the salient features of each of the eleven zones,

which are then again summarised in table

4.

TROPICAL DIVISION

1. Rainforest zone. Where rainfall exceeds

3500 mm a year in the lower part of the

tropical division, and if topography does not

militate against it, the climax vegetation be-

comes a fully developed rainforest system

approaching that called giant evergreen by

Champion & Seth (1968). There is a closed

canopy and several storeys with an amazing

faunal and floral diversity. Most of the Nadgani

Ghat and some areas near Mukkali at the base

of the Silent Valley access road belong to

this zone. Were it not for the fact that the

dry period from January to April was very

severe the number of species would be even

greater, but some species without diapause

mechanisms cannot survive. A significant num-

ber of the most interesting South Indian

butterflies are effectively limited to this zone.

2. Wet Evergreen zone. This zone is charac-

terised by a lower rainfall than the previous

30

BUTTERFLIES OF THE NILGIRI MOUNTAINS

one, ranging from 1800 mm a year to 4000 mm

at the higher levels of the tropical division.

Visually it is less grandiose, the trees lower,

and the typical storied structure of true rain-

forest has become a confused tangle. It covers

the West Coast Tropical Evergreen of Cham-

pion & Seth (1968), and in the zone I also

include the patches of evergreen found on the

southern slopes which they would classify as

Semi-Evergreen. From a butterfly point of

view it is characterised by a great degree of

species diversity, the absence, or near absence,

of the true rainforest elements, and the fact

that species from the more open formations

still do not penetrate. Much of the forests of

the western slopes fall into this category. I

have not worked any spot in this zone con-

sistently, but Kallar forms a transition between

it and the mixed deciduous forest (see 4

below).

3. Wet Agricultural zone. Till a few cen-

turies ago the entire western slopes and good

parts of the southern slopes were covered with

evergreen forests of the two types discussed

above. Both are very fragile eco-systen^s that

do not take kindly to human interference and

where wood has been exploited virtual eco-

logical deserts result, clad only in grasses and

one or two dominant dicotyledons. Such areas

are entomologically useless, but unfortunately

they prevail over long stretches. Much of the

area has been under agriculture for the past

few centuries, with rice, bananas, turmeric,

rubber, palm crops, pistachio and various

fruits as the main crops. Plantation teak is the

worst of all. Any butterflies that remain are

found in fringing riverine forest and untidy

gardens, but they are not many. A very few

skippers thrive on the rice and some common

butterflies of the more open formations manage

to establish populations, but on the whole wet

agricultural lands are disappointing for butter-

flies, except that the occasional forest species

maintains a surprising foot-hold here and

there.

4. Mixed Deciduous Forest zone. Much of

the lowland southern slopes and the central

parts of the northern slopes consist of dense

mixed deciduous forest with considerable floral

and faunal diversity and a wide range of

micro-habitats in response to climatic and

topographical variation. Wherever permanent

moisture is available there is an admixture of

evergreen plants, occasionally leading to local

almost evergreen patches. The rainfall regime

is usually in the range of 1000 to 1800 mm.

The physical aspect is a low, but very dense

forest, at times almost impenetrable. At the

upper levels it merges gradually with the sub-

tropical evergreen forest on the southern

slopes. The mixed deciduous forest is very

different from the moist-deciduous forests of

the Wynaad and the Mudumalai Wildlife

Sanctuary, which are very open with tall teak

as the dominant tree. Moist-deciduous forest is

a relatively poor butterfly habitat which is

weakly represented in the Nilgiris proper and

not dealt with separately in this paper, though

notes are given at relevant points in the syste-

matic part. The whole of the Kotagiri Ghat

below Kunjapannai is typical of the mixed

deciduous forest zone.

5. Thorn Forest zone. Once rainfall drops

to between 500 and 1000 mm the natural

climax vegetation becomes some form of

savanna-like forest which visually is not dis-

similar to parts of East Africa and where, in

fact, much of the dominant vegetation has

African affinities, not least species of Acacia.

This is also the home of one of the Indian

antelopes, the Blackbuck. The floral and faunal

composition is much less diverse and varied

than the mixed deciduous forest. Most of the

area between Masinagudi and Bhavanisagar

via the Moyar Gorge forms part of this zone,

but it is also represented at many places in

31

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

the foothills of the southern slopes, when rain-

fall is not high enough to support mixed

deciduous forest. It is also the home of the

most typically African of the South Indian

butterflies.

6. Dry Agricultural zone. Most of the land

at the foot of the forested slopes now consists

of non-irrigated farmlands, much of which

would have been thorn forest, and in favoured

localities mixed deciduous forest. Often rocky

outcrops and natural fences are left untended.

Such areas contain many of the species charac-

teristic of thorn forest, but by and large it is

a very poor butterfly habitat. Here and there

irrigated agriculture prevails. Except where

trees and palms are grown, such areas mainly

contain the most common plains species and

those that have adapted to man-made environ-

ments. At Kallar, with its extensive planta-

tions, some forest species, even those of the

evergreen forests, survive. Most of these would

probably be absent if Kallar were not imme-

diately adjacent to undisturbed forest.

SUBTROPICAL DIVISION

7. Subtropical Evergreen Forest zone. The

exact altitudinal boundaries of the subtropical

zone vary from 1200/1500 m to 1800/1900 m

depending on local circumstances, and once

these elevations are reached the extreme diffe-

rences in annual precipitation that create such

diversity in the tropical zone have largely

disappeared. Consequently the range of vege-

tation zones has also narrowed, and the pre-

dominant climax vegetation of the western and

the southern slopes is subtropical evergreen

forest, except where the scarps are so steep

that forest cannot cling to them. Where the

subtropical forests abut tropical evergreen

forests (e.g. at Silent Valley) the transition

between the two is both visually and floristi-

cally very gradual. Elsewhere the transition

between mixed deciduous and subtropical ever-

green is abrupt and very noticeable, such as

on the Kotagiri Ghat. Climax subtropical ever-

green forest is often almost impenetrable. As

far as the butterflies are concerned this zone

has a small, but conspicuous, special element.

Most of my own experience of this zone is

from the forests at Glenburn and Kunjapannai.

8. Sub-climax Subtropical Forest zone. In

areas of low rainfall on the northern slopes

and in disturbed and rocky areas of the south-

ern slopes the subtropical evergreen forest

cannot develop fully. The result is some im-

poverished associations drawing from the

evergreen and deciduous zones the most hardy

species, interspersed with grasslands. These are

very poor habitats with no major influence on

the Nilgiri fauna.

9. Subtropical Agricultural zone. Most of

the subtropical zone has been converted to

plantation agriculture, with coffee at the lower

reaches and tea at the upper. On the whole

these are poor butterfly habitats, not least

because weedicides and pesticides are applied

liberally, frequently and consistently. Butter-

flies are only found along streams which, at

least in the coffee country, often has a belt of

fringing, riverine vegetation. Tea country is a

virtual ecological desert, only a handful of

butterflies managing to survive. The grassland

areas are indicators of recent forest destruction

and are not natural habitats for any species.

MONTANE DIVISION

10. Montane Evergreen Temperate Forest

zone. The plateau of the Nilgiris, mainly

above 1900 m, has two characteristic types of

natural vegetation. The evergreen forests

known as sholas, often growing only in shel-

tered places, and rolling open grasslands with

only a few Rhododendron trees. The sholas are

dark, almost impenetrable forests, their edges

often shaped by the prevailing winds. Typical

sholas develop under rainfall regimes from as

32

BUTTERFLIES OF THE NILGIRI MOUNTAINS

low as 1200 mm to as high as 6000 mm+

without displaying much difference in floral

composition or physical aspect. The reason for

this is that the sholcts in the most heavy rain-

fall area have a rapid run-off of excess water.

Patches of shola remain over most of the

plateau, and extensive forests may still be

found in the Kundahs, at Avalanche and in

the Mukurti Peak area. One of the prettiest

and most accessible is the Longwood Shola

near Kotagiri, which has a most attractive

bog at its centre. The floral composition is

characterised by a large proportion of plants

with Palaearctic and Oriental montane affi-

nities.

11. Montane Grasslands zone. In between

the sholas the normal vegetation consists of

rolling grasslands, but this is now in evidence

only in the remoter parts towards the western

escarpment between the Sispara Pass and

Mukurti Peak. Elsewhere they have been con-

verted to tea. These grasslands are the home

of the famed Nilgiri Tahr. Champion & Seth

(1968) are insistent that the grasslands are

derived and that the entire plateau was once

clad in forest. The presence everywhere of

the fire resistant Rhododendron is their main

evidence. Ranganathan (1938) takes issue

with this (referring to the first edition of

Champion), pointing out that wind and frost

would preclude shola vegetation in some parts

of the plateau. Blaseo (1971) would appear

to support the latter viewpoint. I do not have

the expertise to settle the matter, but certainly

the vegetation of the grasslands is both more

complex and more varied than in the recent

grasslands of Sumatra and Papua New Guinea

which are definitely the result of human inter-

vention. Grasslands were extensively present

when the first explorers visited the Nilgiris just

150 years ago. It is difficult to see how a

tiny handful of Toda pastoralists could, or for

that matter should, have indulged in massive

deforestation.

12. Montane Agricultural zone. Whether or

not the montane grasslands are derived, it is

a fact that more than three quarters of the

plateau has been heavily modified by human

intervention. Beginning in 1832 large areas

have been forested with exotic trees, espe-

cially of Australian origin ( Acacia dealbata,

A . decurrens, A. melanoxylon and E. globu-

lus). One of the latter is now at 78 m one of

the tallest trees in all of India. Somewhat later

a number of temperate conifers were also

planted. Virtually any tree from anywhere

that was of potential promise has been tried

out in the Nilgiris and remnants are some-

times found in surprising places. The amount

of tree planting may well have exceeded the

deforestation in some areas, but diversity was

sacrificed for monoculture. In the Ooty area

some homesick English gentlemen spent their

summers strewing about the seeds of broom

and gorse on the downs, with the result that

these plants are locally dominant. Since their

arrival some 400 years ago the local Badaga

have concentrated on the cultivation of vege-

tables with potatoes, carrots and cabbages as

the main crops, though wild boar is a pere-

nnial problem. However, in terms of both area

and visual impression it is tea, tea and yet

more tea that predominates as a never-ending

ecological desert.

Summary

This concludes the brief overview of the

Nilgiri ecology and outlines the twelve ecolo-

gical zone which will later be used for an

analysis of the ecology and zoogeography of

the butterfly fauna. Table 4 below gives a

brief summary of the information. As

already mentioned a somewhat robust approach

is called for when dealing with major ecolo-

gical zones in a area where so much varia-

tion can be found within a few kilometres.

33

3

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol 84

Only a robust approach can impose some

element of order into what sometimes

appears to be anarchy; it should not be used

to obscure the fact that genuine anomalies

exist and that surprises do occur. It is also

necessary to highlight once again the fact that

much of the area, especially that above 1200

m, has been the subject of very intensive

human intervention for the past 150 years. It

its good to know, though, that representative

portions have been maintained for posterity.

The proposed inclusion of the Nilgiris in the

World Biosphere Reserves scheme will hope-

fully ensure that the environment is gradually

improved. Few places contain as much diver-

sity within so little space as the Nilgiris. As

an ecosystem it demands our respect.

Table 4

Summary of the twelve major ecological zones

of the Nilgiris

Tropical Division (100-1500 m)

1 . rainforest zone

Rainfall 3500 mm+; temperature range 22-30°.

Tall closed canopy forest with little undergrowth

except where bamboo prevails. Hopea, Dipterocar-

pus, many Guttiferae, many Anacardiaceae, Sapota-

ceae, Meliaceae, etc. Great floral and faunal diver-

sity. Strong affinities to Sundaland. Only western

slopes and not usually above 1000 m.

2. wet evergreen zone

Rainfall 1800-3500 m; temperature range 18-28°.

Dense closed forest, lower than previous zone, and

often with well-developed undergrowth. Much local

variation and transition to moist-deciduous or to

semi-evergreen depending on specific circumstances.

Most of western slopes and part of southern slopes.

3. wet agricultural zone

Rainfall 2000-5000 mm; temperature range 18-30°.

Main crops rice, coconut, coffee, cardamom, fruits

and plantation trees. Mainly at the foot of the

western slopes. Some of the slopes are grasslands of

little diversity slowly regenerating to forest.

4. MIXED DECIDUOUS FOREST ZONE

Rainfall 1000-1800 mm; temperature range 20-30°.

Low dense forest with thick undergrowth though

patches of teak and figs may be taller. Typical trees

Anogeissus, Boswelfia, Tamarindus, Santalum,

Moringa. Some penetration of African derived flora.

Mainly southern slopes and parts of Mudumalai, but

also in western slopes rain-shadows.

5. thorn forest zone

Rainfall 500-1000 mm; temperature range 22-33°.

Open savanna forest with Acacia, Zizyphus, Euphor-

bia and other African elements as dominants and

much admixture of Afrotropical flora. The eastern

half of the northern slopes, the eastern third of the

southern slopes, and locally on the southern foot-

hills where not disturbed by agriculture.

6. dry agricultural zone

Rainfall 500-2000 mm; temperature 20-35°. Crops

are mainly rain-fed millets and certain pulses. Many

tropical weeds and some remnants of thorn forest

vegetation. Very dry during dry season, prone to

drought. Much of the plains adjacent to the southern

and northern Nilgiris where forest has been lost.

Subtropical Division (1300-1900 m)

7. SUBTROPICAL EVERGREEN FOREST ZONE

Rainfall 1300-4000 mm; temperature range 15-25°.

In some respects transitional between tropical and

montane evergreen but with some special elements.

Olea dioica is typical. Dense dark forests, now much

reduced in extent. Most of western and southern

slopes where not cut down, and parts of the NW

slopes above the Wynaad and Ouchterlony.

8. SUBCLIMAX FOREST FORMATIONS ZONE

Rainfall 1000-1600 mm; temperature range 18-30°.

Impoverished version of (7) above where rainfall

and soil is deficient. Often clumps of isolated trees

in grasslands. Here and there on northern and

southern slopes but of marginal importance.

9. SUBTROPICAL AGRICULTURAL ZONE

Rainfall 1100-4000 mm; temperatures 15-30°. Coffee

at lower levels, tea at upper reaches, with vestiges

of forest along rivers and where too steep for plan-

tation crops.

Montane Division (1800 m++)

10. MONTANE EVERGREEN FOREST ZONE

Rainfall 1400-6000 mm; temperature range 10-20°.

Dense evergreen forest with Oriental montane and

many Palaearctic plants. Among trees members of

genera such as Ternstroemia, Eugenia, Micheiia,

34

BUTTERFLIES OF THE NILG1R1 MOUNTAINS

Gordonia, Rhododendron, etc. Many low montane

plants ( Begonia , Lobelia, Impatiens ) and Palaearctic

plants ( Fragaria , Viola, Rubus). Dotted throughout

the plateau, but much reduced in extent.

11. MONTANE GRASSLANDS ZONE

Rainfall 1300-3500 mm; temperature range 10-20°.

Open rolling grasslands with a somewhat complex

structure, much variation in structure according to

soil, exposure, drainage and rainfall. Only fire resis-

tant tree is Rhododendron but exotic gorse and

broom prevalent in many areas. Now mainly from

western escarpment system to Mukurti.

12. MONTANE AGRICULTURAL ZONE

Rainfall 1300-3500 mm; temperature range 10-20°.

Three types: monoculture of exotic Acacia, Euca-

lyptus etc; monoculture of tea; and vegetable gar-

dening (carrot, potato, cabbage and others) with

some fallow land. Presently covers most of the

plateau.

History of exploration

General

The Nilgiri Mountains long lay as an iso-

lated and unknown ‘jewel of nature’ protected

by a wide band of malarious jungles, guarded

by tigers and fierce elephants, largely unknown

to the outside world. On the plateau lived

the pastoral Todas exploiting the extensive

grasslands, with primitive hunters and

gatherers in the surrounding forests, but never

exceeding the 10.000 mark as far as popu-

lation was concerned. Their impact on the

environment must have been marginal, but

they developed a culture and a matrilineal

kinship system that has been the delight of

anthropologists. More must have been written

per capita of the Todas than of any other

people. The Kotas provided supplementary

skills in the type of symbiotic relationship that

pastoralists often develop with other groups.

Lower down the mountain lived primitive

hunter-gatherer societies, Irula, Kurumba,

Panniya and related tribes. Their numbers

were, in total, somewhat larger than that of

the Todas, but they lived in harmony with

nature and had neither the skill nor the in-

clination to encroach in any major way on

the natural conditions.

Some four hundred years ago the Badaga

appeared on the scene. They were Kannada

speaking people fleeing some political up-

heaval in what is now Karnataka, possibly

the repercussions of the Bijapur Muslim con-

quest of the Vijayanagar empire. They were

the first agriculturalists on the plateau and

settled in the villages that still carry their

original names, digging their fields, but not

maintaining much contact with their area of

origin. Though probably more in number than

the resident Todas, it seems that a modus

vivendi was arrived at so that frictions bet-

ween the radically different life styles of the

two groups were minimised. Life probably

went on much as it had always done, except

for the compact Badaga villages and their

agriculture. We do not know, it has to be

said, because information on the Nilgiris was

almost nil and no written tradition exists.

Just around 1600 the modem world made

its way to the Nilgiris. Rumours of Christian

communities in the distant mountains, derived

from the visit of the Apostle Thomas in the

early days of Christianity, reached the court

of the Bishop of Calicut. He sent off a group

of Portuguese priests to investigate matters,

but his judgment in choice of emissaries was

not too sound. They came back with a report

that was not ‘so sure and complete as was

desirable’, a rather nice turn of phrase. So a

few years later, in 1603, heavier guns were

brought to bear. The Jesuit Father, Jacomo

Ferreira, led an expedition on behalf of the

Bishop of Malabar in Calicut. He brought

back a fair amount of circumstantial and anec-

dotal information, but of one thing he was

certain .... there were no Christians in the

Blue Mountains, and so the Portuguese lost

interest.

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JOURNAL , BOMBAY NATURAL HIST. SOCIETY, Vol. 84

In 1799, after the defeat of Tipu Sultan at

the hands of the East India Company, the

Nilgiris were part of the territory ceded to

the Company through the treaty of Sriranga-

patna, though probably Tipu never held much

actual sway north of the Wynaad. The moun-

tain was visible to the large British garrison

and civil establishment at Coimbatore, but it

was only in 1812 that the first representative

of the colonial power visited the mountain

itself. This was in the form of a somewhat

unimaginative tax collector whose only com-

ment was that the Nilgiris were not worth any

particular efforts on his behalf A more official

and less single-minded party set off during the

winter of 1818-1819. They reported back with

enthusiasm on the climate, vegetation and

wildlife of the plateau, in fact describing a

paradise on earth. They were perhaps over-

enthusiastic, because their reports were flatly

disbelieved at Fort St. George in Madras ....

or at least as flatly as bureaucratic niceties

would allow: It is somewhat difficult to accept

the understandable enthusiasm of a

group of men notwithstanding .... etc.

But truth will not be concealed, and soon

the Collector of Coimbatore became the driv-

ing force in the establishment of the Nilgiris

as a major hill resort. By 1827 there were

seventeen European houses in Ooty and five

in Kotagiri. Ten years later large scale esta-

blishment of plantations was in full swing and

the destiny of the Nilgiris was changed fore-

ver. The population swelled with an influx of

Tamil speaking plains people, needed for the

tea and coffee plantations. The main towns of

Ooty, Coonoor and Kotagiri became trading

centres over and above their recreational

function. By 1860 the foundations of what the

Nilgiris still are to-day were laid.

Entomological exploration

Butterflies from India reached the scientific

community already at the time when Carl von

Linne published his tenth edition of systema

naturae (1758), the starting point of zoolo-

gical nomenclature. This book contains many

Indian species, and many more were describ-

ed by Fabricius and Cramer in 1775. Soon

after the Nilgiris were opened up they were

visited by explorers and naturalists, both pro-

fessional and amateur. The first of these was

the Austrian nobleman and naturalist. Baron

von Hiigel in the late 183Gies. His material

was described by Kollar and the Felders. It

is, however, striking how relatively late some

of the more prominent South Indian endemic

butterflies were actually described as evidenc-

ed by the list below :

Pachliopta pandiyana Moore 1881

Papilio Uomedon Moore 1874

Papilio dravidarum Wood-Mason 1880

Papilio buddha Westwood 1872

Prioneris sita Felder & Felder 1865

Colias nilagiriensis Felder & Felder 1859

Celatoxia albidisca Moore 1884

Parantica nilagiriensis Moore 1877

Idea malabarica Moore 1843

Mycalesis adolphei Guerin-Meneville 1843

Ypthima chenui Guerin-Meneville 1843

Calaenorrhinus ambareesa Moore 1867

Thoressa honor ei de Niceville 1887

Oriens concinna Elwes & Edwards 1897

The first systematic account of the Nilgiri

butterflies dates back almost exactly a century

when Sir George Hampson [1888 (1889)]

made a comprehensive list based on his five

years of residence in the Nilgiri Wynaad as a

coffee planter. He collected mainly in the

northwestern corner, including the Nadgani

Ghat, but only in that part which lay in the

Madras Presidency, and not the Malabar side,

now in Kerala State. His energies were sub-

sequently transferred to moths. On his return

to the United Kingdom he joined the British

36

BUTTERFLIES OF THE N1LGIRI MOUNTAINS

Museum (Natural History) and produced the

magnificent series on moths published as part

of FAUNA OF BRITISH INDIA. His total output

outstrips most other entomologists that have

ever lived, yet despite this, the quality and

accuracy of his work is legendary.

Hampson’s list contained 275 entries and

he expected that ‘no more than about twenty

species would be added to it’. On current

taxonomical view the list actually contains

somewhat less than 260 species. In all just

about forty species have in fact been added

since, but as a first effort it certainly is no

mean achievement. It is a great shame that

none of his copious field notes were included.

They are available in the British Museum

(Natural History) but I have not been in a

position to avail myself of them.

It is hardly surprising that the nomencla-

ture adopted by Hampson is often difficult to

decipher for a contemporary reader. We are

therefore indebted to Yates (1935) for ‘trans-

lating’ Hampson’s list to the language used

by Evans (1932) which is largely intelligiblle

to-day, and for adding quite a few species

from smaller contemporary collections. Many

of Yates’s additions were species from the

lower parts of the Nadgani Ghat. Yates’s list

comes to 282 entries, some of which are now

considered forms or synonyms.

The next list of Nilgiri butterflies is that of

Wynter-Blyth (1944, 1946). At the time he

was headmaster at the school at Ketti below

Coonoor. Most of his collecting was done at

Ketti, and on the Ghat between Coonoor and

the plains, and not least at Kallar, a magni-

ficent place for collecting butterflies to this

very day. He did make a number of visits to

the Nadgani Ghat, but because of wartime

petrol rationing he did not visit the lower

parts on the Kerala side. In his day, of course,

the forests stretched much more east, almost

to Gudalur.

His list came to 290 entries, later supple-

mented with a further twenty or so. Some of

these were again synonyms or forms, and one

or two were erroneous. His paper is a very

solid one with which I have few quarrels.

Wynter-Blyth was later to publish the most

recent guide to Indian Butterflies (1957) and

the influence of his work in the Nilgiris is

clearly visible in his book.

Various taxonomic works have since scoop-

ed up old unpublished material, and I have

collected six or seven species never before

recorded from the Nilgiris, so that the pre-

sent list contains 299 valid species. As shown

at the end of the list of Nilgiri butterflies there

are about a score of South Indian butterflies

that have not yet been recorded from the

Nilgiris. Some of these, probably about a

dozen, will one day be found also there. They

will be an incentive to future collectors, but

more precise ecological data and better etho-

logical observations than my own should be

the priority, and a much more satisfying pros-

pect that the record of a few additional

species.

Current study

I deliberately chose to study the Nilgiri

fauna because it was already relatively well

known, and because I had some opportunities

for comparison stretching back a hundred

years. It was clear from the outset that the

number of new records would be relatively

small, but I was more interested in the gene-

ral ecology of the area. After publishing this

systematic account of the Nilgiri butterflies it

is my plan to convert the data into a more

formal ecological-cum-biogeographical analysis

over the next few years. Also of interest was

an assessment of the extent to which the eco-

logical degradation that has taken place in

the Nilgiris over the past 100 years had in-

fluenced the butterfly fauna. When choosing

37

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

the Nilgiris I was not unmindful of the fact

that most of the southern mountains in India

are basically very similar as far as the butterfly

fauna is concerned, except that if tropical

rainforest is absent, then so are several of the

more interesting butterflies. Had I chosen the

Ghats below Sultan’s Battery to somewhere in

the South Sahyadri I would have had the

pleasure of filling out some — largely predic-

table — distributional blanks, but it would

have been at the cost of data on the ecology

of the butterflies concerned.

I was in the Nilgiris, with occasional visits

to the Annamalais, the Biligiriranga Moun-

tains, and Kanara, from 11. iv to 18.x. 1986.

During this time I spent roughly half my days

in the field visiting all the ecological zones as

often as possible. My main areas of work were

the Longwood Shola at Kotagiri, the forests

around Glenburn, Kotagiri Ghat, Kallar, Nad-

gani Ghat and especially the little river at its

foot, the area around Masinagudi and the

Coonoor Ghat. Many other localities were

visited once or twice.

It is a pity that I was not able to spend a

whole year in the area. The profound dry

season from January till April has a very

definite effect on the butterflies and it would

have been interesting to study this not ieast

in the Nadgani Ghat area. Some species are

mainly on the wing during dry season in the

rainforest zone. However, I do not think this

loss of information has introduced serious

bias into the systematic part that follows.

Acknowledgements

I have rarely been so well and generously

treated as by my friends and colleagues in the

Nilgiris, none of whom I knew before my

arrival at base in Kotagiri. The Asokan family,

Bunty and Rudi Bosen, the Devadars, senior

and junior, Mr. and Mrs. Dharman, Palni and

Uma Palaniappan, and Mr. Radcliffe. From

many others I received much help and kind-

ness.

Gordon Thompson provided hospitality,

good company and many interesting specimens

throughout, and my debts to him are high.

Thanks are also due to Nancy Fee for gene-

ral encouragement and the necessary impetus

to do things that would not otherwise have

been done. Augustine and Dominic in Kota-

giri did all they could to make my stay at ‘The

Nest’ pleasant.

I spent some interesting and very pleasant

days at the Peechi Forest Research Institute

in Kerala, being looked after in the best

possible way by Dr. K. S. S. Nair and Dr. G.

Matthew.

Mr. J. C. Daniel of the Bombay Natural

History Society aided with a number of prac-

tical issues, as has the Royal Danish Embassy

in New Delhi.

Philip Ackery and Dick Vane-Wright, Pam

Gilbert, Rienk de Jong, Niels Peder Kristen-

sen and Brenda Leonard kindly provided a

lifeline to library resources and advice over

long distances.

In Copenhagen Harish Gaonkar kindly

undertook the task of screening the manus-

cript which resulted in helpful advice on a

number of matters and the eradication of

some errors that would otherwise have

survived.

I am indebted, as often before, to the

Danish Carlsberg Foundation whose grants

have assisted in making this and other pro-

jects economically feasible.

I have not bracketed the names of authors

where the genus has been reassigned since the

original description since virtually all Indian

butterflies are no longer in their original genus.

Systematic part

The systematic part that follows gives the

basic data on each of the species that I accept

38

BUTTERFLIES OF THE NILG1RI MOUNTAINS

as having been genuinely reported from the

Nilgiris. The basic sequence is that of Eliot

(1978) except that the Lycaenidae have been

placed before the Nymphalidae in accordance

with modern usage. The subfamilies of the

Nymphalidae were in older literature usually

accorded family rank, but there is increasing

agreement that they should not be.

Every effort has been made to refer to the

butterflies according to the most modern

nomenclature, much of it adopted from Eliot

(1978) and a number of recent revisions. It

has not been thought the place to discuss

nomenclature in any detail, except where ex-

cessive confusion might be engendered, or

where I have deviated from current practice.

Where a name in Wynter-Blyth (1957) differs

in genus or species, the name used by him is

given in brackets below the current one.

Under each species I have endeavoured to

give the status and the distribution in the

Nilgiris, with whatever other information that

might be of special interest. I have also sum-

marised in brief terms the global distribution

of the Nilgiri species. These are not meant to

be definitive, but simply to give an impression

of how the South Indian fauna is related to

its neighbouring regions.

Systematic list of the Nilgiri butterflies

Papilionidae

Papilioninae

troidini

001 . Troides niinos Cramer

( Troides Helena)

The southern birdwing is the largest of the

Nilgiri butterflies, being endemic to southern

India and the Western Ghats. It is replaced

by Troides darsius Gray in Sri Lanka, while

its closest relative in the Himalayas is Troides

aecus Felder & Felder rather than, as often

assumed, Troides helena Linne. Haugum &

Low (1982-1985) discuss these relationships.

The species is not rare and may even be

common on the western slopes during the

monsoon and immediate post-monsoon

months. The main habitat is lowland ever-

green forest, but the species is at home also

in mixed deciduous forest and the subtropical

evergreen. It is a good coloniser of agricultu-

ral lands and is often common in coffee

plantations. Wandering specimens are not in-

frequently met with on the plateau itself

though it cannot breed there. The biggest con-

centration I ever saw was on some land that

had been cleared for coffee a few years ago

at Kokode Estate near Sholarmuttam. Hun-

dreds were feeding from Lantana growing

among the forest trees that had been left as

shade trees. The species is active in the early

morning, coming down from the tree-tops to

feed from Lantana and Mussaenda. By 10.30

they once again ascend and are then very diffi-

cult to catch. Females will often be found

sitting with the wings open with up to three

males assiduously courting them. The adults

do not come to water. In captivity, on cold

mornings, the butterflies will increase their

body temperature by wing-quivering in the

manner of many moths, being able to fly

actively under conditions where other low-

land species are immobilised. The food plants

are Aristolochia indica and in the wet zone

especially Thottea wallichi, a Troides food

plant not mentioned by Haugum & Low

(1982-85). Much concern has been express-

ed over the conservation status of Bird wing

butterflies worldwide, and the European

Economic Community has banned their im-

port and export even for scientific use. The

Southern Birdwing is by no means a threaten-

ed species, and in some cases its numbers

increase through the agricultural activities of

man.

39

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

002. Pachliopta pandiyana Moore

(Tros pandiyana)

The malabar rose is an unusual swallow-

tail which is endemic to the wetter parts of the

Western Ghats system and it is closely related

to the Sri Lankan endemic Pachliopta jophon

Gray. It appears to be strictly limited to the

wettest type of rainforest, where it is some-

what local, but sometimes very numerous

indeed. I have seen hundreds of fresh speci-

mens of both sexes at Lantana on the Nadgani

Ghat. Activity began as early as 06.45 when

the only other butterflies about were skippers,

and it was not unusual to find specimens

flying even in heavy rain. It normally flies

in dense forest but will visit clearings and

roads where flowers grow; at such times the

three South Indian Pachliopta may be seen

side by side though they are normally ecolo-

gically segregated. In the forest the flight of

the species is very slow at the lower level of

the canopy and despite their great difference

of pattern the resemblance to Idea malaharica

is great, though I will not go so far as to

postulate a co-mimicry relationship. A speci-

men escaped from a photo session in Kotagiri

and was immediately snapped up by a Red-

whiskered Bulbul (Pycnonotus jocosus). By

the time the bird had moved out of sight the

wings of the butterfly had been crumpled up,

but the body not yet damaged. I unfortu-

nately did not see whether the butterfly was

actually eaten. It is obviously a protected

species, but the bulbul would not have known

this.

As far as is known the only larval food

plants are Thottea siliquosa\ according to Jason

Weintraub this is characteristic for the group

to which this species belongs.

003. Pachliopta aristolochiae aristolochiae

Fabricius

( T ros aristolochiae )

The common rose is often referred to as

being very common, but this is not normally

the case in my experience, though it is wide-

spread and obvious. Normally numbers are

not all that large, though I did see a huge

migration at Kotagiri in autumn of 1957 in-

cluding millions of this species and of Pach-

liopta hector in roughly equal proportion. It

is at home in most types of habitat with the

exception of dense, wet forest, but it occurs

on the plateau only as a straggler. Contrary

to the two other members of the genus the

Common Rose will visit water, usually early

in the morning well before the Pierids and the

Papilio begin. Large quantities of water are

ingested and immediately excreted through

the anus, so obviously salts or nitrogenous

substances are extracted metabolically. The

flight is slow and deliberate. Flowers are fre-

quently visited and the species often spends

the night in communal roosts with P. hector.

The species is found practically throughout

the Oriental region, being replaced by P.

polydorus Linne in New Guinea. One of the

female forms of Papilio polytes is an excellent

mimic of this butterfly, and in nature it is

often necessary to look twice before being

certain which of the two species is involved.

The female of the Zygaenid moth Histia

nilgira Moore is a fine example of co-mimcry.

I saw it only once, at Glenburn, and it took

some considerable time before I realised that

it was not a small P. aristolochiae. The moth

is certainly aposematic and exuded a foul

smelling yellow substance from the tegulae

when handled. At rest it adopted the normal

Zygaenid posture and all resemblance to a

swallowtail was lost.

004. Pachliopta hector Linne

( Tros hector )

The large and beautiful crimson rose is

endemic to Sri Lanka and South India, being

found mainly south of the Godavery river and

in West Bengal. There are records also from

40

BUTTERFLIES OF THE NILGIRl MOUNTAINS

eastern Burma and the Andaman Islands. It

is sometimes very common indeed, may

migrate by the million, and congregates in

small forests and fruit tree groves in commu-

nal roosts during winter. One such roost at

Mahabalipuram in December 1985 must have

contained many tens of thousands and possi-

bly more than 100,000. I have never seen as

many large butterflies in one place [though of

course the wintering sites of the American

Monarch Danaus plexippus (Linne) would

vastly surpass it in numbers]. P. hector is

very fond of flowers, but never comes to

water. The main habitat is open dry decidu-

ous forest and ill-kept agricultural land at low

altitudes, though migrants and vagrants will

be found to the highest peaks. I once saw the

species literally fall out of the sky at Kallar.

On closer investigation it turned out that a

supernumerary male had attached itself to a

copulating couple, clinging so tenaciously to it

that the threesome could be lifted off the

ground by holding a wing of any of the speci-

mens. The species would repay systematic

investigation into diapause and migration.

PAPILIONINI

005. Chilasa dytia clytia Linne

The common mime, in both sexes, occurs

in two forms. The typical form, clytia, is an

excellent mimic of species of Euploea, while

form dissirnilis is an equally good mimic of

the two Tirumala and of Parantica aglea. The

slow and deliberate flight of the Danaid

models is beautifully copied, but try to miss

a specimen with the net — off it goes at a

speed which would give even a Charaxid a

good run for its money. In nature the mimicry,

as it often the case, is much more convinc-

ing than in cabinet specimens, and I have

rarely been quite certain whether I actually

had a Mime in front of me before taking it

out of the net. Working on Sri Lankan mate-

rial Clarke & Karunaratne (1967) found that

/. clytia was the ancestral form, and as in

Sri Lanka my own Nilgiri observations indi-

cate that there are two specimens of dissi-

milis to every specimen of the ancestral form.

As in Papilio polytes the relative proportion

of the two forms may vary from place to

place. Near Karkala in South Kanara Gordon

Thompson and I found /. clytia to be more

common than dissirnilis. In the Nilgiris the

butterfly is quite rare and at best two or three

are seen on any one day. Their main habitat

seems to be the mixed dry-deciduous forests

at low levels and the lowland evergreen forest.

It penetrates the lower level of the subtropical

evergreen zone and at Kallar may be found

in the Arecanut plantations, probably because

the larval food plant Cinnamonum is grown

there. The adult butterfly will come to bolh

flowers and to water where its habit of hover-

ing above the flower or puddle will tell it

apart from the models. The larva is very con-

spicuous (see frontispiece in Woodhouse)

while the twig-like pupa is one of the natural

masterpieces of camouflage. The species is

found throughout the Oriental region, each

subspecies carefully tuned to mimetic resem-

blance of the local Danaids. It would doubt-

less be a fascinating laboratory insect for

further genetic research.

006. Papilio demoleus Linne

The lime butterfly is common and may

be found anywhere in the Nilgiris, though it

does not spontaneously enter the densest and

the wettest of the lowland evergreen forests.

However, numbers fluctuate considerably in

time and space in a somewhat unpredictable

way. It is a pest on cultivated citrus, but it

also feeds readily on any type of wild ruta-

ceous plant. I have not found it on cultivated

fennel and it is perhaps surprising that none

of the South Indian swallowtails has managed

to transfer to this plant. In the immediate

pre-monsoon period P. demoleus is an avid

41

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

participant in mudpuddling assemblies, as

many as one hundred being found in tight

groups, shoulder to shoulder. The distribution

of this Oriental species is somewhat puzzling.

The core area includes India to Malaya and

southern China, with a secondary area in the

Australian region. During the past seven

hundred years it has colonised Arabia (Lar-

sen 1983) where it feeds exclusively on culti-

vated Citrus and where it is almost in contact

with the African vicariant P. demodocus Esper.

It invaded southern Iraq from Iran only since

1957 (Larsen 1977a). In recent years it has

established populations also in Sumatra and

on the Philippines, almost certainly in res-

ponse to human interference with the natural

environment. It is greatly to be hoped that

the recent invasions will be well chronicled.

007. Papilio liomedon Moore

The MALABAR BANDED SWALLOWTAIL is a close

relative of the Oriental Papilio demolion

Cramer, but is definitely specifically distinct

and endemic to the Western Ghats, chiefly in

the wettest parts of the lowland evergreen

forest zone. It is a rather scarce butterfly

which is also somewhat localised. The few

Nilgiri records are all from the Nadgani Ghat,

and here, too, I saw my own only Nilgiri

specimen on flowers (19.0) just on the TN/

Kerala border. In behaviour it is much like

the other swallowtails, though according to

Gordon Thompson it does not come to water.

It is certainly much more common in Kanara

than in the Nilgiris, and in late September

to early October good series may be collected

on the globular red flower heads of Clero-

dendron paniculatum in suitable localities.

008. Papilio dravidaram Wood-Mason

The malabar raven is another endemic of

the Western Ghats system, its closest relative

being a species from Assam and Thailand.

It is rather more common than P. liomedon,

but is not frequently met with. The habitat is

chiefly the wettest of the lowland evergreen

forests and in the Nilgiris the main locality

is Nadgani Ghat, though Gordon Thompson

has occasionally taken it at Kallar. The colour

pattern is reminiscent of the Euploea but the

flight is very rapid and whether a genuine

mimetic relationship is involved is perhaps

doubtful. The butterfly has the disconcerting

habit of suddenly emerging at furious speed

from dense jungle through a road or clearing,

only to vanish immediately again. It may be

hard to distinguish from P. polytes on the

wing. Wynter-Blyth (1957) states that it never

seems to visit flowers, but that is not the case.

I have caught several of my few specimens

on Lantana. The species is regularly found at

mud patches during the drier months and on

hot days.

009. Papilio helenus daksha Moore

The red Helen is the third largest butterfly

of the Nilgiris and is common in the subtro-

pical evergreen forests, relatively common on

the plateau, and less common at lower ele-

vations. It is a good coloniser and exists well

outside of natural forest, and it is one of the

few large butterflies to be found in tea planta-

tions. It is, however, unable to survive in the

drier thorn forests. It is very fond of Lantana

blossoms and will sometimes assemble on

mud patches, usually in the company of other

swallowtails, but not usually with the large

agglomerations of Pieridae. The weaving,

random flight looks most haphazard, but in

fact the butterfly makes rapid progress and is

very difficult to catch. It seems to feed on

virtually all available rutaceous plants and is

easily bred. The global distribution covers

almost the entire Oriental region as well as

the southern fringes of the Palaearctic in China

and Japan. Colour pattern apart, the simi-

larity between this species and P. paris tami-

lana is strong, and it is interesting that the

42

BUTTERFLIES OF THE NILGIR1 MOUNTAINS

South Indian subspecies of both should be

the largest of them all.

010. Papilio polytes polytes Linne

The common mormon is indeed a common

butterfly at lower and middle heights, but it

does not ordinarily breed on the plateau,

though from time to time it is seen even on

the highest peaks. Like others of its kind it

does not really penetrate virgin evergreen

forest, though it colonises whenever it is dis-

turbed by human activity. It comes to flowers

readily and sometimes large numbers of males

are seen mudpuddling. Such assemblages are

usually independent of the Pierid agglomera-

tions, occupying somewhat shady spots, often

in the company of P. crino and other swallow-

tails. The largest assembly that I have per-

sonally witnessed consisted of 34 P. polytes,

5 P. crino, 1 P. demoleus, and 2 Graphium

doson. Two of the three female forms are ex-

cellent mimics of two of the red-bodied

swallowtails, the Common and the Crimson

Rose respectively, while the third female form

is almost like the male. The flight pattern of

the mimetic forms is also a wonderful copy

of the slow and deliberate flight of the models.

During my frequent visits to Kallar I noted

all females seen in nature, though in many

cases it was necessary to check the body

colour to be quite certain whether model or

mimic was involved. At Kallar the two mimetic

forms predominated, the male-like form be-

ing rare and constituting less than 5 percent

of the total female population. The two

mimetic forms were equally common. The

precise data are as follows : hector- mimic

50.4%, aristolochiae mimic 45.7%, and male-

like 3.9%. This is in marked contrast to the

Delhi population of the same species, where

some 60% were mimics of aristolochiae and

about 40% male-like. P. hector does not occur

in Delhi and the mimetic form is very rare,

certainly less than one in a thousand (Larsen

1987). Such differences constitute powerful

support for the concept of balanced poly-

morphism as discussed by Ford (1975). Frjer

(1913) bred the species extensively in Ceylon

to work out the genetics and found that the

male-like form constituted nearly 50% of

the total. These results were puzzling to me

till I realised that he was breeding the species

at an altitude where the models are scarce

indeed, just as in Delhi. The Common Mor-

mon is, by any standards, among the world’s

most interesting butterflies and an excellent

insect for both field and genetic studies. An

indispensable starting point for any research

is the paper by Clarke & Sheppard (1972) on

the genetics of the species.

Oil. Papilio polymnestor Cramer

The blue mormon is the second largest of

the South Indian butterflies, and it is a plea-

sure that such a large and handsome butterfly

should be common practically everywhere.

Only in the driest lowland habitats is it gene-

rally absent, though stray specimens will turn

up even there. I have seen the occasional

specimen at Masinagudi. It is endemic to Sri

Lanka and peninsular India, but is closely

related to the widely distributed Oriental P.

memnon Linne. The latter species has a host

of mimetic female forms, some being quite

similar to Pachliopta pandiyana, and it is per-

haps surprising that the female of the Blue

Mormon is similar to the male. Occasional

females have bright red spots at the base of

the forewing upperside, a feature which enters

the mimetic pattern of some memnon forms.

This red spot is very much more frequent in

Sri Lanka than in the Nilgiris. Possibly the

very narrow range and habitat choice of P.

pandiyana makes it an unsuitable model. The

Blue Mormon is found in all types of habitat,

flying with a rapid weaving flight, covering

long distances. It visits flowers avidly and

comes freely to water. Unlike many swallow-

43

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

tails it comes to foul substances, otter drop-

pings being a great favourite. I once found

three males deep inside a cave where a group

of otters had been dismembering crabs. The

eggs are laid on all the natural rutaceous

plants as well as on cultivated citrus.

012, Papilio paris tamilana Moore

The large and brightly coloured South

Indian subspecies of the paris peacock is the

largest of all, and it qualifies among the finest

butterflies anywhere in the world. It is found in

all types of evergreen forest from near sea

level till at least 2400 m, but I suspect its

main habitat is the subtropical evergreen

forest. It is found locally also in the moist-

deciduous forests of the Wynaad and Mudu-

malai. Wandering specimens may be en-

countered in open country, even in the centre

of Ooty, but this really is exceptional.

Normally it does not stray from dense forest.

The species is not really rare, though never

numerous, but it may be very difficult to

catch. Early in the morning it may be caught

at flowers, and on hot days it visits damp

patches, sometimes settling with the wings

held flat against the substrate, which is some-

what unusual in the family. Unusual, too, is

the fact that females are often found in this

way. At Sholayar in the Annamalai Moun-

tains I found the species hilltopping along a

ridge in primary forest and saw two copulat-

ing pairs. I can subscribe to the view of old

observers that P. paris has fixed patrol routes

in its jungle habitats .... if you miss a speci-

men it is likely to reappear an hour or so

later from the same direction. They are fond

of flowers, and when half a dozen or so are

feeding on the same Lantana patch, the sight

is a highlight of all that is enjoyable in natural

history. The species is found in South India

(but not Sri Lanka), in Orissa, and then from

Kumaon east to most of the Oriental region.

It is curiously absent from Malaysia.

013. Papilio crino Fabricius

The common banded peacock is endemic

to peninsular India and Sri Lanka. I have

seen no phylogenetic reconstruction of the

group, but it would appear that the species is

fairly isolated. In South India it is more or

less restricted to lowland mixed deciduous

forest, with only the slightest of colonising

ability. I have, however, in December 1983

seen it in the centre of Mysore, from where

it was not recorded in lists from early this

century (Watson 1890). The closest South

Indian relative of P. crino is P. buddha which

is essentially limited to the wettest parts of

the evergreen zone with the result that the

two species are hardly ever sympatric. P.

crino is somewhat migratory so it may on

occasion be found far from its natural habitats.

The species is not very common in the

Nilgiris. except at Kallar, where it may abound

about the time of the onset of the monsoon.

A few years ago several thousands were

caught during one season on behalf of Japa-

nese commercial collectors, but the popula-

tion does not seem to have suffered. The

species is very fond of mudpuddling and I

have seen more than a dozen assembled,

though none was a female. Both sexes come

to Lantana, though visits to flowers are infre-

quent at the time when mudpuddling is at its

maximum. The courtship display, with the

male hovering below the female, is very simi-

lar to that of P. polytes. The larval food plant

is known to be Chloroxylon, the Satinwood

tree, but it seems that no-one has found the

larva recently, and I was wholly unsuccessful

in this respect.

014. Papilio buddha Westwood

The buddha peacock is arguably the finest

butterfly in South India. The general pattern

is not unlike that of P. crino and on photo-

graphs they may look alike. In real life the

Buddha is so much brighter and more

44

BUTTERFLIES OF THE NILG1RI MOUNTAINS

beautiful than the other gloss swallowtails, and

it is a sight of rare beauty. The species is

limited to wet lowland evergreen forest but

is not usually found where rainfall is much

less than 4000 mm a year, and it seems unable

to survive in severely disturbed forest. On the

Kanara Ghats, however, where paddy fields

are interspersed with primary forest, the

Buddha Peacock emerges from the forest to

feed on Lantana and on Clerodendron pani-

culatum. The latter plant is also a great

favourite of the Purplebacked Sunbird, which

attacks the Buddha in the belief that the

butterfly is a supernormal rival, since the

green colour of Buddha matches the green

crown and the wing shoulders of the bird

(Larsen 1987c). Normally the butterfly flies

in dense forest at canopy level with a furious

flight and is almost impossible to net except

when visiting flowers. There are no records

from damp patches, though P. crino is an avid

mudpuddler. In the Nilgiris it is limited to ihe

western slopes and though not uncommon is

difficult to collect. It is more common in

Kanara. The chief season is just after the end

of the SW monsoon in late September and

early October, but individuals may be met

with at any time. Bell et ad. have data on

diapause in Kanara. It should be mentioned

that Gordon Thompson once caught a male

at Glenburn, an illustration of the potential

dispersal power of such a strong butterfly, I

have given a more detailed account on the

ecology and habits of this species elsewhere

(Larsen 1987).

LE PTOCIRCI N I

015. Graphmm sarpedon teredos! Felder &

Felder

(Zetides sarpedon)

The common bluebottle is a genuinely

common butterfly in the Nilgiris from the

lowest levels to the highest peaks, and it is

the one species able to colonise agricultural

lands. However, it will not survive perma-

nently in the driest tracts. It is an avid visitor

to both flowers and damp patches, but at

lower levels it is often outnumbered by G.

doson. By disposition it is a very nervous

insect and is not easy to catch. More than

most swallowtails it is attracted to natural

baits such as rotting grasshoppers and cicadas.

I have found the larva on Cinnamonum, but

it feeds on numerous other plants as well.

The world-wide distribution covers practi-

cally all of the Oriental region.

016. GrapMum doson eSeius Fruhstorfer

( Zetides doson )

The common jay is essentially a butterfly

of the evergreen forest zone at both tropical

and subtropical levels, but it is occasionally

found away from forest proper. At low levels

in evergreen forest it is sometimes more nume-

rous than is G. sarpedon. Large numbers may

be seen mudpuddling just before the onset of

the SW monsoon, and when the monsoon

changes in Sep. /Oct. My childhood records

indicate that we did occasionally see it as

high as 2000 m, but this is exceptional. The

world-wide distribution covers Sri Lanka and

southern India, suitable spots of the Eastern

Ghats., and then most of the Oriental region.

017. Graphium agamemnon sneakies Felder &

Felder

( Zetides agamemnon)

The tailed jay with its apple green mark-

ings is among the more attractive Nilgiri but-

terflies, but on the whole it is less common

than the previous two members of the genus.

Its relative scarcity is a bit surprising since

it is common in cities such as Mysore, Banga-

lore and Madras. It is absent from the very

driest tracts and not normally resident on the

plateau, though it may breed there during

summer. While it is an avid visitor to flowers

45

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

it is much less of a compulsive mudpuddler

than the two preceding species. The global

distribution covers the entire Oriental region.

018. Pathysa nomius nomius Esper

The spot swordtail may be very common

during the dry season at the foot of

the Kotagiri Ghat and at Kallar, but

it is rare elsewhere and at any other time.

It seems to be a species of dry mixed deci-

duous forest, sharing this habitat with Papilio

crino. Just as the latter species has hardly

any overlap with Papilio buddha, so F.

nomius hardly overlaps with the rainforest

species F. anti pat hes. Hampson caught only a

few specimens on the northern slopes, and on

the Nadgani Ghat I have only caught one,

though F. antipathes may be common indeed.

Normally the species flies high and fast, but

in spring (March to early June) it is an avid

visitor to damp patches. It is a migrant and

specimens are seen on the plateau from time

to time, but it does not breed there. The global

distribution covers virtually the entire Orien-

tal region. I agree with Eliot (1978) that it is

legitimate to maintain the generic name Pathysa

for the Swordtails, rather than to subsume

them under Graphium.

019. Pathysa antipathes akibiades Fabricius

The five bar swordtail is a dramatic in-

sect that is limited to the wettest rainforests

of southern India and Sri Lanka. Generally it

is considered to be scarce, but in the right

spots it may actually be very common indeed.

The flight is fast and furious, and when on

the wing the butterfly looks more like a fast

Pierid than like a swallowtail. During Febru-

ary/March, the main season, it is an invete-

rate mudpuddler. When disturbed on a mud

patch specimens will often perch on vegeta-

tion close by. It is reputed to visit flowers but

I have never personally witnessed this. The

species is found in Sri Lanka, S. India, then

again from Nepal east to most of the Oriental

region. The South Indian subspecies has

usually been referred to as ssp. naira Moore.

However, Fabricius described ssp. alcibiades

after specimens from ‘Tranquebar’, then a

Danish colony and examination of the Fabri-

cian types in Copenhagen by Harish Gaonkar

clearly show them to be of South Indian origin

(though Tranquebar cannot be the correct

locality).

PlERIDAE

PlERINAE

020. Delias eucharis Drury

The common jezebel is one of the most

striking Indian butterflies and it is generally

not rare, occasionally being locally abundant.

It may be found practically everywhere in the

Nilgiris, and it seems that the absence of

frost and the presence of the larval food plant,

Loranthus are the only ecological require-

ments. The adult butterfly is very fond of

Lantana flowers, but unlike the montane mem-

bers of the genus in Papua New Guinea it is

not attracted to water. The species is endemic

to India and Sri Lanka and unlike other mem-

bers of the genus which are forest dwellers, D.

eucharis is common even in major towns and

cities. This trait is shared with D. hyparete

Linne which replaces it from Burma eastwards.

When handled the species feigns death and

it is almost certainly aposematic. In Prioneris

sita it has a beautiful mimic.

021. Leptosia nina nina Fabricius

The psyche is the only Oriental represen-

tative of an African genus with half a score

or so of very similar species. Eliot (1978)

suggests that the Oriental species might be

conspecific with the African L. alcesta but this

seems unlikely in view of significant differ-

ences in haploid chromosome numbers (n=19

in nina and n=12 in alcesta). The species is

very common all year at Kallar but otherwise

46

BUTTERFLIES OF THE NILGIRl MOUNTAINS

it appears to be remarkably scarce in the

Nilgiris, though I have found it as high as

1900 m near Kotagiri in the 1950ies. It seems

to avoid the wettest of the evergreen forests

and I have not come across it at Nadgani.

The flight is weak and fluttering, probably

the most feeble of any South Indian butterfly.

Flowers are visited, with Tridax as a firm

favourite, while water is only occasionally

attractive. I have once seen an African

member of the genus actually alight on a

pool of water. The species is distributed

throughout the Oriental region.

022. Prioneris sita Felder & Felder

The painted sawtooth is not rare on the

western slopes of the Nilgiris and on occa-

sion it may be very common on the Nadgani

Ghat during the dry season when large num-

bers are to be found on damp patches in the

company of Graphium doson and Pathysa

antipathes. It is distinctly uncommon during

the rainy season and is only very rarely found

outside of the wettest evergreen forest zone.

I have taken a male (23. v) and a female

(4.vi) at Kallar, but this is most exceptional.

Both sexes are excellent mimics of Delias

eucharis , a butterfly with a much wider range.

The male, it is true, sometimes flies much

faster than the model, in the manner of

Hebomoia glaucippe, but at other times, such

as when circling round a damp patch, the

flight is quite like that of the Jezebel model.

The species is endemic to Sri Lanka and the

Western Ghats system.

023. Artogeia canidia canis Evans

( Pieris canidia )

The Indian cabbage white is a Palaearctic

butterfly that is very common on the plateau

above 1800 m, flying in a series of broods

throughout the year. The original habitat

must have been the edges of sholas, but it is

now especially common in agricultural areas

where it is a moderately serious pest of cab-

bages. In northern India the species is migra-

tory and visits the plains in winter (Larsen

1986a), but I have seen no evidence of migra-

tion in the Nilgiris. The Bangalore record that

so puzzled Wynter-Blyth (1957) probably

came as a pupa with agricultural produce.

The butterfly spends most of its time flying

around in search of flowers and will occa-

sionally visit damp patches on hot, dry days.

It is found on the higher South Indian

mountains but is absent from Sri Lanka. In

all probability the total area above 1900 m

on that island was too small to support a

viable population. Outside of South India it

is found from the Himalaya east to Japan,

just penetrating the tropics in suitable loca-

lities.

024. Cepora nerissa phryne Fabricius

( Huphina nerissa)

The common gull is chiefly a butterfly of

the drier lowland habitats, though it may be

found in clearings at middle heights. Mixed

deciduous forest, ill-kept agricultural land, and

thorn forest are the main haunts of this gene-

rally common butterfly. Due to strong migra-

tory tendencies it is also met with on the

plateau from time to time but I cannot agree

with Wynter-Blyth (1946, 1957) that it is

resident much above 1400 m. In spring large

numbers may be seen mudpuddling with other

Pierids. In behaviour it is more or less a tropi-

cal replacement for the Small Cabbage White

( Artogeia rapae) and it is much less of a

quarrelsome species than the two lxias. The

distribution covers practically the entire Orien-

tal region.

025. Cepora nadina remba Moore

{Huphina nadina)

The lesser gull is limited to the wetter

parts of lowland evergreen forest and is never

seen in open country. Hampson noted that

47

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

it was sometimes common, but generally it is

uncommon. Usually only single specimens are

met with. It spends most of its time flying in

the forest where it is almost impossible to

collect, but one or two can usually be found

on a good mudpuddling patch and occasion-

ally on flowers. Its behaviour and general

aspect in nature contrast so strongly with

those of C. nerissa that it is difficult to accept

them as congeneric. Apart from in Sri Lanka

and South India the species is found from

Nepal east to Taiwan and Sumatra.

026. Anaphaeis aurota Fabricius

The caper white is linked to the drier

tropical habitats, but owing to a great migra-

tory potential it may occasionally be found

in numbers practically anywhere. Vast swarms

are sometimes encountered and, like many

migrants, numbers fluctuate considerably in a

most unpredictable way. In the Nilgiris it is

most consistently common in the thorn forests

around Masinagudi. Under good conditions

this butterfly breeds faster and more profuse-

ly than practically any other and it may com-

pletely strip all available food plants for miles

around. The species is Palaeotropical, being

found on the Indian subcontinent, Arabia and

all over Africa. It occasionally invades the

Mediterranean area (Larsen 1986b).

027. Appias indra shiva Swinhoe

The plain puffin is a rather scarce butterfly

that seems to be centred on the subtropical

evergreen forest zone, though it may be found

both below and above this level. There are

few Nilgiri records. I took one at Tamizagham

in Ooty (ii . 1984) and it seems to have a

headquarters of sorts on the Coonoor Ghat

between Wenlock Bridge and Benhope where

Wynter-Blyth caught most of his material and

where J have seen it on several occasions. I

have seen it once only at Kallar (31. viii) and

Gordon Thompson has a few taken at water

on the Nadgani Ghat where I never saw it.

No clear pattern emerges. It comes freely to

flowers. In Sri Lanka the species is considered

even more of a rarity than is the case in

South India. Otherwise it ranges from Nepal

east to most of the Oriental region where it

is often common.

028. Appias libythea libythea Fabricius

The name striped albatross is something

of a misnomer for the South Indian popula-

tion where the males are practically imma-

culate except for some dark apical shading.

The underside of the hindwing is chalky

white lacking the cream overlay of the other-

wise similar A. albina. A certain distinction

between these two species lies in the cell of

the forewings. A. libythea has the end cell vein

forming a 90° angle to the costal edge, while

in other Appias the angle is only 30°. Hamp-

son considered it to be a rare species while

Wynter-Blyth failed to find it. I have taken

it on the Kotagiri Ghat, at Glenburn, Kallar,

Ronningtown and even on Nadgani Ghat. It

appears to be most unpredictable, possibly

because it is migratory, but its headquarters

are probably the mixed deciduous forests,

though it might be without a permanent head-

quarters. It is certainly most erratic in the

Delhi area (Larsen 1987b). The range covers

Sri Lanka and India, east to the Philippines

and Malaysia. Numbers have increased signi-

ficantly in Malaysia this century (Eliot 1978)

probably because the butterfly does well in

secondary vegetation in the wetter tropics.

029. Appias lyncida Inti fascia Moore

The chocolate albatross is easily recog-

nised by the deep yellow colour of the hind-

wing underside and the broad chocolate

borders. These are not obscured by the con-

siderable level of individual and seasonal

variation. I find it a most enigmatic butterfly

in the Nilgiris, difficult to interpret in ecolo-

48

BUTTERFLIES OF THE NILG1RI MOUNTAINS

gical and distributional terms. Hampson re-

cords it simply in the words ‘1000 to 3000 ft’.

Wynter-Blyth took only three at Kallar, com-

menting that it was reputed to be common

on the Kotagiri Ghat, where I never saw it.

On 1 . v and 2 . v I saw single males flying

towards the SW in Kotagiri town in the

manner of normal migrants. I have seen

small numbers on about half of my visits to

Nadgani Ghat, where it sometimes came to

water. Four were collected in mixed decidu-

ous forest at Ronningtown. This is a pattern’

singularly lacking in consistency. The world-

wide distribution covers Sri Lanka, the

Western Ghats, and then from Nepal east to

most of the Oriental region. An apparently

isolated population exists in Orissa.

030. Appias albina darada Felder & Felder

The common albatross is indeed the most

common of the Nilgiri Appias, but like other

members of the genus it is somewhat unpre-

dictable. I would not, however, concur with

Wynter-Blyth’s statement that it is ‘most

abundant everywhere’. I have taken specimens

in most of the localities visited from time to

time, though this species will not be found

in the drier lowland forests where A. libythea

may be met with. The headquarters appears

to be in the drier parts of the lowland ever-

green forest where it merges with the mixed

deciduous. It may be caught at flowers but

more usually at damp patches. In the large

migration that took place in late May and

early June of 1986 (Larsen 1987b) this

species contributed less than 0 . 5% of the

total (some 20.000 individuals). In the migra-

tions of my childhood (Larsen 1978a) there

were millions. It ranges from Sri Lanka and

the Western Ghats, through suitable places in

peninsular India to Sikkim, and from there

east to practically the entire Oriental region,

New Guinea and NE Australia.

031. Appias wardii Moore

The lesser albatross is inappropriately

named since males are usually larger and

more dramatically marked with black than

those of the other species of the genus. The

taxonomy of this and related species is diffi-

cult. There is something to be said for uniting

the large number of often disjunct and distinc-

tive taxa under the ‘ paulina umbrella, but the

South Indian form is so different from the

Sri Lankan that specific status seems the best

solution till a full revision of the species-

group has taken place. In the wet season form

the South Indian taxon is easily identified

since it has the black markings normally

characteristic of female A. albina . The dry

season forms are very similar to male A.

albina, though vestiges of the black apical

markings sometimes remain. Hampson con-

fused the genus so thoroughly that whether

he obtained this species or not is uncertain.

Wynter-Blyth did not mention it in his main

Nilgiri paper. I have caught it on several

occasions at Glenburn, once near Naduvattam,

and on a few occasions at Nadgani. It is

decidedly uncommon. The range is similar to

that of A. albina, but the species is much

less common, confined to dense evergreen

forest, and populations are often disjunct.

032. Colotis amafa amata Fabricius

( Colotis Calais)

The South Indian subspecies of the small

salmon arab has traditionally been known as

ssp. modesta Butler, but the Fabrician type is

from South India and has been checked by

Harish Gaonkar. At best the name modesta

is applicable to the Sri Lankan population,

but that hardly merits a name of its own. From

North India and west through Arabia and the

African Sahel it flies in the very different ssp.

Calais Cramer. The Small Salmon Arab is

chiefly found in the thorn forest formations

and surrounding agricultural land, though I

49

4

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

have seen specimens also in the mixed deci-

duous forest of the Kotagiri Ghat. The South

and Central Indian subspecies is very different

from the North Indian and African one, and

its main food plant seems to be Azima rather

than the more normal Salvador a. There might

be a case for considering the two taxa speci-

fically distinct, and certainly the presence of

the two subspecies in India would appear to

be due to a multiple invasion from Africa. In

the Nilgiris area the species is quite localised

but usually common where found, often in

the company of the other Colotis species. The

distribution covers all of tropical Africa, much

of Arabia, the drier parts of the Indian sub-

continent. and NE Sri Lanka.

033. Colotis etrida Boisduval

The small orange tip is a common butter-

fly in the drier lowland habitats, but it pushes

further up the mountain and further into the

wetter zones than do the other members of

genus. There is a considerable degree of

individual and seasonal variation, and the

species is often wholly absent during the

wettest months of the year. Occasional speci-

mens are met with on the plateau indicating

some capacity for dispersal. The butterfly

comes avidly to flowers, especially Tridax, but

neither this nor other members of the genus

come to water, despite being found in very

dry habitats. The genus is African, but the

species is endemic to Sri Lanka and the

Indian peninsula.

034. Colotis eucharis eucharis Fabricius

The plain orange tip is a fairly local

butterfly, but it is usually not rare where its

food plant, Cadaba indica grows. This is nor-

mally on rocky ground in the thorn forest of

the foothills, or along the hedges growing at

the edge of fields. It is almost invariably

found in the company of Colotis danae with

which it shares the larval food plant. Usually

most of their time is spent flying about stands

of Cadaba in search of partners, but they will

feed from flowers. On exceptionally hot days

the warmest hours are spent in the shade of

dense trees, usually right down amongst the

roots. The species is an intruder from the

Afrotropical region, where it is found through-

out the tropical zone. In India it is limited

to the Deccan and southern India, being

found in NE Sri Lanka as well, but not in

Sind and Saurashtra where the Afrotropical

species usually occur. This is a classical

Sudano-Deccanian distribution pattern.

035. Colotis danae danae Fabricius

In South India crimson tip butterfly is quite

parallel in distribution and habits to Colotis

eucharis but it is rather more common. If only

one of the species is present it is almost invaria-

bly C. danae. The nominate subspecies is found

on Sri Lanka and in peninsular India as far

north as Madhya Pradesh. A different sub-

species inhabits the area from Saurashtra to

Baluchistan. Usually known as ssp. dulcis

Butler, I find it impossible to separate it from

the Arabian ssp. eupompe Klug. Different

subspecies are found throughout dry, tropical

Africa.

036. Madais fausta fulvia Wallengren

( Colotis fausta)

The salmon arab is a pretty and vivacious

butterfly of the drier foothills where it may be,

on occasion, quite common. It penetrates

higher up the mountains and deeper into the

evergreen forests than most of the related

Colotis species. The flight is rapid and danc-

ing and it is often difficult to capture a good

series of specimens. As elsewhere in India, the

larval food plants are Maerua, though the

nominate subspecies from the Middle East

feeds on Capparis as well. The species visits

flowers freely, not least Tridax, but never

comes to damp patches. The subspecies from

50

BUTTERFLIES OF THE N1LGIRI MOUNTAINS

peninsular India differs from the others

through invariably having white females. In

northern India ssp. fulvia has dimorphic

females, while in the nominate subspecies all

females are salmon. The Indian populations

appear to be highly sedentary though the

nominate subspecies in Arabia and the Middle

East is strongly migratory. The genus is

monobasic, very close to Colotis, and the single

species is found in Arabia, the Middle East

and the Indian subcontinent, just penetrating

the East African coastal regions north of

Kenya.

037. Ixias marianne Cramer

The two Indian Ixias are almost identical

in distribution and habits, though in most of

India the white orange tip, /. marianne, is

slightly less common than /. pyrene. They are

both common in the mixed deciduous forest,

in thorn forest and in ill-kept agricultural land

at low levels. There is little penetration into

the subtropical zone, nor into the lowland

evergreen forests. In May and June both are

among the most prominent species in the mud-

puddling assemblages, and both were promi-

nent in the large migration that I observed in

late May and early June (Larsen 1987b).

Normally the butterflies are seen flying about

the open scrubland in search of flowers, mates

or food plants. /. marianne is endemic to

India and Sri Lanka.

038. Ixias pyrene sesia Fabricius

The use of subspecific names for the yellow

orange tip in India has been the subject of

some confusion. I follow Gabriel (1943) in

using the name sesia. There is, in any case,

so much seasonal and individual variation

that the designation of subspecies is difficult.

The habits are like those of the preceding

species and were the ground colour of the two

not different they would be impossible to

distinguish in the field. The range stretches

from Sri Lanka, throughout India east to

Hong Kong and Malaysia.

039. Hebomoia glaucippe australis Butler.

The giant orange tip is one of my favou-

rite butterflies. It is the largest of the Indian

Pierids, and it is a beautiful sight to see it

swooping down the mountains along densely

clad water courses with the wings held

three-fourths open. It is generally common

enough in mixed deciduous and open ever-

green forest at low levels, sometimes being

found also in subtropical evergreen forest. It

is not a good coloniser of agricultural land

and is rarely found in disturbed areas. It

comes to flowers but is then very wary. Often

it hovers on the flower in the manner of the

Papilionidae, the only Pierid to do so. The

species is an avid mudpuddler actively follow-

ing river systems on the look-out for good

spots which may well be situated some dis-

tance from the forest edge. In the afternoon

they may be seen moving up-river again.

When mudpuddling, together with myriads of

other Pierids and some Papilionids, the

camouflage pattern of the underside may be

seen at its best. Never mind how carefully

you study the assemblage, one or two H.

glaucippe will be overlooked. The larva is a

startlingly effective snake mimic. When handl-

ed it rears up its head, revealing a couple of

blue eye-spots that are concealed when the

larva is at rest. Most likely this is a defence

against the Bonnet Macaque Monkey whose

instinctive fear of snakes is almost comical

in its manifestations (try throwing a little bit

of rope out of a car and watch the results).

The species is distributed in suitable country

throughout the Oriental region.

040. Pareronia Valeria hippia Fabricius

( Parenonia ( sic! ) Valeria)

The common wanderer appears to be very

rare in the Nilgiris. Hampson mentions it with

51

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

no detail, Wynter-Blyth and I failed to find

it, though there is a genuine specimen from

Kallar in the Bombay Natural History So-

ciety collections. This scarcity is difficult to

understand as it is common enough in forest-

ed country around Bangalore and as the forests

on the northern slopes seem very suitable for

the species. It is worth mentioning that the

South Indian population throws up the occa-

sional female of the form mimicking Parantica

aspasia Fabricius, a Danaid that does not occur

in peninsular India. The species is not found

in Sri Lanka, but stretches from India deep

into the Oriental region.

041 . Pareronia ceylonica Felder & Felder

( Parenonia (sic!) ceylonica)

The dark wanderer is endemic to Sri

Lanka and southern India where it is more

of a wet zone butterfly than is the preceding

species. The South Indian population is listed

as ssp. pingasa Moore by d’Abrera, but I

agree with Talbot that it is not worthwhile

separating it from the nominate Sri Lankan.

It will be found in the mixed deciduous forests

as well as in tropical evergreen at Nadgani.

It is normally somewhat scarce but may be

quite common at Kallar. The flight is fast and

restless through dense vegetation and even

when coming to flowers it is wary, so pro-

curing a good series is not always easy. I have

never seen it at water, though I have seen a

picture of one of the Pareronia in a mudpud-

dling assemblage. On the wing the female is

a most effective mimic of Parantica aglea,

very much more so than one would suspect

from cabinet specimens.

COLIADINAE

042. Catopsilia pomona Fabricius

(C. pomona & crocale)

The lemon emigrant is a large and power-

ful butterfly that is strongly migratory. Almost

a million participated in the relatively modest

migration that I documented in May /June of

1986 (Larsen 1987b), and many millions were

involved in a month or more of intermittent,

generally southwards migration during Sept-

ember and October. Its main base is the drier

lowland formations, but it can be found

breeding practically anywhere except on the

highest plateau. It is an avid mudpuddler and

has a great liking for flowers. There are two

chief forms: f. crocale has black antennae and

has immaculate undersides; f. pomona has

reddish antennae and the underside has dark

irroration as well as spots at the end of the

cell. Intermediates are found but they are not

very common. There is no doubt that they

represent forms of one species, but the rela-

tive frequency of the two forms varies and

their respective functions are quite unknown.

The large numbers that settled on damp

patches at Kallar in May 1986 were 95%

crocale, while during the wet season the two

forms were roughly equal in number. Both

forms participate in the migrations so we are

not faced with the type of phases so well

known from migratory locusts. A thorough

study of the dimorphism in this species

would be most interesting. The range covers

the entire Oriental region. New Guinea, parts

of Australia to well out in the Pacific. A

Malagasy species ( Catopsilia thaurama Rea-

kirt) is sometimes, in my view not correctly,

linked to C. pomona.

043. Catopsilia pyranthe Linne

(C. pyranthe & C. florella)

The mottled emigrant is a common butter-

fly with both sexes showing a high degree of

variation. The florella form with very narrow

forewing borders and with well-developed,

red-ringed silver spots at the end cell of the

underside has traditionally been accorded

specific status, especially since it is relatively

constant in Africa, while the pyranthe form

is fairly constant in eastern Asia. However,

52

BUTTERFLIES OF THE NILG1RI MOUNTAINS

in India’s drier tracts both forms and inter-

mediates occur with little consistency, except

that the florella form is most frequent during

the dry season. The species is common in the

Nilgiris, sometimes very much so, usually in

the py rant he morph. The species is strongly

migratory and the progeny of migrants some-

times breed even on the plateau, though it is

mainly a species of the lowland drier tracts.

It is fond of both flowers and damp patches.

The distribution covers all of tropical Africa,

southern Arabia, the entire Oriental region,

New Guinea and parts of Australia.

044. Eurema brigitta rubella Wallace

( Terias libythea)

The small yellow is a common butterfly

with the widest possible of distributions, both

geographically and ecologically. In the Nilgiris

it may be found at all levels in most types

of habitat, though it tends to avoid the dense

evergreen forests. It is, in fact, one of those

butterflies that seem to thrive particularly well

in areas disturbed by the activities of man.

It is not very common and somewhat local

in the Nilgiris. Flowers are avidly visited, but

the species is rarely seen at damp patches. It

has been known to migrate but I have seen

no evidence of this in the Nilgiris. E. brigitta

was one of the few permanent residents of

my compound in Kotagiri, the population

usually consisting of four to eights individuals

at any given time. One specimen was seen

captured in flight and eaten by the Whitespott-

ed Fantail Flycatcher ( Rhipidura albicollis).

The range covers all of tropical Africa, south-

western Arabia, most of the Oriental region

and parts of New Guinea and Australia. From

Sundaland eastwards it becomes a rather

scarce and local grasslands species, possibly

because of competition from the many other

members of the genus in that area (Holloway

1973 gives an interesting review of the genus).

045. Eurema laeta laeta Boisduval

( Terias laeta)

The spotless grass yellow is normally a

common species, but on the whole this is not

the case in the Nilgiris. Wynter-Blyth was a

long time before finding it in numbers near

Coonoor and at Gudalur. I have found only

modest colonies in one or two places on the

Nadgani Ghat and at Masinagudi. Obviously

colonies may be found in nearly all types of

terrain. There is strong seasonal dimorphism,

but 1 have seen only wet season forms in the

Nilgiris. The species has a limited distribution

compared to some of the other Eurema, be-

ing found only in Sri Lanka and from India

to Burma, Thailand and Indo-China.

046. Eurema hecabe simulata Moore

( Terias hecabe)

The common grass yellow is just that,

very common practically everywhere. Since it

also has a vast range it is probably among

the top ten of the world’s most numerous

butterflies. In the Nilgiris it may be found

literally anywhere, though it is least common

in dense evergreen forest where it is replaced

by E. blanda. Indeed, E. hecabe is often parti-

cularly common in habitats that have been

modified by human activity. The species is a

known migrant, but large scale migration has

not been observed in the Nilgiris, though

some did follow the May 1986 migration

(Larsen 1987b). It visits flowers and often

comes to damp patches. The range covers

the entire old world tropical zone from Africa

and Arabia to India and the Oriental region,

to New Guinea, Australia, Japan, Fiji and

Tonga.

047. Eurema blanda silhetana Wallace

( Terias blanda)

The three spot grass yellow is restricted

to lowland forests where it is sometimes very

common indeed. The flight is often higher

53

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

above the ground than in the other species

of the genus, probably because the larval food

plants are tall trees like Albizzia and asso-

ciated creepers like Wagatea spicata. Contrary

to the other Eurema the present species lays

its eggs in large batches and the larvae and

the pupae are gregarious throughout, the

black pupae being situated so close to each

other as to almost touch, sometimes up to

fifty at a time. The species may assume pest

proportions in Albizzia plantations. E. blanda

is found in Sri Lanka, South India’s wetter

tracts. Eastern Ghats and then east to practi-

cally all the Oriental region to at least Papua

New Guinea.

048. Eurema andersonii ormistoni Watkins

(Terias andersoni)

The one spot grass yelijOW is the odd man

out among the South Indian Eurema inasmuch

as it is a genuinely rare and local species. In

the Nilgiris it seems limited to evergreen

tropical forest below 1500 m, which means

that it just penetrates the subtropical level.

Wynter-Blyth collected a few at Kallar where

I have also taken the species. I also have a

few from the Glenburn forests as well as one

from an evergreen patch on the Kotagiri Ghat.

The flight is weak and irresolute, the butterfly

coming to both flowers and wet patches. It

occurs from Sri Lanka through suitable wet

tracts of peninsular India to Malaysia, and

it seems to be scarce and local everywhere.

049. Colias nilagiriensis Felder & Felder

( Colias erate )

The nilgiri clouded yellow is quite

common on the plateau above 1900 m, flying

throughout the year, especially in places with

some moisture. The biggest concentration I

have seen is in the beautiful bog at Longwood

Shola near Kotagiri where hundreds may be

seen on a good day, buzzing about just above

the surface. The larval food plant here is

Parochetus communis , though I suspect other

plants are used elsewhere. All South Indian

females of this butterfly are white, though

in all its closest relatives females occur in

both white and yellow forms. While related

to the Palaearctic C. erate Esper I can see little

reason for not accepting the isolated and

morphologically very distinctive South Indian

taxon as specifically distinct. As such it is a

South Indian endemic species of obvious

Palaearctic origin.

{to be continued)

54

NOTES ON COMPARATIVE BODY SIZE,

REPRODUCTIVE EFFORT AND AREAS OF MANAGE-

MENT PRIORITY FOR THREE SPECIES OF KACHUGA

(REPTILIA, CHELONIA) IN THE NATIONAL

CHAMBAL SANCTUARY1

R. J. Rao2 and L. A. K. Singh3

(With a plate & three text-figures)

Record lengths of carapace have been reported from the River Chambal for

Kachuga tentoria circumdata (26.5 cm), K. dhongoka (48.0 cm) and K. kachuga

(49.0 cm). The ratios, carapace length/breadth and plastron length/breadth were

similar in all three species. The ratio carapace length/ shell height was less in K.

tentoria as it bears a high dome. Out of 13 areas of nesting identified for manage-

ment-priority, 23.0% were shared by all three species and 46.1% by K. tentoria and

K. dhongoka. Although biometrical features relating ‘body capacity’ to ‘egg mass

occupancy’ in the body indicate similarities between K. tentoria and K. kachuga,

only K. tentotia is known to lay more than one clutch per season while others lay

only one. The incubation period is 5-8 months for tentoria as winter-laying is there,

but is 2 months for the others. The clutch sizes are 4-9 (tentoria), 21-35 (dhongoka)

and 11-18 (kachuga). Egg length x breadth cm x weight g are: 4.7 x 2.7 X 21.4

(tentoria), 5.9 x 3.6 x 44.2 (dhongoka) and 7.0 x 4.1 x 57.4 (kachuga).

I NTRODUCTION

Three species of Kachuga (Emydidae)

namely, K. tentoria circumdata, K. dhongoka

and K. kachuga occur in the National Cham-

bal Sanctuary along the River Chambal of

the Gangetic system (Rao and Singh in press).

In the following we present data on the

body size of these chelonians with preliminary

analysis of the observations on their repro-

duction. Besides, a list is given of the

nesting areas that are of significance to a

Manager for the species in the Sanctuary.

The National Chambal Sanctuary, created

in 1978 extends from Jawaharsagar Dam

(Rajasthan) to Kota barrage (Rajasthan) and

1 Accepted September 1984.

2 Camp: National Chambal Sanctuary. Post Box

11, Morena 476 001, Madhya Pradesh.

Crocodile Research Centre, Hyderabad 500 264.

after a gap of 18 km, from Keshoraipatan

(Rajasthan) through Pali (M.P./ Rajasthan) to

Pachhnada (Uttar Pradesh). The total length

of the river inside the Sanctuary is about

600 km.

Materials and Methods

Three live and 22 caracases of K. t. circum-

data and three caracases each of K. dhongoka

and K. kachuga were collected outside water

during survey trips made by boat and foot

between October 1983 and July 1984. The

shell measurements of all specimens were

taken as described by McRae et al. (1981)

and the data were used to compare the maxi-

mum sizes reported in literature for different

species (Smith 1933, Pritchard 1979, and

Daniel 1983), and determine the similarities

in certain biometrical ratios. The carapace

55

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

length and carapace width (= plastron width)

were taken as straightline measurements.

Four gravid females, represented by two of

tentoria and one from each of the two other

species were examined at the ovary (for

methods see, Moll and Legler 1971. Rao 1982)

to determine the nesting frequency and collect

comparative data on clutch and egg sizes in

relation to the length of the females.

In order to compare the species-wise re-

productive effort, the approximate body capa-

cities (BV) have been considered against the

approximate egg-clutch mass (EM) and egg-

clutch occupancy (EV). The following methods

were used in calculating BV, EM and EV.

PL x PB x SH,

BV f ,

2

EM = CSxEW, and EV = CSxELxEB

In the above, PL and PB are the mean

lengths and breadths of plastron, SH, the mean

shell height; CS, the mean clutch size (mean

number of eggs per clutch); EL and EB, the

mean length and breadth of eggs, and EW

the mean egg weight (g). All measurements

are in centimetres.

The incubation period was determined by

direct corroboration of nesting activities,

period of commencement of embryonic deve-

lopment and periods of sightings of hatchlings.

Different nesting areas and their manage-

ment priorities were determined during the

surveys from the extent of nesting activities

and predation pressure.

Results

Size and biometrical ratios. The average

shell measurements of dead and live speci-

mens of hardshelled turtles collected in the

Chambal river are presented in Table 1. The

mean carapace lengths were 24.65 cm

(Kachuga tentoria circumdata), 44.6 cm. (K.

dhongoka ) and 47.6 cm ( K . kachuga). The

maximum length of the carapace in the three

species were 26.5 cm, 48.0 cm and 49.0 cm,

respectively. The plastron was always smaller

than the carapace but attached to the latter

at such a point in the front that it protruded

out a little behind the rear end of the cara-

pace. Therefore, the total lengths (Table 1)

were larger than the carapace length. The

ratios carapace length/width were 1.38, 1.38

and 1 . 32 for K. tentoria circumdata, K.

dhongoka and K. kachuga respectively (Table

1). The ratio, plastron length /width were 1.26,

1 . 27 and 1 . 25 respectively and the ratio,

carapace length /shell height (body depth)

were 2.24, 2.73 and 2.34. For one male K.

kachuga the ratios were: carapace length/

width 1.25, plastron length /width 1.14 and

carapace length/shell height 2.36. The male

(29.5 cm carapace length) was caught by

net near Babu Singh ka gher (Table 2) on

5 December 1983. The turtle, being in its

breeding colours, had six red longitudinal

stripes along the neck, a pair of oblong yellow

spots on the throat and the head was brilliantly

red on the top and bluish on the sides.

Nesting areas, Distribution pattern. Kachuga

are very scarce in the river upstream of Pali.

Downstream of Pali although pre-nesting acti-

vities were seen, no nests were located. The

first major nesting site was located at Baroli

(Table 2), 57 km downstream of Pali.

Out of the total 13 identified nesting sites

(Table 2) three (23.0%) were common to

all three species, and six (46.1%) were com-

mon to K. tentoria and K. dhongoka. There

were no regular distance-intervals between

any two nesting sites (Figs. 1 and 3).

Nesting frequency. Three predated nests,

about a week old, discovered on 29 October

1983 were the earliest record of nesting by

Kachuga tentoria. Fresh nests (confirmed

from fresh turtle tracks) were located on 5

56

NOTES ON THREE SPECIES OF KACHUGA

Table 1

Body and egg biometrics of Kachuga tentoria circumdata, K. dhongoka and K. kachuga. for SL. no.

1 through 8 n = 25 (tentoria), 3 ( dhongoka ) and 3 (kachuga). sizes in cm and weight in g (range

in parentheses)

57

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vo!. 84

Table 2

Major nesting sites of Kachuga sp. identified in national ciiambal sanctuary, 1983-84. d, K.

dhongoka\ k, K. kachuga and t, K. tentoria circumdata. for tentoria, ** first nesting, ***second nest-

ing. al, alluvium; bs, bush and os, open sand.

* In reference to Palighat (Parbati-Chambal confluence) (Fig. 1).

December (10), 3 January (1) and 28 Janu-

ary (1). After 28 January no further nesting

activities were noted. From all the above nest-

ing records, it was presumed that nesting

season of Kachuga tentoria circumdata extend-

ed from October through January (Fig. 2).

A female Kachuga tentoria circumdata (25

cm carapace length) wandering on land at

0730 hrs on 24 November contained six

(3+3 right and left) oviductal eggs

with a mean weight of 23.3 g and length x

width 50±0.564 (50.0-51.5) x 29.08±0.205

(29.0-29.5) mm. The ovaries contained only

six fresh corpora lutea (right 4 and left 2).

Four ovarian follicles of ovulatory size (25-26

mm) were also present suggesting the pro-

duction of at least two clutches per season.

On the same day (24 November) at 1600

hrs, a female turtle (carapace length 23.0 cm)

was captured on land when it was returning

after egg laying. The nest contained 7 eggs

with the mean measurements of 46.0 ±0.925

(45.0-47.0) x 28. 0_ 0.534 (27.0-29.0) mm.

Mean egg weight was 22. 3 ±0.527 (22.0-

58

NOTES ON THREE SPECIES OF KACHUGA

Fig. 1. River Chambal (National Chambal Sanctuary) from Pali (Chambal-Parbati

confluence) to Pachhnada showing important nesting areas of Kachuga sp. (1-13, as

in Table 2). M.P., Raj. and U.P. are the states of Madhya Pradesh, Rajasthan and

Uttar Pradesh through which the river flows.

Inset: Position of River Chambal (arrows) in the Gangetic system.

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

60

Fig. 2. Relationship among climatic conditions and breeding activities in small (S) and large (L) Kachuga turtles. N,

nesting; H, hatching. Horizontal scale shows months from August (A) 1983 through July (J) 1984. Maximum tempera-

ture (°C) shown as dotted line and minimum temperature (°C) as solid line through hollow circles. Hour of sunshine

(hr) : dotted line through solid circles, relative humidity (%) : solid line through hollow triangles, and rainfall (mm) :

solid line through crosses.

NOTES ON THREE SPECIES OF KACHUGA

l J J_J L _J

5SSSS58!

I

r-ssai

G\

6 2?

** <4-1

<U O

00

2 J

co to

1 a

fl

H O 2

•8-S*

61

JOURNAL , BOMBAY NATURAL HIST. SOCIETY, Vol. 84

23.5) g. The ovaries contained seven corpora

lutea (right 4 and left 3) and four ovarian

follicles of ovulatory size (24-26 mm) indi-

cating two clutches per season.

Clutch size and egg size of Kachuga t.

circumdata are shown in Table 1. The clutch

size was determined mostly from the exami-

nation of natural nests (18 no.) and also from

the data on eggs found in the oviduct (n=l)

of live- and in the body cavity (n=l) of dead-

specimen (see above). Mean clutch size was

5.95 with a range of 4-9. Eggs were elliptical

and with a very thin shell crack-

ing to pressure. Mean egg length was

47.5 mm, width 27.54 mm and weight 21.40

g. One female K. t. circumdata, dead presu-

mably a week before location (12 December

1983) had eight shelled eggs free inside the

body. The female measured 26.5 cm in cara-

pace length and the eggs 46.1 x 29.8 mm

(Plate 1).

The breeding season in K. kachuga is ex-

pected to commence by the beginning of

December because the male caught on 5 Dec-

ember was already in its breeding colours. The

nesting season of both the large Kachuga sp.

( dhongoka and kachuga) was March- April.

The first nest of Kachuga dhongoka was

found on 11 March 1984 and after 31 March

no fresh nesting was recorded. The nests of

large Kachuga sp. looked the same as that of

small Kachuga sp. except that these were of

large dimension and most often on ground

with relatively more silt.

Eggs in the size range of 66-75 mm in

length with small clutch size (11-18) were

the eggs of Kachuga kachuga. These eggs were

37-45 mm wide and 56-58 g in weight. Mean

clutch size of Kachuga dhongoka was 23.64

and of Kachuga kachuga was 15.5. Egg

dimensions and clutch size are shown in Table

1. Eggs of K. dhongoka were small, 51-66 mm

x 32-41 mm x 36-57 g (Plate 1).

The body cavity of one K. dhongoka was

examined late in the nesting season. The

turtle was dead about a week before location

(31 March 1984) and had 21 free eggs in-

side the body. The carapace length of the

female was 40.5 cm and the eggs were 60 . 7 x

38.7 cm.

A Kachuga kachuga suspected to be dead

the night before location (5.4.1984) had

well calcified eggs, 4 in the right oviduct and

7 in the left. Except a few atretic follicles

there were no follicles greater than 20 mm

but there were 3 fresh corpora lutea in the

right ovary and 8 in the left ovary. The

female measured 49.0 cm in carapace length,

and the eggs were 66.5 mm x 37.6 mm.

Incubation period. The minimum and maxi-

mum ambient temperature during the nesting

season of K. tentoria (October- January) rang-

ed as 3.5-29°C and 17-39°C, respectively

(Fig. 2). The prevalent conditions of humi-

dity, rainfall and photoperiod (sunshine) dur-

ing nesting are shown in Fig. 2.

The nest temperature during winter nesting

was very low (17°C-25°C) for embryo deve-

lopment. Periodical checks of eggs in an open

hatchery revealed that there was no embryo

development until March but on 3 April, 1984

(30°C nest temperature) development had

already commenced. In the wild hatchlings

have been seen in May.

During the nesting season of K. dhongoka

and K. kachuga (March-April) the ambient

temperature ranged between 11°C and 46°^

(Fig. 2), and the nest temperature 27.5°C-

32°C. The development of embryo was notic-

ed to have started within two days after egg-

laying. Emerged hatchlings of K. dhongoka

were recovered from nests at Tigri-Rithaura

on 4.6.1984. Nesting at Tigri-Rithaura

had occurred during the last week of March.

The incubation period is estimated to be about

sixty days.

62

J. Bombay nat. Hist. Soc. 84

Rao & Singh: Kachuga spp.

Plate 1

Above : Kachuga terttoria circumdata with eggs recovered from the body.

Below. A nest pit of Kachuga dhongoka with clutch of 23 eggs.

NOTES ON THREE SPECIES OF KACHUGA

Comparative reproductive effort. The appro-

ximate BV of K. tentoria, K. dhongoka and

K. kachuga were 2195.38, 10835.16 and

16443 respectively. The EM and EV were

127.33 and 77.91 ( K . tentoria), 1045.12 and

511.58 ( K . dhongoka) and 890.01 and 454.62

(K. kachuga) (Table 1).

The factor showing the proportion (%)

of EM in BV were 5.79, 9.64 and 5.41 for

tentoria , dhongoka and kachuga respectively

(Table 1). The factor P2 showing EV in BV

were 3.5% {tentoria), 4.7% {dhongoka) and

2.7% {kachuga) (Table 1).

Discussion

The largest specimen of K. t. circumdata,

K. dhongoka and K. kachuga were 26.5 cm,

48.0 cm and 49.0 cm respectively. These

sizes are distinctly larger than the sizes given

in literature (Table 1). We would be inte-

rested to know when larger specimens are

reported.

The ratios CL/CB and PL/CB are almost

the same in all three species, perhaps a basic

requirement in the design of the hardshelled

turtles. Differences in CL/SH are evidently

due to the extent to which the dome is raised

above the plastron (SH) and this makes the

differences in body-capacities.

The production of multiple clutches is to

be expected for forms that have less body

capacity and lay only a few eggs relatively

less secure. In K. t. circumdata the nests are

83 . 5% times on flat sand banks close to main-

land (Rao and Singh 1984) and the clutch

of 4-9 eggs are only 16.13 cm below the

surface. Therefore, a double effort is made

in the species to ensure enough annual re-

cruitment after sustaining all kinds of loss.

These efforts are in the number of turtles and

in laying double clutches spaced over a period

of few months. Singh (1985) has also

reported double nesting by K. tentoria in

Mahanadi.

In the case of K. dhongoka the necessity

to lay more than a clutch is waived because

the size of the clutch and eggs are large. K.

kachuga, although larger in size than K.

dhongoka and produces larger and heavier

eggs, the clutch size is small. The factors Px

and P2 as determined by comparing the body

capacities and clutch-occupancies, are closely

similar in the case of K. tentoria circumdata

and K. kachuga indicating that K. kachuga

may be having the potentiality to produce

more than one clutch like K. tentoria. If, how-

ever, the results of examination of the ovaries

and oviduct in the specimen of 5.4.1984 are

any indication then we can at the best con-

clude that between October and July K.

kachuga lays only one clutch and it is not

known if a second clutch is ever laid.

Hatchling turtles of Chrysemys picta and

Pseudemys scripta (Gibbons and Nelson 1978)

and Chelydra serpentina (Newman 1906) are

known to over-winter in the nest itself, and

Congdon et al. (1983) suspect a direct rela-

tionship between the emergence tactics of

hatchlings and the level of lipid in the eggs.

Singh (1985), based on his notes on ter-

restrial activities in K. tentoria of the River

Mahanadi, suspected that the species

perhaps hibernates. Based on the present

study it is suspected that the eggs of K. t. cir-

cumdata laid early in the season (October)

may be undergoing some development before

the onset of the extremely low temperatures

of December -January but during the winter,

development proceeds at an extremely slow

rate. The eggs laid during November and later

have to remain dormant until the tem-

perature rises to 30°C as has been observed

in developing embryos of April. The above

situation would lead to the hatching of K. t.

circumdata spread over a few months that

would ensure less competition. In any case,

the incubation period is suspected to be 5-8

63

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

months depending on the date of egg laying

and temperature. In large Kachuga sp. since

the temperature is already high development

commences soon after egg-laying. This situa-

tion calls for the greatest care in the event of

a necessity to shift or translocate these eggs.

More stringent than the practice with croco-

dilian eggs, the eggs of K. dhongoka and K.

kachuga may be shifted only on the day fol-

lowing deposition or after 50-55 days

incubation.

The reason why gravid females were dead

at the nesting site without even commencing

to dig a nest-pit, is difficult to explain. Egg-

bound deaths are known in captive reptilians

which are under stress due to lack of suitable

nesting environment but why should such

parallel, though stray cases, occur in nature

is an aspect to study in future. At the pre-

sent, these data provide stimulation to accu-

mulate similar information to compare in

future the body size to the clutch and egg

sizes.

Singh (1978) has found that in the

case of gharial ( Gavialis gangeticus) even

if egg sizes vary among nests there is a

specific “upper-limit” to the “egg capacity”

(considered as the product of egg length and

breadth). In the case of the three species of

Kachuga studied by us the egg-design is simi-

Refer

Congdon, J. D., Tinkle, D. W. & Rosen, P. C.

(1983): Egg components and utilization during

development in aquatic turtles. Copeia 1983 (1):

264-268.

Daniel, J. C. (1983) : The Book of Indian Rep-

tiles. Bombay Natural History Society, Bombay.

Gibbons, J. W. & Nelson, D. H. (1978): The

evolutionary significance of delayed emergences

from the nest by hatchling turtles. Evolution 32:

297-303.

McRae, W. A., Landers, J. L. & Cleveland, G.

D. (1981): Sexual dimorphism in the Gopher tor-

lar to the gharial’ s, longish with round ends,

and there does appear certain extent of varia-

tion in the egg sizes within any Kachuga

species but at present it is not known if there

is an “upper limit” to the egg-capacity.

Most of the Chelonian nesting areas are

either common among themselves (Table 2)

or with the gharial and are under heavy pre-

dation pressure (Rao and Singh in press).

Therefore, a Manager involved in the conser-

vation of the gharial can, with very little

extended effort, attend to the nesting areas of

the turtles too. Once the action is implement-

ed it could form the basis to take up ex-

panded conservation management.

Acknowledgements

We express our gratitude to Government of

India (Wildlife Institute of India), Food and

Agriculture Organisation of the United

Nations, and Government of Madhya Pradesh

(Wildlife wing) for the provision of various

facilities and the permission by the latter to

conduct the study. Several staff members and

colleagues gave assistance in the field. RJR

remained a CSIR Senior Research Fellow for

most part of the study duration and LAKS

is with the Crocodile Research Centre of the

Wildlife Institute of India.

E N CE S

toise ( Gopherus polyphemus) . Herpetologica 36(3):

46-52.

Moll, E. O. & Legler, J. M. (1971): The life

history of a Neotropical slider turtle, Pseudemys

scripta (Schoepff), in Panama. Nat. Hist. Mus. Los

Angeles Cty. Sci. Bull. 11: 1-102.

Newman, H. H. (1906): The habits of certain

tortoises J. Comp. Neuro. Psych. 16: 126-152.

Pritchard, P. C. H. (1979): Encyclopedia of

Turtles. T.F.H. Publications Inc., Neptune, New

Jersey.

Rao, R. J. (1982) : Studies on the neuroanatomy,

64

NOTES ON THREE SPECIES OF KACHUGA

cytology and histochemistry of the reproductive sys-

tem of the freshwater turtle, Trionyx gangeticus

(Cuvier) with special references to the seasonal

cyclicity. Ph.D Thesis, Bhopal University, Bhopal.

& Singh, L.A.K. (1984): Ecological

Relationship among Turtles in National Chambal

Sanctuary. Interim Study Report 1, CRC/NCS/21-

A, August 1984, Field Camp of the Crocodile Re-

search Centre, Wildlife Institute of India, Govern-

ment of India, Deori, Morena, Madhya Pradesh,

iii+54 pp, 2 figs.

— — — — __________ (jVj press) : Notes

on ecological relationship in basking and nesting

site utilisations among Kachuga sp. (Reptilia, Chelo-

nia) and Gavialis gangeticus (Reptilia, Crocodilia)

in National Chambal Sanctuary. /. Bombay nat.

Hist. Soc.

Singh, L.A.K. (1978) : Ecological studies on the

gharial, Gavialis gangeticus (Gmelin) (Reptilia, Cro-

codilia). Ph.D thesis, Utkai University, Bhubaneswar.

(1985): Notes on tracking and

terrestrial activities of the freshwater turtle, Kachuga

tent or ia in River Mahanadi, Orissa. /. Bombay nat.

Hist. Soc. 82(2) : 414-417.

Smith, M. A. (1933) : The Fauna of British

India including Ceylon and Burma. Reptilia and

Amphibia. Vol. I. Loricata, Testudines. Taylor and

Francis, London.

Note (12 June 1987) : Further studies have con-

firmed double-nesting in K. kachuga between Dee-

ember and March.

65

5

THE GENUS PIPER LINN. IN KARNATAKA, INDIA1

B. A. Rahiman2 and M. K. Nair3

(With eight text-figures)

The family Piperaceae, until 1940, was limited to two genera viz., Piper Linn.,

and Peperamia Ruiz. & Pav. In 1843, F.A.W. Miquel subdivided it into several

ranks and described about 600 species under 20 different segregate genera. Since

then, the family has undergone considerable changes in the circumscription of various

taxonomic ranks. In India, 108 species have been reported from two inde-

pendent ‘centres of distribution’. They are 1) the region of Sub-Hifnalayan and

North-eastern hill ranges and 2) South Deccan. In the present paper an attempt is

made, to revise the taxonomy of Piper species occurring in the Karnataka region

of Deccan, based on the authors’ survey of the region under study and collection

of over 300 herbarium specimens. In all 8 species have been described. An artificial

key for the Karnataka species, field notes, nomenclatural notes, comments on affinity,

world distribution, distribution in India and in Karnataka and type locality are

provided.

The genus Piper Linn., the largest in the

family Piperaceae, occurs throughout the tropi-

cal and subtropical regions. More than 3000

species are on record (index kewensis 1895-

1970, Rahiman 1983). Because of the large

number of species, wide distribution, very

minute, achlamydous and closely packed

flowers, unisexual nature of many of the

species and lack of any recent critical phyletic

study (Hooker 1886; Gamble 1925; Law-

rence 1951), a valid acceptable species con-

cept could not be established till to date. Until

1940, Piperaceae was limited to two genera

namely Piper Linn, and Peperomia Ruiz. &

Pav. In 1843, F.A.W. Miquel in his monograph

system a piperacearum subdivided the family

into several ranks such as tribe, section, cohorts,

1 Accepted November 1984.

* Post Graduate Department in Bio-Sciences,

Mangalore University, Mangala Gangotri 574 152,

Karnataka, India.

3 Central Plantation Crops Research Institute, Re-

gional Station, Marikkunnu, Calicut, 673 012, Kerala,

India.

genus and species and described about 600

species of Piper under 20 different segregate

genera. As the report of new species from all

over the world accumulated and the number

of species went on increasing, the segregate

genera started losing their distinctions, mainly

because of the lack of sharp taxonomic dis-

continuities within the family. In 1869, C. de

Candolle in dc prodromus merged these

segregate genera into 9 major genera. Hooker

(1886) further reduced the number of genera

to three and described the various Piper

species under 4 sections. Contemporary taxo-

nomists discontinued even, the practice of sub-

dividing the genus into sections and dealt the

species directly under the genus. From taxo-

nomic point of view, Piperaceae has been

considered as a most difficult family.

According to Howard (1973), the family is

one of the worst messes in plant taxonomy.

Because of all these problems and also due

to the poor quality of the herbarium speci-

mens of pioneering workers. Sir Joseph

Hooker (1886) advised the local botanists in

66

GENUS PIPER LINN. IN KARNATAKA

various ‘centres of distribution’ of species in

Indo-Malayan region to “examine the plants

on the spot, with a view to matching the

sexes, and flowering with fruiting specimens,

and to observing the transition from young

to old foliage, and the effects of locality and

climate on the character of each species”. In

India, two major, independent centres of dis-

tribution of species are recognized. They are:

1) the region of Sub-Himalayan and North-

eastern hill ranges and 2) Southern Deccan.

More than 108 species are recorded from

India (Rahiman & Nair 1983). In the pre-

sent investigation, systematics of all the species

occurring in the Karnataka region of Deccan

were taken up. However, the study does not

include P. betle Linn, (betel vine) which has

not been recorded in the wild state in India

and cultivars of P. nigrum Linn, (common

black pepper vine) . The observations are based

on a survey of the Piper species in Karnataka

by the authors (Rahiman el al 1979, 1981) in

which more than 300 herbarium specimens

were specially collected for the study.

Key to the Species

1 . Spike pendulous, berry large or medium-sized,

bracts adnate or decurrent at lower half, not

pedicelled :

2. Bracts adnate to the rachis at the medial

position, margin narrow but free, berry

does not attain red colour:

3. Female spikes long, developing berry

oblong attenucutum

3 . Female spikes short, developing berry

ovate :

4 . Leaves, peduncle and young branches

hirtellous hookeri

4. Leaves, peduncle and branches glabrous,

rarely minutely puberulous

ctrgyrophyllum

2. Bracts decurrent at lower half, berry large,

becomes reddish nigrum

2. Bracts transformed into a fleshy cup:

5. Outer surface of the cup puberous

trichostachyon

5. Outer surface of the cup glabrous

galeatum

1 . Spike erect, berry small, bracts peltate with a

short pedicel:

6. Fruiting spike sub-globose, leaf base

acute mullesua

6. Fruiting spike cylindric, leaf base symme-

trically or asymmetrically cordate

Ion gum

P. attenuation Ham. ex Miq., Syst. Pip. 306.

1843 & in FI. Ind. Bat. 1(2): 451. 1859;

J. Hooker, FI. Brit. India 5: 92. 1886;

Gamble, FI. Madras 1205. 1925. P. diffusum

Vahl. Enum. 1: 333. 1804. P. Karok Blume,

Cat. Gew. Buitenz 33. 1823. P. malamaris

Roxb., FI. Ind. 1: 160. 1832. P. Sirium C.

DC., in DC., Prodr. 16(1): 160. 1869.

(Fig. 1).

A slender climber, dioecious. Leaves thin,

pressed ones membranous-chartaceous, ovate,

rarely cordate, 9.5-17.5 cm long and 4. 0-8. 5

cm broad, glabrous, sometimes minute white

dots seen on the dorsal surface, ventral side

green, dorsal, dark green, 2-3 pairs of promi-

nent lateral ribs, all arising from the base or

very near to it; stipule adnate, deciduous.

Flowering spike narrow, filiform, c? upto 26.0

cm, $ upto 10.0 cm, fruiting spike upto 21.0

cm, peduncle glabrous. Bracts linearly-obovate

to elliptic, sessile, adnate in the medial posi-

tion with narrow but free margin. Stamens 3,

rarely 4. Carpel single, astylocarpellous, ovary

oblong, stigma mostly 4-lobed, lobes short,

ovule solitary, errect. Berry, mature ones

spherical, developing ones characteristically

oblong, smaller than the commercial black

pepper, 0. 3-0.4 cm in diameter, bitter in taste.

Climbs over the supporting trees with the

help of strong adventitious roots, not more

than 5-9 m in height. The fruit is an indehis-

cent drupe but commonly treated as berry.

67

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

Fig. 1. P. attenuatum Ham.

A. Branch with fruiting spikes; B. Branch with male spikes; C. Female spike; D. Male

spike; E. Portion of female spike; F. Portion of male spike; Ga. Female bract — side

view; Gb. Female bract — bottom view; Gc. Male bract — bottom view; Ha. Ovary; Hb.

Ovary — L.S.; I. Stamen; Ja. Young berry; Jb. Mature berry.

68

GENUS PIPER LINN. IN KARNATAKA

Young berry green in colour, mature ones

black. Unlike P. nigrum Linn, the berries do

not attain red colour during the development.

Number of mature berries drop before fully

mature. Runners arc in plenty, cuttings of the

runners start flowering in the very next season

of planting. Flowers in May- July period. Off

season flowerings common.

Saldanha and Nicolson (1976) considered

P. attenuatum as synonym of P. trioecum

Roxb. and Gamble (1925) considered the

latter under P. attenuatum. Miquel (1843)

and C. de Candolle (1869) considered them

as two distinct species. F. trioecum was esta-

blished by Roxburgh (1820). In his flora

indica he described a male, a female and a

bisexual vine. A close perusal of the descrip-

tions of these 3 vines furnished by him

clearly showed contrasting diagnostic charac-

ters — the bisexual vine showed 2 stamens

and the berries which are pungent, ripened

perfectly in the spike and the male one

showed 3 stamens and the berries which are

non-pungent, dropped before full maturity in

the spike. (The former set of characters are

similar to the characters observed in F.

nigrum and the latter, are similar to that of

F. attenuatum). From this it is obvious that

these vines do not belong to one and the same

species as William Roxburgh believed. The

type specimen of F. trioecum is not available

in any Indian Herbarium and the figure given

by Wight (1853) in the icones plantarum

indiae orientalis, though said to be taken

directly from Roxburgh’s drawings, is not of

much help to determine the identity. Hence

the epithet F. trioecum is ignored in the pre-

sent study.

Distribution'. Bangladesh, Bhutan, Sri

Lanka and Malaysia. In India, the Hima-

laya, Sikkim, Assam, Meghalaya and the

Western Ghats. In Karnataka, very common

in low level forests (lower than 200 m MSL)

of Uttara Kannada and Dakshina Kannada

districts.

Type locality. The Himalaya.

Selected specimens examined. Wallich 6642

D (CNH); Ansari 78563, Subramanian 77003

(BSI); Barber 5051; Henry 17394, 48263,

Hooker & Thomson ex ciccata sin. num., acc.

no. 70410, Joseph 17190, 44488, Narayana-

swami 3537, 5373, Raju & Naganathan 18146,

18156, 18206, Ramamoorthy 16153, Sebastine

15688, 16520, 25055, Subbarao 24528., 30052,

32877, 44384, 42575, Subramanyan 3846,

8066, Vajravelu 32131, Wight sin. num., acc.

no. 43777, 43778, 43779, 10881, 15723 (MH);

Nicolson & Ramamoorthy 246, Saldanha

10068, 11412, 13899, 14232, 14415 (CTS)4;

Rahiman 29, 33, 34, 37, 40, 49, 50, 65, 69,

70, 74, 115, 142, 143, 208, 229, 240, 254, 259,

262, 268 (MUK)5.

P. hookeri Miq., London J. Bot. 4: 437.

1845; J. Hooker, FI. Brit. India 5: 88. 1886;

Gamble, FI. Madras 1204. 1925. P. hyme-

nophyllum Miq., London J. Bot. 4: 437.

1845. F. lanatum Wight ex Miq., London

J. Bot. 5: 533. 1846. non Roxb. 1832. F.

nilighirianum C. DC, in DC, Prodr. 16(1):

364. 1869. F. Wightii Miq., London J. Bot.

5: 552. 1846 (Fig. 2).

Allied to F. attenuatum Ham. ex Miq. and

F. argyrophyllum Miq. Resembles F. atte-

nuatum in majority of morphological charac-

ters. The major differences are the presence

of crisp hairs on young branches, entire

ventral surface and along the major ribs on

dorsal surface of the leaves, petiole and bracts

in this species. Leaves thinly coriaceous, upto

17.0 cm long and 7.0 cm broad, very much

variable in size, shape and thickness.

cT spikes upto 14.0 cm, fruiting spikes upto

20.0 cm, stamens 3 in number.

4 CTS — Centre for Taxonomic Studies, Banga-

lore, Karnataka.

5 MUK — Mangalore University, Mangalore

(Konaje), Karnataka.

69

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

Fig. 2. P. hookeri Miq.

A. Branch with fruiting spikes; B. Branch with male spikes; Ca. Portion of leaf — dorsal

side showing hairs; Cb. Portion of leaf — ventral side showing hairs; Cc. Tip of the

branch showing hairs; D. Female spike; E. Male spike; F. Portion of female spike;

G. Portion of male spike; Ha. Stamen; Hb. Anther lobes — top view; la. Ovary;

lb. Ovary — L.S.; Ja. Bract — bottom view; Jb. Bract — side view; Ka. Young

berry; Kb. Mature berry.

GENUS PIPER LINN. IN KARNATAKA

Some problem pertaining to the identity of

this taxa was encountered in the literature.

Brandis (1906) and Cooke (1903) described

a species with hairy vegetative and floral parts

from the Western Ghats and considered it as

P. hookeri. Saldanha and Nicolson (1976),

on the other hand, considered a similar taxa

from Karnataka as P. hymenophyllum. Hooker

(1886) and Gamble (1925) described two

such taxa from the Western Ghats and iden-

tified them as two distinct species, namely

P. hookeri and P. hymenophyllum. Gamble

(1925) reported the former from Bababudan

hills (Bababudan hills are situated in the

Karnataka region of the Western Ghats) and

the latter from the Western Ghats of

Mysore (Karnataka). We studied a num-

ber of specimens with hairy nature from

all over Karnataka including Bababudan

hills. The morphological characters showed

wide variation but did not show any sharp

discontinuities to merit differentiation into two

species. P. hookeri and P. hymenophyllum

were established by Miquel in 1845 and 1846

respectively. In the original diagnosis, the

differences mentioned were the presence of

adnate bract in P. hookeri and linear-oblong,

adnate bract with undulated margin in P.

hymenophyllum and the coriaceous to mem-

branous leaves in the former and finely mem-

branous and transparent leaves in the latter.

In all other characters the two were identical.

The leaf characters, as Miquel himself remark-

ed were variable in these two species. In the

light of variability noticed in the morpholo-

gical characters in the genus, the differences

mentioned in the original diagnosis are quite

inadequate to differentiate these two into two

different species. Therefore it would be rea-

listic to consider these two species as conspe-

cific. Since P. hookeri was validly published

in 1845 and P. hymenophyllum , in 1846, the

former is to be considered as the valid specific

epithet. Therefore, in the present work, the

hairy species is considered as P. hookeri.

Distribution : Reported only from the

Western Ghats and Biligirirangan hills.

In Karnataka occurs in Uttara Kannada,

Shimoga, Hassan, Kodagu, Chikmagalur and

Mysore districts, mostly in the forests situated

at more than 200 m above sea level.

Type locality. Bombay.

Selected specimens examined : Arora 46275,

55232, Chibber sin num., Janardhan 72150,

Mahajan 24792, Pan 14462, Rao 79970, Rolla

87487, Raghavan 62477, 67853, 80838, 86982,

94119, Talbot 1602, 2459, Reddi 97261, 97826,

Exiccata sin. num., acc. no. 6962 (BSI); Barber

5450, 5489, 7555, 7539, 7559, 7560, 7563, 7595,

7346, Bourne 6112, 6085, Ellis 16957, Jacob

391, 427, 17666, Ramamoorthy 18188, 22882,

Sebastine 17275, Viswanathan 691 (MH).

Nicolson, Ramamoorthy & Gandhi 2884,

Ramamoorthy & Gandhi 2608, Saldanha

9049, 9053, 10704, 13581, 16753 (CTS); Rahi-

man 12, 890, 893, 905, 910, 911, 957, 958,

984, 998, 1000, Fc 230, 231, 236, 248, 258

(MUK).

P. argyrophyllum Miq., Syst. Pip. 330. 1843;

J. Hooker, FI. Brit. India 5: 93. 1886;

Gamble, FI. Madras 1205. 1925. P. walkeri

Miq., London J. Bot. 4: 438. 1845. P.

Wightii Miq., London J. Bot. 5: 552 in

part 1846. (Fig. 3).

Allied to P. attenuatum. It differs from P.

attenuatum by the presence of silvery scales

or blotches on the ventral surface of the

leaves. Leaves chartaceous to thinly coriace-

ous, upto 21.0 cm long and 7.0 cm broad,

sometimes sparsely distributed minute hairs

are also seen on the ventral side. Stamens 3

in number. Rarely P. argyrophyllum may lack

silvery blotches and P. attenuatum may show

similar blotches in some leaves and as such it

is very difficult to separate these two species,

especially based on herbarium specimens. It

71

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol 84

Fig. 3. P. argyrophyllum Miq.

A. Branch with fruiting spikes; B. Branch with male spikes; C. Female spike; D. Male

spike; E. Portion of female spike; F. Portion of male spike; G. Portion of male spike

showing stamens; Ha. Ovary; Hb. Ovary — L.S.; la. Bract — side view; lb. Bract —

bottom view; Ja. Young berry; Jb. Mature berry.

GENUS PIPER LINN. IN KARNATAKA

is also allied to P. hookeri and because of

the presence of sparsely distributed hairs,

poses difficulty in separating these two species

as well.

Distribution : In India reported only from

the Western and the Eastern Ghats. In Karna-

taka, occurs in Chikmagalur, Hassan, Shimoga,

Kodagu, Uttar Kannada and Mysore districts.

Occurs in higher altitude forests. Both P.

argyrophyllum and P. hookeri are seen in the

same locality.

Type locality. The Western Ghats.

Selected specimens examined. Fisher sin.

num., Singh 124569 (BSI); Barber 6498,

7202, 7313, 7344, 8710, 8712, 8713, 8715,

Ellis 34891, Gamble 18392, Henry 16282,

16286, Joseph 12705, 12793, 13789, Karthi-

keyan 26834, Narayanaswamy 3875, Shetty

32327, Rao 31999, Vajravelu 33840, 35123,

41724 (MH); Nicolson & Ramamoorthy 2866

(CTS); Rahiman 165, 167, 172, 207, 212, 213,

228, 230, 240, 244, 253, 255, 257, 258, 263 (all

the specimens fully glabrous), 209, 231, 232,

240, 242, 243, 264, 272 (all are puberulous in

young vegetative parts) (MUK).

P. nigrum Linn., Sp. PI. 28. 1753; Roxb., FI.

Ind. 1: 150. 1832; Gamble, FI. Madras

1204. 1925; Piper rotundum nigrum Caspa-

rus, FI. Mai. 54. 1696. P. aromaticum Lam.,

Illust. 1: 79. 1791. P. baccatum C. DC., in

DC., Prod. 16(1): 242. 1869. P. colonum

Presl, Bot. Bemerk. 112. 1844. P. fallax

Vahl, Enum. 1: 335. 1804. P. glyphicum

Hoffmag. ex Kunth, in Linnaea 13: 573.

1839. P. malabarense C.DC., in DC., Prodr.

16(1): 242. 1869. P. spurium Link, Enum.

Hort. Berol. 1: 37. 1821. Muldera multi -

nervis Miq., London J. Bot. 5: 557. 1846.

M. Wightiana Miq., l.c. 558. 1846. (Fig. 4).

A vigorous vine, stem thick, rough in tex-

ture. Leaves thick, pressed ones coriaceous,

broadly ovate to elliptic, 9.0-21.0 cm long

and 2.5-13.0 cm broad, in males usually

smaller and narrower, 2-3 pairs of prominent

lateral ribs, the anteriormost pair emerges

alternately about 2. 0-3. 5 cm above the leaf

base, entirely glabrous, dorsal side green,

ventral light coloured, stipules deciduous,

adnate to the petiole. Spike narrow, filiform,

pendulous, young ones green, mature ones

yellowish, upto 16.0 cm and 9 12.5 cm,

female spikes usually much smaller, peduncle

glabrous. Bracts in the cf linearly oblong, de-

current, sessile, upper half with free margin,

in $ same as in the male but the upper

portion forms a thin hemispherical cup-like

depression probably due to the presence of the

spherical ovary, outer surface glabrous. Stamens

consistently 2. Carpel single, ovary spherical,

style represented by a mere constriction, stigma

3-5 lobed, lobes elongate, papillate. Berry

spherical 0.5-0. 6 cm in diameter, pungent.

Very common in the forest slopes of the

Western Ghats, found climbing over the sup-

porting trees both by twining and striking

roots at the nodes, ascends to a height of 10 m

or more. Flowers during May- June period,

off season flowering rare. Fruit in ripening

undergoes a colour change from green to red

to black. More than 75 cultivars are known

to be cultivated for their fruits, which are

marketed as the famous ‘black pepper’. The

cultivars of black pepper are quite similar to

the wild ones. The only major difference is

the monoecious nature of cultivars and dio-

ecious nature of the wild vines. However,

hermaphrodite forms of wild vines and female

forms of cultivars are also known.

Distribution : Indonesia, Malaysia, and

Brazil. In India, occurs wild in the forests

of the Western Ghats and the Eastern

Ghats and cultivated in Kerala, Karnataka,

Tamilnadu and Maharashtra. In Karnataka,

both cultivated and wild forms occur in

Dakshina Kannada, Uttara Kannada, Shimoga,

Kodagu, Chikmagalur and Hassan districts.

73

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

Fig. 4. P. nigrum Linn.

A. Branch with fruiting spike; B. Branch with male spike; C. Female spike; D. Male

spike; E. Portion of female spike; F. Portion of male spike; Ga. Stamen; Gb. Anther

lobes after anthesis; Ha. Ovary; Hb. Ovary — L.S.; la. Female bract — side view;

lb. Female bract — bottom view; Ic. Male bract — side view; Id. Male bract —

bottom view; J. Mature berry.

GENUS PIPER LINN. IN KARNATAKA

Type locality. India.

Selected specimens examined. Kanodia

9626, Patil 2921, Puri 1114, Rolla 84917,

Subramanian 70796, 71600, Wadwa 4313,

109793 (BSI); Barber 5466, 5945, 7410, 7414,

8708, 8709, Bourne 369, Naithani 24170 (MH);

Saldanha 11474, 12599, 13274, 14634, 15071,

Ramamoorthy & Gandhi 2655 (CTS); Rahi-

man 3, 4, 5, 6, 19, 22, 36, 38, 42, 43, 73, 75,

76, 85, 86, 95, 111, 112, 114, 116, 117, 122,

130, 132, 148, 200, 210, 211, 239 (MUK).

P. trkhostachyon (Miq.) C.DC, in DC., Prod.

16(1): 242. 1869. J. Hooker, FI. Brit. India

5: 80. 1886; Gamble, FI. Madras 1206.

1925. Muldera trichostachya Miq., London.

J. Bot. 5: 556. 1846. (Fig. 5).

A stout-stemmed climber. Leaves alternate,

coriaceous, entirely glabrous, usually oblong

but variable from ovate to lanceolate, upto

20.0 cm long and 10.0 cm broad, in males

leaves smaller and narrower, 2-3 pairs of pro-

minent lateral ribs, the anteriormost pair

emerges from the midrib simultaneously about

2-3 cm above leaf base, nerves strong beneath,

dorsal side green, ventral, light coloured,

glaucous in appearance. Spike narrow, filiform,

cf upto 9.5 cm, 9 upto 7.0 cm, fruiting

spike upto 10.0 cm, peduncle glabrous except

for a strip of hairy tissue on the upper por-

tion which might represent the decurrent part

of the lowermost bract. Bract decurrent at

the base, upper portion transformed into a

fleshy obconical or hemispherical cup with a

narrow slit like mouth, entire bract puberu-

lous or hirtellous. Stamens consistently 2,

short. Carpel single, ovary obovate, style

absent, stigma 3-4 lobed, lobes short. Berry

spherical or oblong, larger than commercial

black pepper, 0.6-0. 8 cm in diameter.

Climbs to a height of 10 m or more, stem

may grow to a thickness of more than 6.0

cm, covered with thick cork which in old

vines, is longitudinally furrowed. Occurs only in

higher altitude forests (more than 700 m above

sea level). Compared to all the other species,

this one and P. galeatum (Miq.) C.DC., are

hardy species and can withstand slightly dry

climate. These two species are occasionally

encountered along with cultivated Pepper in

some plantations. Flowers in May- June period,

off season flowerings observed. Fruits in ripen-

ing undergo a colour change from green to

yellow to orange to red, slightly less pungent

than commercial black pepper and are some-

times used as adulterant while marketing the

latter.

Distribution : India and Malaya (Malaysia).

In India reported only from the Western

Ghats. In Karnataka, Shimoga, Chikmagalur,

Coorg, southern part of Uttara Kannada and

western part of Hassan districts.

Type locality. Malabar (Kerala).

Selected specimens examined. Cooke 46350

A, Arora 85576, Rao 79977 (BSI); Jacob

537, Narayanaswami 3538, 5407, Subra-

manyam 27703, Vivekanathan 45642, Wight

exciccata sin. num., acc. no 43008, 43009

(MH); Saldanha 12247, 15775, 15798, Nicol-

son, Saldanha & Ramamoorthy 37 (CTS);

Rahiman 62, 105, 107, 121, 138, 144, 147,

155, 162, 163, 178, 187, 191, 196, 201, 202,

203, 210, 213, 233, 234, 235, 237, 246, 266

(MUK).

P. galeatum (Miq.) C.DC., in DC., Prodr.

16(1): 242. 1869; J. Hooker, FI. Brit. India

5: 80. 1886; Gamble, FI. Madras 1206.

1925. P. Talbotii C.DC., in Fedde, Repert.

10: 523. 1912 nomen. Muldera galeata

Miq., London J. Bot. 5: 557. 1846. (Fig. 6).

Similar to P. trichostachyon in all the

characters except for the absence of hairs

on the outer surface of the bracts. Young

spikes are pink in colour in some vines and

green in others.

Several botanists including Miquel (1846),

C. de Candolle (1869), Hooker (1886) and

75

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

Fig. 5. P. trichostachyon C. DC.

A. Branch with fruiting spike; B. Branch with male spike; C. Female spike; D. Male

spike; E. Portion of female spike; F. Portion of male spike; G. Stamens; Ha. Flower

showing stamens; Hb. Flower with stamen — L.S.; la. Ovary; lb. Ovary — L.S.;

J. Bract; K. Mature berry.

GENUS PIPER LINN. IN KARNATAKA

Fig. 6. P. galeatum C. DC.

A. Branch with fruiting spikes; B. Branch with male spikes; C. Female spike; D. Male

spike; E. Portion of female spike; F. Portion of male spike; G. Stamens; Ha. Female

flower; Hb. Ovary — L.S.; I. Bract; J. Mature berry.

77

JOURNAL , BOMBAY NATURAL HIST. SOCIETY, Vol. 84

Gamble (1925) distinguished P. galeatum

from P. trichostachyon by distantly arranged

flowers, presence of sessile bracts and

absence of hairs on the outer surface of the

bracts in contrast to the closely placed flowers,

stipitate bracts and presence of hairs on

bracts. Of these, the first two characters are

of relative nature. While the typical P. galea-

tum from Cardamom hills shows stipitate and

distantly placed flowers, the same species

occurring in Anamalai hills shows almost

sessile and comparatively closely placed

flowers6. P. galeatum from Karnataka region

also shows almost sessile and very closely

placed flowers. The third character, namely

the presence and absence of hairs, on the

other hand, is an absolute character and hence

could alone be taken as the diagnostic charac-

ter to distinguish these two species.

Distribution : India and Java. In India, the

Western Ghats from Bombay to Yellapur of

Karnataka and from Anamalai hills to

Travancore hills. In Karnataka, northern part

of Uttara Kannada district.

Type locality. Peninsular India (Courtallum

of Tamilnadu?)

Selected specimens examined. Talbot 1219

(CNH); Ahuja 31626, 47696, Cooke exciccata

sin. num., acc. no. 12333, 16689, Hemadri

104440, Janardhan 70108, 76614, 81777, Maha-

jan 17176, Puri 12605, Reddi 85946,

95813, 99326, Ryan 1757, Rolla 83427, Talbot

1593, 2888, Vasavada 9320 (BSI); Barber

5426, 5441, 5467, 5483, 5484, 5485, 5486, 7192,

Ramamoorthy 16145, Wight exciccata sin.

num., acc. no 42973 (MH); Rahiman 53, 54,

55, 56, 57, 58, 59, 270, 273, 297 (MUK).

P. mullesua Ham., ex D.Don, Prod. FI. Nep.

20. 1825; C.DC., in DC., Prod. 14(1): 338.

1869. P. brachystachyum Wall., in Wall. Cat.

6656 in part. 1832. P. guigual Ham. ex D.

6 C. de Candolle (1912) unpublished document

available at Botanical Survey of India, Southern

Circle, Coimbatore.

Don, Prod. FI. Nep. 20. 1825. P. vasculosum

Wall., Cat. 6660. 1832. Chavica Guigual

Miq., Syst. Pip. 280. 1843. C. Mullesua Miq.,

l c. 280. 1843. C. spherostachya Miq., l.c. 278.

1843. (Fig. 7).

Slender-branched extensive climber, stem

rarely attains more than 1.5 cm thickness,

branches entirely glabrous, runners many,

characteristically puberulous. Leaves alternate,

coriaceous, elliptic, upto 14.5 cm in length

and upto 5.5 cm in breadth, usually much

smaller in size, tip caudate-acuminate, bent, 2

pairs of prominent lateral ribs of which the

anteriormost one emerges about one-thirds

above the leaf base, the others emerge from

the base, nerves strong beneath, dorsal side

green, ventral dark green. Spike erect, cylin-

dric in male, upto 4.5 cm in length, female

very small, oblong, white, about 0.4 cm in

length and 0.3 cm in breadth, fruiting spike

upto 1 . 2 cm in length and 0.7 cm in breadth.

Bracts orbicular, peltate, pedicelled. Stamens

2, filament short, thick, anther lobes single re-

niform, attached transversely at the tip of the

filament, pollen sacs 2, dehisce by conspicuous

longitudinal cleft at the crest and the wall of

the sacs recurve and form an umbrella like

structure. Carpel single, ovary ellipsoid, style

represented by a constriction, stigma mostly

3-lobed, lobes minute. Berry very small,

spherical-obovate, very pungent, gives a burn-

ing sensation when chewed. Flowers during

April-May period, off season flowering com-

mon. This species occurs only in very high

altitude forests (more than 700 m above MSL).

This species has been commonly treated as

P. brachystachyum Wall, by many of

botanists including Hooker (1886) and Gam-

ble (1925). The specific epithet 'brachysta-

chyum! was given by Wallich in his catalogue

(1832). Later Hooker (1886) published P.

brachystachyum with requisite diagnosis. How-

ever, long before these, i.e. in 1825 itself, Hamil-

78

GENUS PIPER LINN. IN KARNATAKA

Fig. 7. P. mullesua Ham.

A. Branch with fruiting spikes; B. Branch with male spikes; C. Male spike; D. Female

spike; E. Portion of male spike; F. Portion of female spike; Ga. Stamen; Gb. Anther

lobes — top view; Gc. Anther lobes after anthesis — top view; Gd. Anther lobes

after anthesis — side view; Ha. Ovary; Hb. Ovary — L.S.; la. Male bract — bottom

view; lb. Male bract — side view; Ic. Female bract — bottom view; Id. Female

bract — side view; J. Berry.

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

ton validly published this species as F. mullesua

in Don’s prodromus florae nepalensis.

Raizada (1966) corrected this by considering

F. mullesua as the valid name.

Distribution : India, Nepal, Bhutan and

Eastern Islands. In India, the Himalayas from

Simla to Bhutan, Khashi Hills and the Western

Ghats. In Karnataka, Chikmagalur, Kodagu

and Shimoga districts.

Type locality. Nepal.

Selected specimens examined. Wallich 6656

(CNH); Gamble 20631, Subramanian 71165,

Talbot 3193 (BSI); Balakrishnan 129, Barber

1213, 5436, 5437, 5438, 5447, 5447 a, 5447 b,

5448, 5472, 6546, 6547, 7206, 7207, 7208, 7220,

7314, 7578, Beddome exciccata sin. num., acc.

no. 43091, Bharghavan 47443, Bourne 2374,

Deb 31543, Ellis 34727, Gamble 20630, Henry

16421, Hooker exciccata sin. num., acc. no.

70403, 70405, Jacob 16069, 17562, Lawson

exciccata sin. num., acc. no. 43078, 43088,

Narayanswami 3801, 4440, Ramakrishnan

39108, Ramamurthy 18129, Rao 40472, 40489,

Sebastine 2561, 3212, 4466, 18444, 24994,

Sharma 35873, Vajravelu 29196, 35110, 36833,

38213, 38239, 43171, Viswanadhan 807, 975,

Vivekanathan 40614, 40645, Wight exciccata

sin. num., acc. no. 43081, 43082 (MH); Rahi-

man 11, 15, 16, 96, 97, 101, 103, 140, 172, 174.

175, 208, 224 (MUK).

P. longum Linn. Sp. PI. 29. 1753; Roxb., FI.

Ind. 1: 154. 1832; Gamble, FI. Madras

1203. 1925. F. sarmentosum Wall. Cat. 6641.

1832. F. latifolium Hunter, in As. Res. 9:

390. 1809. F. tubinatum Noronha, in Verh.

Batav. Gen. 5: 25, 1790. Chavica Rox-

burghii Miq., Syst. Pip. 230. 1843. C. sar-

mentosa Miq., London J. Bot. 4: 433. 1845;

5: 531 non Syst. Pip. 1846. (Fig. 8).

A slender undershrub, vegetative branches

prostrate, rarely climbing to a short height,

flowering ones erect or subscandent, young

branches puberulous, hair minute but densely

distributed. Leaves on the vegetative branches

upto 13.0 cm long and 7.0 cm broad, cordate,

equilateral with large auricles, leaves on the

fruiting branches cordate to oblong-lanceolate,

with inequilateral lamina, base deeply cordate

with unequal but deep auricle, 3-4 pairs of

prominent lateral ribs, all arise from leaf base,

ventral side pale green, downy or puberulous,

dorsal, light green, glabrous, petiole downy.

Spike cylindric, erect, upto 10.0 cm long

and 0.4 cm broad, $ upto 3.5 cm long and

1 . 0 cm broad, peduncle downy. Bracts peltate,

pedicelled, orbicular, glabrous. Stamens 3-4.

Carpel single, ovary obovate, style represented

by a mere constriction, stigma 3-4 lobed, lobes

short. Berry small, obovate, very pungent,

about 0.2 cm in diameter.

Distribution : India, Sri Lanka and Mala-

yan Islands. In India, east Nepal to Assam,

West Bengal, and west coast of India. In

Karnataka, collected only from Dakshina

Kannada and Shimoga districts. Being an

undershrub which dries and dies in the summer,

it is difficult to locate. Among all the species

in Karnataka, this is the only species which

is found outside the perview of the forests

and woodlands.

Type locality. India?

Selected specimens examined. Ryan sin.

num., Subramanian 70562, 77096, 77144,

Talbot exciccata sin. num., acc. no. 7905,

7906 (BSI). Barber 6078, 6656, 7161, 8728,

Beddome exciccata sin. num., acc. no. 43091,

Bourdillon 449, Deb 30431, 30770, Ellis and

Ramamurthy 18796, Henry 85, 16421, Joseph

17825, Narayanswami 3489, Ramamurthy

18129, 47644, Sebastine 717, 20810, 25369,

4466, 18444, Sharma 42451, Shetty 27381,

Subramanyan 3489, Vajravelu 26277, 33372,

48844, 47443, Wight 886 (MH); Rahiman 1,

77, 78, 215, 222, 267 (MUK).

Several taxonomists described intraspecific

categories such as varieties and forms in a

80

GENUS PIPER LINN. IN KARNATAKA

Fig. 8. P. bn gum Linn.

A. Branch with male spikes; B. Branch with female spikes; C. Runner with a leaf;

D. Male spike; E. Female spike; F. Portion of male spike; G. Portion of female

spike; H. Portion of male spike showing stamens (two of the bract removed); I.

Stamen; J. Ovary — L.S.; Ka. Male bract — bottom view; Kb. Male bract — side

view; Kc. Female bract — bottom view; Kd. Female bract — side view; La. Berry

— side view; Lb. Berry — top view.

81

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

number of species of Piper. Among the Karna-

taka species, only a few such as P. argyro-

phyllum, P. hookeri, P. nigrum and P. mullesua

were found to include subspecific categories.

Hooker (1886) described six varieties under

P. argyrophyllum but gave them just numbers

(1 to 6). Brandis (1906) included a variety

a under P. brachystachyum (P. mullesua) and

a variety /3 under P. hookeri. C. de Candolle

(1912) established a variety mysorensis under

P. nigrum. All these varieties are mainly based

on some minor variations in the length and

breadth of leaves or size variations of spikes

and flower parts. Because of the pre-

sence of extreme variability in the mor-

phological characters in the genus and due

to the absence of sharp taxonomic disconti-

nuities in the characters within the species, it

Refer

*Blume, C. L. (1823) : Catalogus van eenige der

merkwaardigste zoo in-als uit-heemse gewassen, te

vinden in’s Lands Plantentuin te Buitenzorg. Botavia.

Brandis, D. (1906) : Indian Trees. Delhi.

Casparus, C. (1696) : Flora Malabarica sive Horti

Malarici. Lugduni Batavorum.

Cooke, T. (1903) : The Flora of the Presidency

of Bombay. London.

de Candolle, C. ( 1 869) : Piperaceae. In : de Can-

dolle, A.P. (ed.). Prodromus Systematis Regni

Vegetabilis 16(1): 235-471. Germany.

de Candolle, C. (1912): Piperaceae Meebodianae

Herbarii Vratislaviensis. In: Fedde, F. (ed.).

Repertorium Specierum Novarum Regni Vegetabilis

10: 518-523.

Don, D. (1825): Prodromus Florae Nepalensis.

London.

Fedde, F. (1912): Repertorium Specierum Nova-

rum Regni Vegetabilis.

Gamble, J. D. (1925): Flora of the Presidency

of Madras. Calcutta.

Hooker, J. D. (1886) : The Flora of British India.

London.

Howard, R. D. (1973): Notes on the Piperaceae

is not advisable to consider subspecific cate-

gories, unlelss the taxon under study is suffi-

ciently different from the species which it has

to be separated from. In the present study,

none of the species was found to have suffi-

cient variations to merit subdivision into a

distinct variety.

Acknowledgements

We wish to express our sincere thanks to

Dr (Fr) C. J. Saldanha, Director, Centre for

Taxonomic Studies, Bangalore, for his kind

help in the study. This paper is a part of the

Ph.D. thesis (University of Mysore) of the

senior author and he gratefully thanks the

Council of Scientific and Industrial Research,

New Delhi, for financial assistance.

E N CE S

of the Lesser Antilles. J. Arnold Arb. 54(3) : 377-

411.

Hunter, W. (1807): Remarks on the species of

pepper, which are found on Prince of Wale’s Island.

Asia. Res. 9: 383-393.

Index Kewensis Plantarum Phanarogamarum

(1895-1970): 2 Vol. & 14 Suppl. Oxford.

*Kunth, C. S. (1939): Linnaea 13: 651-723.

* Lamarck, J.B.A.P.M. de & Poiret, J.L.M. (1791):

Illustrations des genres.

Lawrence, G.H.M. (1951): Taxonomy of Vascu-

lar Plants. New York.

*Link, J.H.P. (1821): Enumeratio Plantarum Horti

Regni Botanici Berlinensis.

Linnaeus, C. (1753) : Species Plantarum. London.

Miquel, F.A.W. (1843): Systema Piperacearum.

Rottendom.

(1845) : Animadversiones in Pipera-

ceae Herbarii Hookeriani. Lond. J. Bot. 4 : 410-470.

(1846): Annotations in Pipera-

ceae Herbarii Arottiani. ibid. 5: 533.

(1859) : Flora Van Nederlandech

India (Flora Indiae Batavae).

*Presl, K. B. (1844): Botanische Bemerkungen.

82

GENUS PIPER LINN. IN KARNATAKA

Rahim'.v, b. A, Murthy, K. N., Nair, M. K. &

Nayar, i . M. (1979): Distribution, morphology

and ecology of Piper species in Karnataka, India.

J. Plant. Crops 7(2) : 93-100.

, Murthy, K. N. & Nair, M. K. (1981) :

Collection and conservation of Piper nigrum and

related species from Karnataka forests. Indian Cocoa

Areca Spices J. 4(4): 110-112.

& Nair, M. K. (1983): Morpho-

logy, Cytology and Chemical constituents of some

Piper species from India. J. Plant. Crops 11(2) :

72-90.

Raizada, M. B. (1966) : Nomenclatural changes

in Indian Plants. Indian Forester. 92(5 ) : 299-320.

Roxburgh, W. (1832) : Flora Indica. Serampore.

Saldanha, C. J. & Nicolson, D. H. (1976) :

Flora of Hassan District, Karnataka, India. New

Delhi.

Vahl, M. (1804) : Enumeratio Plantarum. Havi-

niae.

Wallich, N. (1832): A numerical list of dried

plants in the East India Company’s Museum

(Wallich’s Catalogue). London.

Wight, R. (1853) : leones Plantarum Indiae

Orientalis. Weinheim.

* Original not seen.

83

POLYCHAETA OF THE PULICAT LAKE (TAMIL NADU)1

S. K. Sunder Raj and P. J. Sanjeeva Raj2

(With four plates)

Polychaete fauna of the Pulicat Lake (Tamil Nadu) consisting of 25 species

belonging 23 genera and 13 families, 20 of which are new records to this lake and

two new to the brackishwaters of India are described, together with their distribution

in the lake, and the taxonomic keys for all of them.

I N TROD U CTIO N

Polychaetes constitute a major component

of the bottom fauna of a lagoon like the Puli-

cat Lake as Raman et al (1975) have shown,

and they contribute an important link in the

food- webs of a brackishwater ecosystem.

Among the major brackishwater bodies of

India, the polychaete fauna of the Gangetic

Delta and of the Chilka Lake have been

thoroughly investigated, but no systematic work

on the polychaetes of the Pulicat Lake, which

is the second largest brackishwater body in

India has been attempted.

The present survey was conducted during

the years 1973-1978. As the Pulicat Lake is a

major brackishwater fishing centre in Tamil

Nadu, a knowledge of the polychaete fauna

of this lake would be helpful from both the

academic as well as from the fisheries points

of view. ^

Topography of the Pulicat Lake. Russel

(1898), Hornell (1910), Chacko et al (1953),

Krishnamurthy & Rao (1970), Joel (1973),

Raman et al (1975), Paul Raj (1976) and

Jhingran (1977) have earlier described the

topography of the Pulicat Lake.

Pulicat Lake (Lat. 13°24' to 13°47'N and

Long. 80° 2' to 80°16'E) covers an average

1 Accepted November 1984.

2 Department of Zoology, Madras Christian Col-

lege, Tambaram, Madras 600 059.

area of about 461 sq. kilometres. The average

depth of the lake is about 1 . 5 metres and the

maximum depth is about 7.0 metres. The lake,

at its southern end, close to Pulicat Town,

opens into the Bay of Bengal by a narrow

mouth (pass). In addition to this, there is also

another seasonal pass near Duggirajapatnam

(northernmost point). In the northern part of

the lake, there are two large islands, Venadu

and Irakkam (Plate 1) and a much smaller

one called Kuruvithittu. On the eastern side,

the Sriharikota Island extends north to south

all along, as a narrow strip of land between

the lake and the Bay of Bengal. After the

establishment of the S.H.A.R. on the Srihari-

kota Island it is connected with Sulurpet

Town on the mainland by a cement road. The

Buckingham Canal runs parallel to the entire

length of the lake and it opens into the lake

here and there.

Hydrology and substrata of the Lake. Hor-

nell (1910), Chacko et al (1953), Michael

(1970), Joel (1973), Srinivasan & Pillay (1972),

Krishnamurthy (1973), Raman et al. (1975)

and Paul Raj (1976) have described the hydro-

logy of the Pulicat Lake.

The interesting feature of the lake is that

during the flood season in November-Decem-

ber, the salinity is extremely low, but during

the summer and post-summer months (April

to September) it is hypersaline. Hornell (1910)

observed fine sandy bottom along the shores

84

J. Bombay nat. Hist Soc. 84 Plate 1

Sunder Raj & Sanjeeva Raj: Polychaeta

1. Pulicat Pass; 2. Karimanal; 3. Dhonirevu; 4. Gunankuppam; 5. Lighthousekup-

pam; 6. Kottaikuppam lock; 7. Koraikuppam; 8. Sattankuppam; 9. Edamani; 10.

Kulathumedu; 11. Moosamani lock; 12. Annamalaicheri; 13. Chunnambukulam; 14.

Arangam; 15. Pulincheri; 16. Zonangipalam; 17. Arambakkam; 18. Irrakkam;

19. Venadu; 20. Berupet; 21. Atakanitippa; 22. Malan; 23. Sriharikotta; 24. Roy-

duruah; 25. Avarivakkam; 26. Tada; 27. Duggirajapatnam.

J. Bombay nat. Hist Soc. 84

Sunder Raj & Sanjeeva Raj: Polychaeta

Plate 2

Zone I: Sand with little admixture of mud.

Zone II: Sand and mud with equal proportions with patches of weeds.

Zone III: Mud.

1. Pulicat Pass; 2. Karimanal; 3. Dhonirevu; 4. Gunankuppam; 5. Lighthouse-

kuppam; 6. Kottaikuppam lock; 7. Koraikuppam; 8. Sattankuppam; 9. Edamani;

10. Kulathumedu; 11. Moosamani lock; 12. Annamalaicheri; 13. Chunnambukulam;

14. Arangam; 15. Pulincheri; 16. Zonangipalam; 17. Arambakkam; 18. Irrakkam;

19. Venadu; 23. Sriharikotta; 25. Avarivakkam; 26. Tada.

POLYCHAETA OF THE PUL1CAT LAKE

of the lake near the lake-mouth (pass).

Krishnamurthy (1971) observed three zones,

based on the nature of the substratum, one

zone characterised by the predominance of

sand in the substratum, with little admixture

of mud, a second zone having sand and mud

in equal proportions with patches of weeds,

and a third zone consisting entirely of mud.

Of these three zones, the first and the second

zones are rich in flora and fauna, and poly-

chaetes particularly are plenty in the second

zone (Plate 2).

Review of previous work on Polychaeta of

India. The earliest work on the Polychaeta of

the Indian region is that of Willey (1905),

describing collections from the Gulf of

Mannar. Later, Potts (1909) studied the Poly-

chaeta of the Indian Ocean. Southern (1921)

and Gravely (1927) described the Polychaeta

occurring in the Krusadai Island, Chilka lake,

Gangetic delta and Cochin backwaters. Rindra

(1927) studied the polychaetes belonging to

the genus Eurythoe from Karachi. Aziz (1938)

studied the polychaetes of Karachi. Brackish-

water polychaetes of Madras were studied by

Panikkar & Aiyar (1937). This was followed

by the contributions of Fauvel (1930, 1930a,

1932 and 1940), which culminated in 1953 in

the publication of his comprehensive account

of the Polychaeta of India in the fauna of

British India series. He described 283 species

from the coasts of India, which includes 47

brackishwater species and 236 marine forms.

Day (1962) reviewed the list of the Polychaeta

in the Western Indian Ocean. Polychaeta from

the south-east coast of India were studied by

Ghosh (1963). Some polychaetous annelids

from the Andaman waters were studied by

Tampi & Rangarajan (1964). Polychaetes from

the Cochin harbour area were studied by

Cheriyan (1966). Polychaeta from Maharashtra

and Goa were studied by Parulekar (1971).

More recently, the polychaetes of the Indian

Ocean were catalogued by Hartman (1974).

Concerning the brackishwater polychaetes

particularly of India, Southern (1921) in his

classical work on the Fauna of the Chilka

Lake described about 20 species of brackish-

water polychaetes. Brackishwater polychaetes

of the Gangetic Delta and Cochin Backwaters

were also described by Southern (1921).

Sewell (1934) described the brackishwater

polychaetes of the salt lakes of Calcutta (Hugli

river). Panikkar & Aiyar (1937) described

the brackishwater polychaetes of the Adyar and

Coovum estuaries near Madras. Alikunhi

(1941, 1942, 1943, 1946, 1947, 1948 and 1951)

studied the interstitial polychaetes of Madras.

Chacko et al <1953) merely listed five

species of Polychaeta from the Pulicat Lake,

namely Lycastis indica Southern, Nereis chilka-

ensis Southern, Marphysa gravelyi Southern,

Lumbriconereis polydesma Southern and Poly-

dora kempi Southern. Krishnamurthy (1963)

described six brackishwater polychaetes from

the Adyar estuary, Madras. Balasubramanyam

(1964), while describing the fauna of the

Vellar estuary (Porto Novo) listed 28 species

of polychaetes. Radhakrishna & Ganapati

(1967) worked out the fauna of the Godavari

estuary describing about 19 polychaete species.

In addition to the above faunistic surveys,

some work has been done on the other aspects

like anatomy, behaviour, reproductive biology,

ecophysiology and biochemistry of some select-

ed polychaetes. Ranganathan (1942) worked

out the anatomy of Glycera embranchiata.

Tampi (1946) worked out the structure of the

eyes and tube-building organs of some selected

polychaetes. Krishnan (1952) studied the

nephridia of Nereidae and also the develop-

ment of Diopatra variabilis. Anatomy and

development of Dasychone cingulata was

studied by Thomas (1955). Physiological

studies on Marphysa gravelyi were conducted

by Krishnamurthy (1962, 1963 and 1968).

85

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

Salinity tolerance and weight regulation in

Lycastis indica were studied by Mary (1965).

Table 1 compares the occurrence of the

brackishwater polychaetes of India and their

distribution.

Taxonomic methods. Southern (1921) has

given more emphasis to parapodial structures

rather than to the appendages of the anterior

end. Fauvel (1953) has given more emphasis

to the head morphology (prostomium, eyes,

tentacles, tentacular cirri, gills, proboscis with

its jaws and denticles). The keys given herein

to identify the families of the polychaetes of

the Pulicat Lake are based mainly on the

head structures, whereas the keys for generic

and specific levels are based on head struc-

tures as well as on parapodial characters.

Laboratory maintenance of polychaetes. A

number of attempts were made to maintain

polychaetes alive in the laboratory at the

Madras Christian College, Tambaram, about

80 km inland from the Pulicat Lake. Most

often the worms survived only for a week or

so. The following method was successfully

evolved subsequently to keep the worms alive

for more than six months, away from their

natural habitat.

Polychaete worms collected from the Puli-

cat Lake were transported by rail and road

in about 4 hours and were brought to Tam-

baram in glass bottles containing the lake

water. A perforated lid was used to allow

ventilation. The lower halves of the glass

bottles were filled with the bottom mud from

the lake and the rest was filled with the lake

water. The worms were seen to be readily

burrowing into the mud at the bottom. After

arriving at the inland lab, the worms, along

with the lake water and mud, were transferr-

ed to glass troughs of five-litre capacity. Two

litres of the lake mud at the bottom and two

litres of the lake water were used in each

glass trough. Dark slushy mud was noticed

to decompose fast, hence shore sand was

found to be ideal. The water in the glass

troughs was constantly aerated, to avoid oxy-

gen depletion. Salinity of the water was main-

tained around 35.01%o by adding distilled

water and sea water as may be required. The

optimum salinity was noted to be 30±5%o.

The dissolved oxygen content in the water was

on the average 3.684 ppm. Temperature of

the water was on an average 26 °C. The worms

were able to live well in varying temperatures,

the optimum temperature being 26dz5°C. In

each such glass trough, about 25 worms were

stocked. Species maintained at the inland lab

were Marphysa gravelyi, Nereis chilkaensis,

Heteromastus similis and Euclymene annan-

dalei. If the number in each trough was raised

above 25, mortality was noticed and this may

be due to overcrowding in a small container.

Stocking density may be 10-12 worms per litre.

Since most of the worms are detritus feeders,

no supplemental food was supplied. They were

presumed to be feeding on the detritus and

on the plankton available in the water. These

worms under such laboratory conditions sur-

vived for more than six months, so that they

could be used for the present work.

Materials and Methods

Polychaetes were collected from nearly 27

stations representing all possible biotopes of

the lake. Fishermen of the Pulicat Lake also

collect polychaetes as bait for angling. Two

methods are in vogue.

(i) Intertidal collection. The body fluids of

crabs like Portunus sp., ZJca sp., by breaking

their appendages are spilt over the sandy shore.

This odour of the crab seems to attract the

worms to come out of their burrows. They

are then gently caught by their heads and

pulled out of their burrows.

(ii) Bottom collection. The villagers use a

86

POLYCHAETA OF THE PUL1CAT LAKE

Table 1

87

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

32. Marphysa gravelyi

33. M. sanguined

34. M. stragulum

35. Lumbriconereis polydesma

36. L. simplex

37. L. pseudo bifilaris

38. L. heteropoda

39. S co l o pi os marsupialis

40. Scoloplos indica

41 . Nerine cirratulus

42. Polydora ciliata

43. P. hornelli

44. P. kempi

45. Sp/o bengalensis

46 . Prionospio krusadensis

47 . P. polybranchiata

48. P. cirrifera

49. Myriochele picta

50. Cossura delta

51. Capitella sp.

52. Heteromastus similis

53. Paraheteromastus tenuis

54 . Mastobranchus indicus

55. Barantolla sculpta

56. Branchiocapitella singularis

57. Euclymene annandalei

58. E. insect a

59. Sternaspis costata

60. Pectinaria crassa

61. Amphicteis gunneri

62. Loirnia medusa

63. P/sfa indica

64. Sabellaria spinulosa

65 . S', pectinata

66 . Laonome indica

67. Potamilla leptochaeta

68. Fabricia spongicola

69. Hydroides norvegica

70. Ficopomatus macrodon

71. Mercierella enigmatica

Table 1 (contd.)

+ — + + + +

+

+

+

+

+

+

88

POLYCHAETA OF THE PULICAT LAKE

special teak plank, like a cricket-bat, made

just for collecting polychaetes. The plank is

about 60 to 75 cm long and about 15 cm

broad, and about 10 cm. in thickness. The top

has a handle, but the bottom is pointed. After

choosing the proper habitat of polychaetes,

they insert this plank deep into the soft ooze

almost up to the handle, incline it outwards

and rotate it in a semi-circle, thus scooping

out the bottom mud. Polychaetes dislodged

from such mud are collected by handpicking.

Polychaetes associated with oyster-shells

were collected by dislodging the shells with a

chisel and hammer. Interstitial polychaetes

were collected by “Corer” as designed by

McIntyre (1968).

Key for Identification

(a) KEY TO IDENTIFY ERRANTIA AND SEDENTARIA

Body vermiform, undivided into two regions; all

segments nearly alike; Free-living ERRANTIA

Body divided into two distinct regions, thorax and

abdomen; usually tubicolous SEDENTARIA

(b) KEY TO IDENTIFY FAMILIES OF ERRANTIA

1 . Elytra present on a limited number of segments

only; the posterior segments carry cirri

Aphroditidae

Elytra absent 2

2. Proboscis armed with four teeth; prostomium

fused with buccal segment; feet uniramous

Pisionidae

Proboscis unarmed 3

3. Tentacles four to five; dorsal and ventral cirri

foliaceous; setae compound Phyllodocidae

Dorsal cirri long and moniliform 4

4. Head with two pairs of eyes; two or three ten-

tacles; palps present or absent Hesionidae

Proboscis with paragnaths 5

5. Proboscis armed with a single pair of toothed

jaws; tentacles two; parapodia biramous

Nereidae

Proboscis armed with two pairs of jaws

tentacles four or more; parapodia

biramous or sesquiramous 6

6. Prostomium conical, ringed with four small ten-

tacles; palps absent Glyceridae

Prostomium distinct and well developed with

tentacles and palps; proboscis complex

Eunicidae

(c) ICEY TO IDENTIFY FAMILIES OF SEDENTARIA

1 . Body clearly divided into regions 2

Body not clearly divided into regions; prosto-

mium without tentacles; palps without suckers;

dorsal and ventral cirri foliaceous; hooded hook-

ed setae Spionidae

2. Prostomium conical without appendages; pro-

boscis unarmed; dorsal and ventral cirri absent.

Capitellidae

Prostomium not conical 3

3. Prostomium rimmed with a cephalic plate; anal

funnel with cirri. No gills Maldanidae

Prostomium trilobed or hidden 4

4. Prostomium trilobed, buccal tentacles long and

retractile into the mouth; three to four pairs of

subulate branchiae inserted on the anterior seg-

ments Ampharaetidae

Prostomium hidden; with or without operculum

5

5. With an operculum; a thoracic membrane; tube

calcareous Serpulidae

Without operculum; no thoracic membrane; tube

membranous Sabellidae

(d) KEY TO IDENTIFY GENERA AND SPECIES

Family 1. Aphroditidae

Eyes four; prostomium bilobed; three tentacles;

dorsal setae stouter than the ventral with biden-

tate tips; sessile; elytra fringed with small

papillae; ventral lamellae conspicuous

Harmothoe ampullifera

Family 2. Pisionidae

Presence of two non-serrated buccal spines bet-

ween the two palps with genital papillae in the

35th segment Pisione complexa

Absence of buccal spines and palps longer than

dorsal cirri of the buccal parapodia

Pisionidens indica

Family 3. Phyllodocidae

Prostomium with two pairs of tentacles and two

pairs of tentacular cirri; proboscis with soft rows

of papillae Eteone barcmtollae

89

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

Family 4. Hesionidae

Prostomium with two tentacles; palps absent;

proboscis unarmed; paired brown spots on each

inter segmental line Hesione intertexta

Family 5. Nereidae

1 . Feet uniramous; eyes arranged in a line; dorsal

setae absent Lycastis indica

Feet biramous 2

2. Paragnaths present; dorsal cirrus longer and

larger than ventral cirrus; no dorsal homogomph

falcigerous bristles in posterior feet

Nereis chilkaensis

Paragnaths absent; ventral setigerous lobe bilob-

ed in few segments; jaws with 12 teeth

Ty loner eis fauvel i

Family 6. Eunicidae

1. Tentacular cirri present; tentacles with cirrato-

phores Diopatra neapolitana

Tentacular cirri absent 2

2. Gills present, pectinate; comb setae arranged in

middle region of the body Marphysa gravelyi

Gills absent 3

3. Gills and eyes absent; feet with wing capillary

setae and hooks absent. . . Lumbriconereis simplex

Hooks present Lumbriconereis polydesma

Family 7. Glyceridae

Gills inserted on the dorsal edge of the foot;

proboscis with four long jaws; Dorsal setae

simple capillary and ventral setae compound and

winged; posterior lobes unequal Glyceraalba

Family 8. Spionidae

Prostomium conical; bidentate; hooded hooks;

gills in anterior segments; anal cup present ....

Nerine cirratulus

Prostomium rounded; gills pinnate; hooded hooks

with four teeth; median anal cirrus present ....

Prionospio krusadensis

Prostomium rounded, but slightly notched in

front and prolonged up to the 3rd segment . . .

Poly dor a ciliata

Family 9. Capitellidae

1 . Thorax with seven segments; dorsal and ventral

hooks begin from tenth segment; gills present in

posterior segments Branchiocapitella

singularis

Thorax with more than seven segments 2

2. Thorax with 11 segments; segments one to five

capillary setae; segments six to 11 long hooks;

short hooks in the rest ..... Heteromastus similis

Thorax with 12 segments; segments two to seven

capillary setae; eight to 12 long crochets; short

crochets in the rest Barantolla sculpta

Family 10. M aldan idae

Head with cephalic plate; anal segment with

anal cirri; total segments 19. Ocelli present in

cephalic plate; median ventral cirrus in caudal

funnel stouter than others

Euclymene annandalei

Absence of ocelli in cephalic plate; median ven-

tral cirrus in caudal funnel longer than others

Euclymene insecta

Family 11. Ampharetidae

Thorax 17 segments; gills four pairs and arrang-

ed on either side of the first two segments

Amphecteis gunner i

Family 12. Sabellidae

Thorax six segments; no pickaxe-shaped setae;

ranchiae four pairs Laonome indica

Thorax seven segments; presence of pickaxe-

shaped setae; Branchiae six pairs

Potamilla leptochaeta

Family 13. Serpulidae

Operculum compound; funnel shaped with a

crown of horny spines; radii of operculum sharp

with more than one pair of lateral processes

Hydroides norvegica

Taxonomy

ERRANTIA

Family Aphroditidae

Species 1. Harmothoe ampullifera (Grube) 1878

(Plate 3, figs, la & lb)

Polynoe ampullifera Grube 1878, p. 35; Lepido-

notus ampullifera Gravier 1901, p. 214; Harmothoe

ampullifera Fauvel 1911, p. 368; Paralepidonotus

ampullifera Horst 1917, p. 76; Harmothoe ampulli-

fera Fauvel 1927, p. 414; 1930a, p. 8; 1930b, p. 508;

1932, p. 22; 1940, p. 254; 1953, p. 43-44; De Silva

1965, p. 538-539; Pillai 1965, p. 117-119. Parulekar

1971, p. 732.

90

j. Bombay nat. Hist Soc. 84

Sunder Raj & Sanjeeva Raj: Polychaeta

Plate 3

{see captions overleaf)

Captions to Plate 3

Harmathoe ampullifera Grube, 1878

Fig. la — Median foot;

Fig. lb — Ventral seta.

Pisionidens indica Aiyar & Alikunhi. 1940

Fig. 2 — A typical parapodium.

Pisione complexa Alikunhi, 1947

Fig. 3 — Anterior end, Dorsal view.

Eteone barantollae Fauvel, 1932.

Fig. 4a — Head, Dorsal view;

Fig. 4b — Proboscis with rows of papillae.

Hesione inter texta Grube, 1878

Fig. 5 — Median parapodium.

Lycastis indica Southern, 1921

Fig. 6a — Heterogomph falciger from median parapodium;

Fig. 6b — Homogomph spiniger from same parapodium.

Nereis chilkaensis Southern, 1921.

Fig. 7 — Median right foot.

Tylonereis fauveli Southern, 1921

Fig. 8a — 5th right foot;

Fig. 8b — posterior end, Dorsal view.

Marphysa gravely i Southern, 1921

Fig. 9 — Branchiate foot.

Diopatra neapolitana Delle Chiaje, 1941

Fig. 10 — 10th right foot with gills.

Lumbriconereis simplex Southern, 1921

Fig. 11a — Anterior foot;

Fig. lib — Anterior end, Dorsal view.

Lumbriconereis polydesma Southern, 1921

Fig. 12a — Anterior end, Dorsal view;

Fig. 12b — Tip of crochet from median parapodium.

Glycera alba Rathke, 1843

Fig. 13a — 8th right foot;

Fig. 13b — jaw enlarged.

Nerine cirratulus Delle Chiaje, 1828.

Fig. 14a — Anterior end. Dorsal view;

Fig. 14b — Ventral bidentate hooded hook.

POLY CH A ETA OF THE PULICAT LAKE

Habitat. Collected from within the crevices

of oyster shells.

Description. Length 25 to 35 mm, including

parapodia on either side. Elytra 15 pairs,

overlapping each other posteriorly. Total num-

ber of parapodia 37 pairs. Dorsal setae

slightly curved and serrated. Ventral setae

long and bidentate. Tentacles black in colour.

Occurrence in Indian waters. Madras, Rame-

swaram and Pam ban (coral reefs).

Distribution outside Indian waters. Philippine

Islands, Persian Gulf and Red Sea.

Remarks. This species is closely related to

Harmothoe imbricata, but it differs from that

in having elongated nephridial papillae and

ventral lamellae.

Family Pisionidae

Species 2. Pisionidens indica Aiyar & Alikunhi

1940

(Plate 3, fig. 2)

Pisionella indica Aiyar & Alikunhi 1940, p. 89;

Fauviella pulchra Tebble 1953, p. 938; Pisionidens

pulchra Day 1957, p. 68; Pisionidens indica Day 1962,

p. 636.

Habitat Pulicat Mouth (Interstitial).

Description. 7 to 12 mm in preserved con-

dition. Body consists of about 35 segments.

Prostomium highly reduced. Eyes not clearly

visible. Parapodia uniramous.

Occurrence in Indian waters. Indian Ocean

and Bay of Bengal.

Distribution outside Indian waters. Durban

and West Indian Ocean.

Remarks. The forms described here are not

fully grown. In fully grown forms the third to

sixth parapodia are non-setigerous, setae pre-

sent from the seventh segment only. However

in this juvenile, almost all the parapodia

bear compound as well as simple setae.

Species 3. Pisione complexa Alikunhi 1947

(Plate 3, fig. 3)

Pisione complexa Alikunhi 1947, p. 105; Rao &

Ganapathi 1968, p. 110.

Habitat. Collected from Pulicat Pass (Inter-

stitial).

Description. 6 to 10 mm long, possessing

about 40 segments. Two non-serrated buccal

spines present between the two palps. A pair

of eyes present. Parapodia uniramous. Each

parapodium bears two long acicula and five

setae, both of simple and compound types.

The anal segment bears a pair of long anal

cirri. Genital papillae are seen in the 35th

segment.

Occurrence in Indian waters. Madras beach,

Waltair coast.

Distribution outside Indian waters. Bay of

Bengal.

Remarks. Out of the 15 worms collected

only 4 worms possess anal cirri. In all the

others, the anal cirri are absent.

Family Pliyllodocidae

Species 4. Eteone barantollae Fauvel 1932

(Plate 3, figs. 4a&4b)

Eteone barantollae Fauvel 1932, p. 72; 1953, p. 127.

Habitat. Collected from Pulicat Pass area,

along with Pisione complexa and some nema-

todes.

Description. Length 10 to 15 mm; breadth

1 to 1.5 mm. The worm possesses about 90

segments. A pair of eyes present. Proboscis

bears rows of papillae. Simple setae absent.

Paired anal cirri present.

Occurrence in Indian waters. Saltwater lakes

near Calcutta.

Distribution outside Indian waters. Not

known.

Remarks. Only two worms were collected

by Fauvel (1932 & 1953) who described that

there are five rows of papillae on the proboscis.

91

7

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

But in the present collection, the papillae are

not arranged in five definite rows, but are seen

to be rather irregularly arranged.

Family Hesionidae

Species 5. Hesione intertexta Grube 1878

(Plate 3, fig. 5)

Hesione intertexta Grube 1878, p. 102; Chamber-

lin 1919, p. 188; Monro 1926; p. 311; Pruvot 1930,

p. 29; Fauvel 1932, p. 60; Monro 1937, p. 270;

Fauvel 1953, p. 105; Tampi 1964. p. 104.

Habitat. Collected from the crevices of

oyster shells.

Description. Prostomium bilobed. Two pairs

of eyes present. A pair of tentacles and eight

pairs of tentacular cirri present with cirrato-

phores. Parapodia uniramous. Setae sickle-

shaped. Total segments 16. Proboscis unarmed.

Segments distinct at the sides.

Occurrence in Indian waters. Gulf of

Mannar, Port Blair, West Indian Ocean.

Distribution outside Indian waters. New

Caledonia, Philippine Islands, Australia.

Remarks. These worms were collected along

with Harmathoe ampullifera.

Family Nereidae

Species 6. Lycastis indica Southern 1921

(Plate 3, figs. 6a & 6b)

Lycastis indica Southern 1921, p. 578; Horst 1924,

p. 4; Fauvel 1930a, p. 19; 1932, p. 82; Aziz 1938,

p. 27; Fauvel 1940, 'p. 257; 1953, p. 167; Ghosh

1963, p. 240; De Silva 1965, p. 5.

Habitat. This species is an euryhaline form.

It was collected from Sattankuppam, Light-

house laippam and Edamani on the Pulicat

Lake.

Description. Length 30 to 40 mm and con-

sists of about 130 setigerous segments. Pro-

boscis without paragnaths. Jaws possess nine

teeth. Feet uniramous. In larger specimens, the

dorsal cirri are long and finger-shaped. The

dorsal cirri consist of a few homogomph

spinigerous setae. The ventral tuft bears both

heterogomph, spinigerous and heterogomph

falcigerous setae.

Occurrence in Indian waters. Chilka Lake,

Salt lakes (Calcutta), Cochin, Madras, Tra-

vancore, Kilakarai, Waltair and Porto-Novo.

Distribution outside Indian waters. Not

known.

Remarks. Southern (1921) describes the

dorsal longitudinal groove ending in a pit, but

the Pulicat species do not show this pit.

Species 7. Nereis chilkaensis Southern 1921

(Plate 3, fig. 7)

Nereis chilkaensis Southern 1921, p. 584; Fauvel

1932, p. 94; Panikkar & Aiyar 1937, p. 293; Fauvel

1940, p. 258; 1953, p. 185; De Silva 1965, p. 543;

Parulekar 1971, p. 739.

Habitat. This species is available in almost

all regions of the Pulicat Lake except at the

pass. It lives in burrows and occasionally

comes out to the surface of the water.

Description. Length 50-75 mm and consists

of about 75 segments. Characters are very

similar to those of Southern’s (1921) descrip-

tion. Feet biramous. Each foot bears about

40-50 setae in general. Variations may occur

due to the varying degrees of sexual maturity.

Occurrence in Indian waters. Chilka Lake,

Ennur backwater, Pamban, Madras coast and

T ravancore.

Distribution outside Indian waters. Sri Lanka.

Remarks. The colour of the worm varies in

different areas of the lake.

Species 8. Tylonereis fauveli Southern 1921

(Plate 3, figs. 8a & 8b)

Tylonereis fauveli Southern 1921, p. 582; Fauvel

1930a, p. 19; 1932, p. 84; 1953, p. 169.

Habitat. It was collected at the following

stations : Venadu, Berupet, Sriharikota, Ataka-

nitippa and Royduruah.

92

POLYCHAETA OF THE PULICAT LAKE

Description. Length 40-80 mm and consists

of about 125 segments. Eyes black. Tentacles,

palps and tentacular cirri well developed. Feet

biramous, setae arranged in three groups. Anal

segment bright red in colour.

Occurrence in Indian waters. Chilka Lake

and Pamban.

Distribution outside Indian waters. Mergui.

Remarks. This species has more resemblance

to Tylonereis bogoyawlonskyi Fauvel (1911),

but it differs from the latter in having a

bilobed ventral setigerous neuropodium, instead

of a trilobed one. The worms are pale pink

in colour.

Family Eunicidae

Species 9. Marphysa gravelyi Southern 1921

(Plate 3, fig. 9)

Marphysa gravelyi Southern 1921. p. 617; Gra-

velyi 1927, p. 19; Fauvel 1932, p. 142; Aiyar 1933,

p. 207; Fauvel 1953, p. 246; Krishnamurthy 1963,

p. 97; Pillai 1965, p. 110-177; Cheriyan 1966, p. 44.

Habitat. These worms live in long burrows

in muddy bottom of the Pulicat Lake. Their

burrows are not vertical but pass through the

soil irregularly in various directions. The open-

ings of their burrows can be easily located

by the presence of a circular ridge of fine sand

all round. During breeding season one can

notice the large cucumber-shaped egg mass of

jelly, attached to the mouth of each burrow.

These worms were collected from the stations

Sattankuppam, Lighthousekuppam, Edamani,

Kottaikuppam lock, Avarivakkam, Dhonirevu,

Annamalaicheri, Moosamani lock, Arangam,

Chunnambukulam, Arambakkam, Irakkam,

Malan, Dugirajapatnam and Royduruah.

Description. Large specimens measure about

300 mm and carry about 500 segments. Ante-

rior end slightly cylindrical up to the 7th seg-

ment behind which it is depressed. Anterior

region greenish in colour, posterior region

blood-red in colour. Gills vary from blood-red

to pale yellow. Out of the five tentacles the

middle one is the longest. In general, two eyes

present in young individuals. Two pairs of

anal cirri, of which one pair larger than the

other. Dental apparatus similar to Southern’s

(1921) description. Feet highly vascularised;

setae arranged in two groups; capillary type.

Their length varies very much. The blades

very minutely serrated.

Occurrence in Indian waters. Chilka Lake,

Adyar estuary, Ennur backwaters, Vellar

estuary and Cochin harbour area.

Distribution outside Indian waters. Philip-

pines and Indonesia.

Remarks. Eyes are usually said to be absent

in adults, but they are well developed in the

adults of Chunnambukulam and Pulincheri

stations. However, adults from Edamani,

Lighthousekuppam and Sattankuppam do not

possess eyes. This is an instance of intraspecific

variation.

Species 10. Diopatra neapolitana Delle Chiaje

1841

(Plate 3, fig. 10)

Diopatra neapolitana Delle Chiaje, 1841; Me In-

tosh 1903, p. 128; Crossland 1903, p. 132; Diopatra

amboensis Willey 1905, p. 214;Diopatra variabilis

Southern 1921. p. 611; Diopatra neapolitana Fauvel

1923, p. 419; 1930, p. 29; 1932, p. 144; 1933, p. 28;

Monro 1933, p. 293; Aziz 1938, p. 39; Fauvel 1953,

p. 252; Tebble 1955, p. 116; Pillai 1961, p. 13;

Cheriyan 1966, p. 45; Diopatra variabilis Hartman

1974, p. 223.

Habitat. It was collected at the Pulicat Pass,

Karimanal, Kottaikuppam lock, Kulathumedu

and Pakkam.

Description. Purple-green in colour. Up to

250 mm and consists of about 320 segments.

Out of the five occipital tentacles the median

one can be stretched up to the middle of the

10th segment. All tentacles bear cirratophores.

Eyes absent. Gills begin from the fourth foot

only. Gill filaments spirally arranged. There is

93

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

no variation in the dental apparatus from

Southern’s description. The tube is membra-

nous and partly buried in sand, the upperpart

thick, tough and more or less encrusted with

broken pieces of molluscan shells.

Occurrence in Indian waters. Gangetic delta,

Orissa coast, Madras coast. Gulf of Mannar

and Cochin harbour area.

Distribution outside Indian waters. Pacific

Ocean, China Sea, Gulf of Siam, Arabian Sea,

Gulf of Oman, Persian Gulf, Red Sea, Atlantic

Ocean and the Mediterranean Sea.

Remarks. According to Fauvel (1953), there

is not much difference between Diopatra

neapolitana Delle Chiaje 1841, and Diopatra

variabilis Southern 1921 except for the teeth

in comb setae.

Species 11. Lumbriconereis simplex Southern

1921

(Plate 3, figs. 11a & lib)

Lumbriconereis simplex Southern 1921, p. 625;

Fauvel 1953, p. 264; Cheriyan 1966, p. 46.

Habitat. Available at Dhonirevu only and

is collected along with Euclymene annandalei.

Description. Prostomium conical. Eyes, palps

and tentacles absent. Feet highly vascularised.

Parapodia absent in the first two segments.

Setae simple winged, capillary type. There is

not much variation in the pharyngeal complex

from Southern’s (1921) description. Length

15 to 30 mm., breadth about 2 mm.

Occurrence in Indian waters. Saltwaters of

Calcutta, Chilka Lake and Cochin harbour.

Distribution outside Indian waters. Not

known.

Remarks. According to Southern (1921),

there is a dark amber coloured spot near the

base of each foot. Such spots are not observed

in the Pulicat forms.

Species 12. Lumbriconereis polydesma Southern

1921

(Plate 3, figs. 12a & 12b)

Lumbriconereis polydesma Southern 1921, p. 632;

Panikkar and Aiyar 1937; Fauvel 1953, p. 264.

Habitat. Available only at Dhonirevu.

Description. Prostomium round; eyes absent.

Two achaetous rings between the head and 1st

setigerous segment. Feet increase in size up to

the 10th segment and then taper gradually

towards the posterior end. Flooks present from

the 29th feet. Flooks or crochets have two

broad wings over the tip with delicate stria-

tions. The setae are stout winged capillaries.

Occurrence in Indian waters. Chilka Lake.

Distribution outside Indian waters. Not

known.

Remarks. According to Southern (1921),

there is a small patch of pigment near the

posterior border of the head, a little on the

right side of the median line. Such a pigment

patch is not observed in the Pulicat forms.

Family Glyceridae

Species 13. Glycera alba Rathke 1843

(Plate 3, figs. 13a & 13b)

Glycera alba Ehlers 1868, p. 661; Moore 1903,

p. 464; Izuka 1912, p. 247; var. cochinensis Southern

1921, p. 627; Fauvel 1923, p. 385; Gravely 1927,

p. 9; Fauvel 1932, p. 126; Aziz 1938, p. 31; Fauvel

1940, p. 261; 1953, p. 292; Cheriyan' 1966, p. 47;

Parulekar 1971, p. 745.

Habitat. This form is confined to Pulicat

Pass (sandy beach) only.

Description. Length 50 to 60 mm about 100

setigerous segments. Body rounded, prosto-

mium conical and pointed with four small ten-

tacles. Proboscis with four hooked jaws and

well developed papillae. Gills simple and in-

serted on the dorsal side of the parapodia.

Anal segment bears a pair of long tapering

cirri. Dorsal setae simple, ventral setae com-

pound.

Occurrence in Indian waters. Gan jam coast.

Cochin backwater and Mormugoa Bay.

94

j. Bombay nat. Hist Soc. 84

Sunder Raj & Sanjeeva Raj: Polychaeta

Plate 4

( see captions overleaf)

Captions to Plate 4

Prionospio krusadensis Fauvel, 1929

Fig. 15 — Median parapodium.

Polydora ciliata Johnston, 1838

Fig. 16 — Anterior end, Dorsal view.

Heteromastus similis Southern, 1921

Fig. 17 — Anterior end, side view.

Barantolla sculpta Southern, 1921

Fig. 18 — Dorsal view of segments with branchiae.

Branchiocapitella singularis Fauvel, 1932

Fig. 19 — Dorsal view of segments with branchial lobes.

Euclymene annandalei Southern, 1921

Fig. 20a — Anterior end, lateral view;

Fig. 20a — Uncinus of the 10th segment.

Clymene ( Euclymene ) insecta Ehlers, 1904

Fig. 21a — Anterior end, side view;

Fig. 21b — Acicular hook.

Amphicteis gunneri Sars, 1851

Fig. 22a — Thoracic foot with dorsal cirrus and pinnules;

Fig. 22b — Thoracic uncinus.

Laonome indica Southern, 1921

Fig. 23a — Anterior end, Dorsal view;

Fig. 23b — Thoracic uncinus.

Potamilla leptochaeta Southern, 1921

Fig. 24a — Tip of a gill;

Fig. 24b — Avicular uncinus.

Hydroides norvegica Gunnereus, 1768

Fig. 25a — A single radiole enlarged;

Fig. 25b — Baynet shaped bristle.

POLYCHAETA OF THE PULICAT LAKE

Distribution outside Indian waters. Red Sea,

Atlantic Ocean and Indian Ocean.

Remarks. The worms are milky white in

colour.

SEDENTARI A

Family Spionidae

Species 14. Nerine cirratulus Delle Chiaje 1828

(Plate 3, figs. 14a & 14b)

Nerine cirratulus Fauvel 1927a, p. 36; 1953, p.

312; Day 1962, p. 648; De Silva 1965, p. 553.

Habitat. These forms are available only on

the sandy shores of the Kottaikuppam lock

but completely absent in the muddy areas.

Description. Worms bright red up to the

anterior 30 segments, rest bluish green in

colour. Length 30-50 mm. Prostomium with

two long tentacle-like palps. A single occipital

tentacle-like keel. Four large eyes arranged in

a single transverse row. Gills begin from the

second setigerous segment, absent in the few

posterior segments. Dorsal lamellae long in the

anterior region but short in the posterior re-

gion. Ventral lamellae narrow. Feet biramous

and possess winged capillary setae. The hood-

ed hooks begin from the 30th segment. The

tip of the hooks is bidentate. Their number

varies in various segments.

Occurrence in Indian waters. Vishakapatnam

channel, Sri Lanka, Indian Ocean.

Distribution outside Indian waters. Atlantic

Ocean, Mediterranean Sea and Madagascar.

Remarks. This form differs from Fauvel’s

(1953) description in the following characters.

1 . The colour of the Pulicat form is red

up to the first 30 segments and the rest

of the segments are bluish green in colour,

but the whole worm is bluish green in

colour according to Fauvel.

2. Four eyes are arranged in a transverse row

in the Pulicat forms. They are arranged in

a trapezium according to Fauvel’s descrip-

tion.

3 . The posterior occipital peak is not well

developed in the Pulicat forms. According

to FauvePs description, this reaches upto

the second segment.

Species 15. Prionospio krusadensis Fauvel 1929

(Plate 4, fig. 15)

Prionospio krusadensis Fauvel 1929, p. 182; 1930,

p. 38; 1953; p. 326.

Habitat. This form is available only in the

sandy shores of the Pulicat Pass.

Description. Body slender, anterior region

slightly enlarged. Prostomium long and round-

ed. Four small eyes arranged in a trapezium.

Branchiae arise from the second setigerous

segment onwards. Gills large and oval in shape.

Setae are capillary. Ventral hooks begin from

the 18th segment onwards. Dorsal hooks be-

gin from the 40th foot. Hooks bear three teeth.

Median anal cirrus present. Length 30 to 50

mm. Breadth about 2 mm.

Occurrence in Indian waters. Gulf of

Mannar, Krusadai Island.

Distribution outside Indian waters. Not

known.

Remarks. These forms are milky white in

colour but turn yellow in alcohol.

Species 16. Polydora ciliata Johnston 1838

(Plate 4, fig. 16)

Polydora ciliata Fauvel 1927a, p. 49; Panikkar

and Aiyar 1937; p. 293; Fauvel 1953, p. 319.

Habitat. This species was collected from the

oyster bed area, along with serpulids and

sabellids.

Description. Prostomium slightly notched in

front and prolonged backwards upto the 3rd

segment. Four small eye-spots present on dor-

sal side. No dorsal setae on the 1st setigerous

segment. Ventral capillary setae well develop-

ed. Ventral bidentate hooks present on the

95

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

7th setigerous segment. Gills from the 7th

segment onwards.

Occurrence in Indian waters. Chandipore,

Orissa coast and Mangalore coast.

Distribution outside Indian waters. Austra-

lia, Indo-China, Red Sea, Atlantic Ocean,

Mediterranean Sea and Falkland Islands.

Remarks. Though they are small in size

(25mm /1mm), they are capable of producing

mud-blisters in the mantle of the edible oyster,

Crassostrea madrasensis (Stephen, 1978b).

They are considered as pests of bivalves. Their

burrow is U-shaped. Although another closely

related species Poly dor a kempi was recorded

by Chacko et al. (1953) in Pulicat Lake, dur-

ing the course of the present investigation over

a period of seven years it was not possible

to record this species.

Family Capitellidae

Species 17. Heteromastus similis Southern 1921

(Plate 4, fig. 17)

Heteromastus similis Southern 1921, p. 640; Grave-

ly 1927, p. 26; Fauvel 1930a, p. 46; 1932, p. 195;

1953, p. 366; De Silva 1965, p. 554.

Habitat. This is widespread in Pulicat Lake.

Description. Bright red in colour and con-

sist of about 200 to 220 segments. Length 60

to 70 mm. The anterior end enlarged and

gradually tapers towards the tail. Peristomium

long and achaetous. Thorax of 11 segments.

Segment two to the sixth possess short capillary

setae. Gills absent. Anal segment bears a

slender clavate tail. Posterior segments possess

dorsal hooks in each foot. The hook has two

large teeth and one small one.

Occurrence in Indian waters. Chilka Lake,

Vishakapatnam and Gulf of Mannar.

Distribution outside Indian waters. Teleh-

Sap, Gulf of Siam.

Remarks. These are very closely related to

Heteromastus filiformis Claperede (1864). The

head as well as the arrangement and structure

of the setae are almost identical. In H. fili-

formis, the anterior abdominal segments are

much longer than the thoracic segments,

whereas in H. similis the segments do not

differ in length.

Species 18. Barantolla scuSpta Southern 1921

(Plate 4, fig. 18)

Barantolla sculpta Southern 1921, p. 643; Fauvel

1932, p. 196; 1953, p. 370.

Habitat. These forms were collected from

Gunankuppam along with Heteromastus simi-

lis from the dark mud.

Description. Reddish brown in colour, about

50 mm long by 2 mm. Total number of seg-

ments about 60. Body wider near fourth or fifth

segment. Eyes absent. First four segments slight-

ly tassellated. Segments two to seven have only

capillary setae. Segments eight to twelve possess

long crochets. These crochets are short in the

abdominal segments. Branchiae arise from the

56th segment onwards. They lie under the dor-

sal parapodial lobes. Number of branchiae is

more towards the anal end. Anal cirrus single

and median. 12 thoracic segments.

Occurrence in Indian waters. Barantolla

near Calcutta.

Distribution outside Indian waters. Taleh-

Sap, Gulf of Siam.

Remarks. The head is contracted and with-

drawn under the peristomium. According to

Southern (1921), there are 12 thoracic seg-

ments, but in Pulicat forms they are not very

distinct from the abdominal segments.

Species 19. Branchiocapitella singulars Fauvel

1932

(Plate 4, fig. 19)

Branchiocapitella singularis Fauvel 1932, p. 197;

1953, p. 371; De Silva 1965, p. 555; Pillai 1965,

p. 1 10-177.

Habitat. These worms are available only in

96

POLYCHAETA OF THE PUL1CAT LAKE

the dark sand near the Lighthousekuppam area.

Description. Total length 50 to 60 mm/

2 mm. Body slender and slightly enlarged in

the thorax. Seven thoracic segments. Dorsal

and ventral hooks start from the tenth segment

onwards. Gills present nearer the anal seg-

ments only. Pygidium ends in a bilobed knob.

Occurrence in Indian waters. Porto-Novo

and Vishakapatnam.

Distribution outside Indian waters. Sri

Lanka.

Remarks. The copulatory spines described

by Fauvel (1953) are not seen on the eighth

and ninth segments.

Family Maldanidae

Species 20. Eudymene annandalei Southern

1921

(Plate 4, figs. 20a & 20b)

Eudymene annandalei Southern 1921, p. 648;

Fauvel 1932, p. 199; 1953, p. 377.

Habitat. They live in sandy tubes and from

the dark muddy areas of the Pulicat Lake,

namely from Sattankuppam, Lighthousekup-

pam, Avarivakkam, Dhonirevu and Moosa-

mani Lock.

Description. Length 20 to 60 mm. Total

segments 21, of which two achaetous. Head

carries the cephalic plate. Numerous ocelli.

Caudal funnel bears short, bluntly rounded

cirri. Parapodia very close to Southern’s

description.

Occurrence in Indian waters. Vellar Estuary,

Kakinada Bay and Chilka Lake. Camorta,

Andaman and Nicobar Islands,

Distribution outside Indian waters. Amoy

(China).

Remarks. The sandy tubes are not as brittle

as Southern (1921) has stated. The segmenta-

tion is very distinct.

Species 21. Clymene (Eudymene) insecta

(Ehlers) 1904

(Plate 4, figs. 21a & 21b)

Clymenella insecta Ehlers 1904, p. 54; Paraxillella

insecta Augener 1926a, p. 192; Clymene ( Eucly -

mene) insecta Fauvel 1932, p. 199; 1953, p. 377;

Krishnamurthy 1963, p. 95.

Habitat. This also lives in sandy tubes. This

is available only from the Chunnambukulam

area.

Description. Bright red in colour. Body deli-

cate, total number of segments about 22.

Nuchal grooves not clearly seen. Prostomium

does not bear any appendages other than the

cephalic plate. The caudal funnel possesses a

number of short cirri, of which one is longer

than the others. The first three setigerous

segments possess acicular hooks. Fourth to

seventh segments bear hooded hooks with two

to three teeth. Capillary and capillary wing

setae present from the eighth segment onwards.

Occurrence in Indian waters. Vishakapatnam

and Madras.

Distribution outside Indian waters. New

Zealand.

Remarks. Caudal funnel is similar to that

of Euclymene annandalei , but it differs in hav-

ing a longer median ventral cirrus. In E.

annandalei, the median ventral cirrus is slight-

ly stouter than in others.

Family Ampharetidae

Species 22. Amphicteis gunneri Sars 1835

(Plate 4, figs. 22a & 22b)

Amphicteis gunned Malmgren 1865, p. 365;

Amphicteis japonica McIntosh 1885, p. 431; Amphi-

cteis gunned Fauvel 1897, p. 411; Hessle 1917, p.

116; 1927, p. 231; 1932, p. 216; Monro 1933, p. 313;

Fauvel 1953, p. 407; Day 1967, p. 695.

Habitat. These forms were collected from the

crevices of oyster shells.

Description. Body divided into thorax and

abdomen. Thorax bears 17 segments and

abdomen bears about 15 to 25 segments. Eyes

or eye-spots completely absent. Thorax pos-

sesses dorsal capillary setae and ventral un-

cinigerous pinnules. Abdomen bears only

97

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 84

uncinigerous pinnules. Four pairs of gills pre-

sent. Anal segment does not bear anal cirri.

Tube is muddy and the length varies from 40

to 60 mm.

Occurrence in Indian waters. Bay of Bengal,

Orissa coast.

Distribution outside Indian waters. Gulf of

Oman, Japan, Indo-China, Atlantic Ocean,

Mediterranean Sea and Antarctic Ocean.

Remarks. The worms are pink in colour.

According to Fauvel (1953), there are nume-

rous eye-spots. Pulicat forms do not possess

any eye-spots.

Family Sabellidae

Species 23. Laonome indica Southern 1921

(Plate 4, figs. 23a & 23b)

Laonome indica Southern 1921, p. 652; Fauvel

1953, p. 446.

Habitat. This was collected from the Kula-

thumedu station, from within the crevices of

oyster shells.

Description. Very thin, about 50 segments.

Posterior ten segments shorter than anterior

ones. Only four pairs of branchial lobes in-

stead of seven as Southern (1921) describes,

but they are symmetrical. Eyes completely

absent. Thorax composed of about six seg-

ments. The abdominal segments possess uncini.

The arrangement of setae agrees with

Southern’s (1921) description.

Occurrence in Indian waters. Chilka Lake.

Distribution outside Indian waters. Not

known.

Remarks. The colour is brown in live con-

dition, but it turns pale when preserved in 5%

formalin.

Species 24. Potamilla leptochaeta Southern

1921

( Plate 4, figs. 24a & 24b)

Potamilla leptochaeta Southern 1921, p. 651;

Gravely 1927, p. 27; Fauvel 1932, p. 231; Aziz

1938, p. 47; Fauvel 1953, p. 449; De Silva 1965,

p. 559.

Habitat. It is commonly available at Moosa-

mani lock, Lighthousekuppam and at Gunan-

kuppam stations.

Description. 20-30 mm long. Prostomium

bears about 10 to 12 branchiae. Each branchia

possesses about 40-45 filaments. Gills without

eyes. Colour of the worm differs from

Southern’s (1921) description. All the worms

are pale green with white stripes. The arrange-

ment of setae agrees with Southern’s (1921)

description. Tube is made up of mud and sand.

Occurrence in Indian waters. Chingrighatta

near Calcutta and Vishakapatnam.

Distribution outside Indian waters. Malay

Archipelago.

Remarks. This species closely resembles

Potamilla ceylonica Augener (1926), but it

differs from P. ceylonica with regard to the

arrangement of setae in both the thoracic as

well as in the abdominal segments.

Family Serpulidae

Species 25. Hydroides norvegica Gunner us

1768

(Plate 4, figs. 25a & 25b)

Eupomatus elegans Haswell 1883, p. 633;

Hydroides multispinosa Marenzeller 1884, p. 21;

Hydroides norvegica Pixell 1913, p. 74; Hydroides

multispinosa Augener 1914, p. 139; Hydroides nor-

vegica Fauvel 1927a, p. 356; 1932, p. 242; 1953, p.

458; Filial 1960, p. 12.

Habitat. Large numbers of the tubes of this

species were collected from the crevices of

oyster shells at the Kulathumedu station, but

worms were present only in some tubes.

Description. 20-25 mm long. Tubes white,

cylindrical and more or less erect. Gills around

the mouth, bearing numerous radicles with

two rov/s of barbules. Thorax possesses dorsal

capillary setae and ventral uncinigerous tori,

but the abdominal segments possesses dorsal

98

POLYCHAETA OF THE PUL1CAT LAKE

uncinigerous tori and ventral capillary setae.

Operculum well developed.

Occurrence in Indian waters. Madras.

Distribution outside Indian waters. Persian

Gulf, Red Sea, Atlantic Ocean and Mediterra-

nean Sea.

Remarks. These worms are purely marine

and sedentary forms.

Distribution and Ecology

The two major factors which influence the

distribution pattern of the polychaetes in the

Pulicat Lake are salinity and the nature of the

substratum. On the basis of salinity, the lake

may be divided broadly into three habitats,

namely marine, brackishwater, and freshwater

habitat.

The marine habitat includes stations like

the Pulicat pass, Karimanal, Gunankuppam,

Lighthousekuppam, Kottaikuppam lock and

Dhonirevu. The polychaetes collected at these

stations include mostly marine forms (Table

2). The salinity at these stations ranges from

25.3l5fc to 38.45%c.

The brackishwater habitat includes the rest

of the stations, except the stations in the three

river beds, namely the Araniar river, Kalinga

river and the Swarnamuki river. The average

salinity at the brackishwater stations varies

from 11.65*o to 48.51%c. Out of the 14

species collected from this habitat, Nerine

cirratulus was collected only from the Kottai-

kuppam lock and Lumbriconereis simplex and

Lumbriconereis polydesma were collected only

from Dhonirevu. This shows that these three

species are designed to live at these stations

alone, mainly because of the nature of the

substratum rather than the salinity at these

stations.

The freshwater habitats of the Swarnamuki

river, Kalinga river and the Araniar river have

the river bed mostly sandy and in some places

it is muddy. Distribution pattern of the poly-

chaete worms in the freshwater habitat is

mainly controlled by salinity. The polychaetes

collected in the freshwater habitat are Mar-

physa gravelyi, Nereis chilkaensis , Lycastis

indica, Heteromastus similis, Potamilla lepto-

chaeta, and Diopatra neapolitana. The species

Potamilla leptochaeta and Diopatra neapolitana

were not available during the monsoon season

in the freshwater habitat. This might have

been due to the lowering of salinities of the

river water during such monsoon seasons.

Besides salinity the other major factor deter-

mining the distribution of the polychaetes in

Pulicat Lake is the nature of the substratum.

On the basis of the type of substratum, Pulicat

Lake may be considered to have four types

of substrata for polychaetes, namely sandy,

sand-silt, weedy and the oyster-bed substrata.

The sandy substratum in Pulicat Lake is

distributed at the following stations, namely

Pulicat Pass, Karimanal, Dhonirevu, Gunan-

kuppam, Lighthousekuppam, Kottaikuppam

lock, Irakkam, Venadu, Sriharikota and Ataka-

nitippa. Interstitial polychaetes like Pisione

complexa, Pisionidens indica and Eteone baran-

tollae were collected only from such sandy

substratum. Other polychaetes in sandy sub-

stratum include Glycera alba, Prionospio

krusadensis, Tylonereis fauveli, Euclymene

annandalei and Euclymene insecta.

The sand-silt substratum is observed at the

following stations, namely Annamalaicheri,

Arambakkam, Tada, Berupet, Malan, Roy-

duruah, Edamanai and Duggirajapatnam. The

muddy substrata at these stations are dark in

colour, and possess decayed vegetable matter.

The polychaetes collected at these stations

were Marphysa gravelyi, Tylonereis fauveli,

Nereis chilkaensis, Lumbriconereis simplex,

Lumbriconereis polydesma, and Heteromastus

similis. This sand-silt or sand-clay substratum

extends over the majority of the lake, and the

concentration of the polychaetes was maximum

in this type of substratum.

99

Marine Stations Brackishwater Stations

JOURNAL , BOMBAY NATURAL HIST. SOCIETY, Vol 84

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POLYCHAETA OF THE PULICAT LAKE

The weedy substratum is observed at a few

stations only. Arangam, Pulincheri, Moosamani

lock and Zonangipalam are the only four

stations where the substratum is covered with

weeds. The bottom macrophytes consist mainly

of rooted submerged plants like Halophila

ovalis and Cymododea isoetifolia and filamen-

tous algae such as Chaetomorpha sp., Entero-

morpha sp., and Acetabularia sp. etc. The

marine angiosperm Halophila ovalis (Gaud)

flourishes in places where soil is dark and

clayey. Generally polychaetes are plenty in

places where this plant is abundant. Polychaetes

are said to be detritus feeders, hence there is

some association noticed between polychaetes

and this vegetation; perhaps these worms feed

on the decayed leaf blades of this plant. Mar-

physa gravelyi in fact was collected more in

areas where this plant thrives.

The oyster beds form another kind of sub-

stratum in Pulicat Lake. They occur in Kula-

thumedu village. A number of polychaetes

were collected only from the oyster beds. The

oysters of Pulicat Lake belong to the species

Crossostrea madrasensis (Preston) and Sacco-

strea cucullata (Born). The oysters have spread

even upto the Kottaikuppam lock. The poly-

chaetes associated with these oysters are Poly-

Refer

Aiyar, R. G. (1933) : On the anatomy of Mar-

physa gravelyi (Southern). Rec. Ind. Mus: 35(3):

287-323.

& Alikunhi, K. H. (1940) : A new

Pisionid from sandy beaches Madras, ibid. 42: 89-

107.

Alikunhi, K. H. (1947) : On a Pisione complexa,

new species from the sandy beach, Madras. Proc.

Nat. Inst. Sc. India. 73(3) : 105-127.

Apurba Ghosh (1963): On a collection of poly-

chaeta from the south-east coast of India with a

new eunicid record. J. Mar. Biol. Ass. India., 5. (2) :

239-245.

dor a ciliala, Hydroides norvegica, Amphicteis

gunneri, Hesione intertexta, Harmathoe ampul -

lifera, Diopatra neapolitana and Nereis chilka-

ensis. Out of these, Hydroides norvegica and

Polydora ciliala are encountered more in num-

ber. The tubes of Hydroides norvegica are

chalky white, cylindrical and slightly convo-

luted. The tubes are more or less erect or

sometimes spirally coiled. Atleast eight to ten

tubes are found attached to a single oyster

shell. Polydora ciliala is a spionid polychaete

which is also collected in large numbers from

this lake. The members of Polydora are said

to be pests of bivalves. They form mud-blisters

on the mantles of bivalves (Stephen, 1978b).

It is now considered to be a major problem

in coastal shellfish aquaculture.

Acknowledgements

We are grateful to Dr K. Rama-

moorthy and Dr Mary Florence for their

valuable suggestions in planning this work.

Our sincere thanks go to Dr P.H.D.H. De

Silva, Director of National Museums, Colombo,

Sri Lanka, for identifying some of the poly-

chaetes encountered during the course of this

study.

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104

A CATALOGUE OF THE BIRDS IN THE COLLECTION

OF BOMBAY NATURAL HISTORY SOCIETY — 32

Humayun Abdulali

Muscicapidae (Turdinae)

[Continued from Vol. 83(2): 359]

This part ending with Synopsis No. 1691

up to Register No. 265171 includes 888 Speci-

mens of 56 species of the Turdinae. There are

8 forms (5 species and 3 subspecies) of which

we have no specimens in our collection. Dr

(Mrs) S. Unnithan has assisted with the work.

1635 Brachypterix stellate stellate Gould

(Sikkim) Gould’s Shortwing 2:16

nil.

1636 Brachypterix hyperythra Jerdon &

Blyth (Darjeeling) Rustybellied Shortwing 2:17

15 (by plumage) Margherita, Assam.

Measurements on p. 117.

1636a. Brachypterix cryptica Ripley (40-

mile Camp east of Miao, Noa Dihing River,

Arunachal Pradesh).

This bird described in 1980 ( JBNHS 77:

p. 1) has been found to be Tickell’s Babbler

[Trichastoma tickelli (Blyth) ( JBNHS 81 p.

700/1]

1637 Brachypterix major major (Jerdon)

(Nilgherries) Rufousbellied Shortwing 2:10

2: $ $ 1 Avalanche, 6700', 1 Longwood Shola,

Kotagiri, Nilgiri.

Measurements on p. 117.

1638 Brachypterix major albiventris (Blan-

ford) (Palni Hills) Whitebellied Shortwing

2:11

2: 1 $ 1 $

1 Moir Point, 6500', 1 Kodaikanal 7000', Palnis.

Measurements on p. 117.

[593]

1639 Brachypterix leucophrys nipalensis

Moore (Nepal) Lesser Shortwing 2:19

9: 2 $ $ 6 9 $ 1 o?

4 Margherita, Assam; 4 N. Cachar Hills; 1 Mt.

Victoria, Burma.

Neither of the two males show any blue

and Fleming (1970) in birds of Nepal, pp.

248 states “birds with blue not yet seen here”.

Koelz when describing geokichla from Manipur

(which is not accepted by Ripley) says he

examined 11 S d none of which were blue.

The unsexed bird from Mt. Victoria is the

smallest in all respects.

Measurements on p. 117,

1640 Brachypterix montana cruralis (Blyth)

(Darjeeling) Whitebrowed Shortwing 2:17

11:6^3 (1 blue) 5 9 9

1* Rangpo, Sikkim; 1 Honka, 1 Gedu, West

Bhutan; 1 Darjeeling, 1 Miao, 1 Deban, 1 Firm

Base, Arunachal Pradesh; 4 Margherita, Upper

Assam.

Five of the six males are in female plumage,

but have larger wings, *the one blue bird hav-

ing the longest. All except one dated 22nd

January 1903 show a white streak over the

eye.

Measurements on p. 117.

1641 Erythmpygia galactotes familiaris

(Menetries) (Kur Southern Caucasus) Rufous

Chat or Greybacked Warbler 2:386

24: 12 5 $ 5 $ $ 8 o?

2 Muscat ; 1 Feluja, 1 Rustem, 1 Fahama, 1 Bagh-

dad, 5 Basra, Mesopotamia: 1 Shiraz, 1 Kasoin,

105

8

JOURNAL, BOMBAY NATURAL HIST. SOCIETY , Vol. 84

Persia ; 1 Chaman, 1 Kandera Pass, east of Mashkai

Valley, Baluchistan; 1 Sakesar, Shahpur Dist., 1 Mul-

tan, Pakistan; 4 Bhujia Fort, 1 Nakhatrana, 1 Shinai,

Anjar, Kutch; 1 Dwarka, Okhamandel, Kathiawar.

Measurements on p. 117.

EL Erythropygia galactotes syriacus Hem-

prich & Ehrenberg (Beyrouth)

1 $ Felujah, Mesopotamia.

This is slightly rufous above and marked

“indistinguishable from syriacus

Measurements on p. 118.

1642 Erithacus megarhynchos hafizi (Severt-

zov) (Turkestan) Persian Nightingale 2:87

nil.

EL Erithacus megarhynchos africana (Fis-

cher and Reichenow) (Klein Arusha, near

Kilimanjara) Nightingale

10: 6 8 8 2 $ $ 2 o? (1* fledgling)

1 Sheikh Saud, 1 Fao Iraq ; 2 Enzeli, Caspian

Coast, 2* Gulahek, Tehran, 3 Shiraz, 1 Bog Miza-

khani.

Specimen No. 3174 from Enzeli is marked

by Cheesmann <fafricana in song and evidently

breeding. This is the new name for the Persian

Nightingale hafizi’ \ This specimen is very dark

and similar to Luscinia luscinia but can be

separated on the characters mentioned there-

under.

Measurements on p. 118.

1643 Erithacus calliope (Pallas) (Yenesei)

Ruby throat 2:91

22: 16 8 8 (2 by pi.) 4 $ 9 2 o?

1 Bharatpur, Rajasthan; 1 Partapur, Nepal; 1 Sir-

pur, 1* Sarun, 1 Assam; 3 Baghowni, Tirhut; 2

Darbhanga, Bihar & Orissa; 1 Tikerpara, 1 Bhusand-

pur, Chilka Lake, 2 Mahendragiri, 1 Berbera, Orissa;

2 Sankrametta, 1 Padevalsa, Vizag Dist.; 1 Goda-

very Delta; 1 Tientsein, 2 Peking (cage birds),

China.

The last two males from Peking have the

largest wings — 81 and 82 mm and the chin

is pale pink in one and almost white in the

other. Other males from Assam, Sarna and

Godaveri Delta have pinkish chins but are no

larger. The earlier specimens are marked

camschatkensis, but this race is no longer

recognised.

Measurements on p. 118.

Erithacus svecicus Bluethroats

This is an extremely difficult group to divide

into the many subspecies named, for to quote

Vaurie (1959) “Some, perhaps most, of the

variation is clinal in some characters but it is

complicated by much local variation, seasonal

changes in coloration and pattern, and sexual

dimorphism in the males of some populations”.

Our material consists largely of non-breed-

ing migrants and this makes it more difficult.

There has been no alternative but to accept

the trinominal identifications made by earlier

workers or distributional limits accepted in

Indian literature. More extensive ringing ac-

companied by appropriate collecting may help

to clarify matters.

1644 Erithacus svecicus svecicus (Linnaeus)

(Sweden) Northern Bluethroat 2:83 and 85

10: 9 8 8 1 o?

1 Peshawar, N.W.F.P., 1 Nawashahar, Jullundur,

1 Ambala, Punjab; 1 Sanchi, Bhopal; 1 Baghownie,

2 Madhubani, Darbhanga dist., Bihar; 1 Cawnpur,

U.P.; 1 Laisingah, Cachar, 1 Peking, China.

Measurements on p. 118.

1645 Erithacus svecicus pallidogularis

Zarudny (Orenburg, Russia, c. 55 E, 52 N.)

Turkestan Bluethroat 2: 85

43: 27 8 8 13 9 $ 3 o?

3 Chitral, N.W.F.P.; 1 Wana? Waziristan; 8 Am-

bala, 1 Shikohpur, Jullundur, 1 Campbellpur, 1

Jagadhri, Punjab; 1 Harunabad, Bahawalpur; 1

Kashmir; 1 Badantpur, Bhaji State, Simla Hills; 2

Hamavas Lake, Pali, Jodhpur, Rajasthan; 2 Delhi;

4 Bahawalpur; 1 Kutch; 1 Amreli, Kathiawar, 1

Mehmedabad, Kaira, 1 Ajwa, Baroda, 1 Dohad,

Gujerat; 1 Wada, 1 Shil, 1 Murbad, Kalyan, 1 Bela-

pur, 1 Andheri, Bombay; 1 Madhubani, 1 Baghowni,

Tirhut, Darbhanga, Behar; 1 Barul, 2 Cawnpur, U.P.;

1 Laisingah, Cachar, 1 Prome, Burma.

106

[594]