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ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

| Go]: bh VOLUME 66

THE TRUSTEES OF THE DIE TRUSTEES VAN DIE

SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

CAPE TOWN ; KAAPSTAD

SMU ANON

1974-1975 MAY 25 1975

iRpAReS

# PRINTED IN THE REPUBLIC OF SOUTH AFRICA BY

THE RUSTICA PRESS (PTY.) LTD., WYNBERG, CAPE

561

ANNALS OF THE

SOUTH AFRICAN MUSEUM

VOLUME 66

LIST OF CONTENTS

BoumeE, H. E.

See

SHaw, E. M. & BOumeE, H. E.

BRANCH, G. M.

A new species and records of Scutellidium (Copepoda, Harpacticoidea) from

South Africa, with a world key to the genus (published February 1975)

BuRRAGE, B. R.

Population structure in Agama atra and Cordylus cordylus cordylus in the vicinity

of De Kelders, Cape Province (published October 1974)

CLUVER, M. A.

The cranial morphology of the Lower Triassic dicynodont Myosaurus gracilis

(published October 1974)

Cooper, M. R.

The Cretaceous stratigraphy of south-central Africa (published November 1974)

Day, J. A.

South African Cumacea. Part 1. Family Bodotriidae, subfamily Vaunthompsoniinae

(published February 1975) ..

GRINDLEY, J. R.

See

MCLACHLAN, A. & GRINDLEY, J. R.

Hesse, A. J.

Additions to the South African species of Phthiriinae and Usiinae (Diptera:

Bombyliidae) with keys to all the known species (published March 1975) ..

HEsseE, A. J.

A new South African representative of the South West African genus Namibimydas

Hesse (Diptera: Mydaidae), with some ecological notes on the habits of the

species (published October 1974) .. 2

KENSLEY, B. F.

Type specimens of Decapoda (Crustacea) in the collections of the South African

Museum (published November 1974)

McLAcuLan, A. & GRINDLEY, J. R.

A new species of Mystacocarida (Crustacea) from Algoa Bay, South Africa

(published November 1974) i. ay ay Be, re

ROELEVELD, M. A.

A revision of Massey’s checklists of ‘South African’ obs Sai ere

March 1975) ; : My ;

SHAW, E. M. & Boume, H. E.

Pedi skin dressing technique (published November 1974)

WINTERBOTTOM, J. M.

The zoogeography of the South African avifauna (published November 1974) ..

PAGE

221

177

257

169

233

151

109

ea 24

‘\ OLUME 66 PART 1 OCTOBER 1974

{

OF THE SOUTH AFRICAN

“= MUSE

CAPE ‘TOWN

— =

)

. =

, re

+f 3

‘

a ..

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 66 Band

October 1974 Oktober

Part 1 Deel

POPULATION STRUCTURE IN

AGAMA ATRA AND CORDYLUS CORDYLUS CORDYLUS

Beene, VICINITY OF DE: KELDERS; C.P.

BRYAN R. BURRAGE

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

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POPULATION STRUCTURE IN

AGAMA ATRA AND CORDYLUS CORDYLUS CORDYLUS

IN THE VICINITY OF DE KELDERS, CAPE PROVINCE

BRYAN R. BURRAGE

South African Museum, Cape Town*

(With 5 tables)

[MS. accepted I October 1973]

CONTENTS

PAGE

Introduction . 5 ‘ : 5 : ; 1

Materials and methods . : 5 5 ; 1

Description of the terrain. 5 ; ‘ ; 2

Observations and discussions

The population and habits of the lizards . 3

The territories and home ranges 5 ; 4

Display patterns : : ; ‘ ; 16

Social interactions . : : : : 16

Summary : : : : : : : 20

References. 4 5 : 3 : : 21

INTRODUCTION

Ecological studies on southern African reptiles have been undertaken since

January 1969. This paper describes the spatial and social organization of the

Cordylus cordylus cordylus and Agama atra inhabiting the vicinity of De Kelders,

Cape Province.

The ecology of many American reptiles has been investigated to the point

where general tenets and recognition of organizational patterns can be made

(Carpenter 1967; Rand 1967; Ruibal 1967). The same cannot be said of Old

World forms, though North African and Australian studies are filling in the

picture. It seems proper to improve our knowledge of southern African forms.

Little has been done on the overall systematics of southern African lizards

since FitzSimons’s (1943) excellent work. The identity and relationships of

some forms are confusing. Before worth-while ecological work can be done, it

is important to identify properly just what species are being worked on.

MATERIALS AND METHODS

Populations of Cordylus cordylus cordylus and Agama atra were studied on

10 hectares of coastal cliffs, 2 hectares of inland rock outcrops, and almost 2

hectares in sandy situations (0,7 ha isolated rocks on sand; 1,188 ha isolated

* Present address: College of the Desert, Palm Desert, California.

]

Ann. S. Afr. Mus. 66 (1), 1974: 1-23, 5 tables.

2 ANNALS OF THE SOUTH AFRICAN MUSEUM

koppies amongst sand dunes). Lizards were captured by ‘chase and grab’, can-

trapping, or stunning by being shot with a rubber band cut from an inner tube.

These animals were then marked (see Table 1) by toe-clipping and branding

with an identification number. Animals were observed from vantage points by

means of binoculars. Points visited by a lizard were marked at the site with a

rock on which the lizard’s number, sex, and observation data were printed in

waterproof ink. The points were simultaneously plotted on a scaled map on

graph paper of each study area. The denser the dots, the greater the activity

of the lizard within a given part of its occupied area, and the peripheral dots on

the map and site markers gave the exact area. Peripheral markers were small

tripod rock cairns upon which a rock (with identity markings) pointed to the

centre of a given occupied area. Connecting the outlying dots gave the periphery

of the occupied area, the extent of which was then computed with a planimeter

from the graph paper. Social interactions were marked on the site where they

occurred and on the graph paper to determine usage of a given occupied area

and what part(s) were most vigorously defended. Territories and home ranges

were computed only on the basis of ten or more captures per lizard.

Table 1

Cordylus cordylus cordylus and Agama atra marked by toe-clipping at various study stations

March 1971 to oo la Mane fe September ee 1972.

ee ro ips ke |

Study station eee. Juveniles 33 99 Total

Coculciiic 20) > eee ee cliffs

Agama atra. ' : 620 78 Li) TH

Cordylus cordylus 406 82 83 S78

Inland rock outcrops

Agama atra. : 204 25 26 235

Cordylus cordylus 312 52 52 416

Rocks among dunes

eee Fae eee cordylus 248 113 A 472

DESCRIPTION OF TERRAIN

The Walker Bay coast from the rocky cliffs at De Kelders, north into the

dunes is the study area. The rocky cliffs are mostly of the Table Mountain

Series, composed of sandstone, quartzite, conglomerate, and shale. To the north

are cliffs and outcrops of calcareous dune sand of the Bredasdorp Beds. While

there are some sheer cliff faces to seawards, cliffs of the Table Mountain Series

often present a sequence of step-like ledges. Cliffs of the calcareous dune sand

present somewhat the same aspect, but with greater overhanging ledges. The

old dune bedding planes are quite clear. Immediately landwards of the cliff

edges are small outcrops which appear as step-like tables.

The shifting dunes are composed of white sand and are arranged in roughly

AGAMA ATRA AND C. C. CORDYLUS IN THE VICINITY OF DE KELDERS, C.P. a

north-south longitudinal series. Among the dunes immediately on the maximum

high tide line are some rock outcrops. Further inland among the dunes are

scattered rock koppies of varied size, but of essentially similar configuration.

There are small koppies (Table 2: koppies ‘A’ & ‘B’) not exceeding 2 m in

height, which may be one unit, as are the larger koppies, or discontinuous with

intervening sand. The larger koppies present an appearance of jumbled terraces

(the “Tiers’ of koppie ‘C’ in Table 2) at different heights from ground level to

less than 5 m high. Also, amongst the dunes are scattered large flat rocks, none

of which is more than | m high. At the eastern edge of the dune area, and in

some places extending into the dune area in finger-like projections, are sandy,

brush-covered hummocks, some having a rock outcrop core.

This area lies in the winter-rainfall area of the coastal western Cape. The

vegetation 1s Coastal Macchia (Acocks 1953), consisting of shrubs, grasses and

other annuals. On the sandy beaches are a variety of succulents. The dunes are

devoid of vegetation, except at their bases, but this may be vegetation which

they are burying. Some koppies appear to have been exhumed as the dunes

move on and these are devoid of woody vegetation, having only a few succu-

lents and grasses. Other koppies seem not to have been recently buried and

these have woody vegetation.

The Forestry Department is presently engaged in reclaiming these dunes,

destroying this habitat in its current appearance.

OBSERVATIONS AND DISCUSSIONS

THE POPULATION AND HABITS OF THE LIZARDS

Agama atra is commonest (155 adults per hectare) on the bare rock surface

of the coastal cliffs, and occurs sparsely (51 adults per hectare) on isolated bare,

flat rock outcrops surrounded by open shrub. It is absent on rocky outcrops

surrounded by dense scrub or large expanses of sand, and on the koppies and

rocks amongst the dunes. Juveniles of varied age groups are up to four times

as dense as the adults.

Cordylus cordylus cordylus occurs in all of the above habitats, as long as

there are some rocks, which are the focal point of its activity and in which the

lizards have their retreats. Those inhabiting the coastal cliffs are predominantly

of a yellowish-phase, but only brownish-gray ones inhabit suitable habitats

among the dunes. On the terraces of large rock koppies the population density

of adult cordylids is as high as 288 per hectare, and as low as 4,2 per hectare in

the area of isolated rocks on sand. Juveniles of varied age groups are two to

three times as dense as the adults. The adult population density of both species

in the various habitats is given in Table 2.

While the spatial organization of Agama appears unvarying, that of Cordylus

differs according to the habitat. Agama and Cordylus co-inhabit the coastal

cliffs, but competition is lessened by the different thermal requisites of each and,

hence, different activity patterns. Cordylus is abroad in the early morning and

4 ANNALS OF THE SOUTH AFRICAN MUSEUM

late afternoon on sunny days, and abroad throughout the day under overcast

conditions. However, Agama prefers higher temperatures and is most active

when Cordylus has sought the cool of its shelters.

THE TERRITORIES AND HOME RANGES

Agama atra adults of both sexes are typically territorial (Table 2). Despo-

tism occurs among the more crowded populations along the immediate shore.

Despotic individuals are males, almost invariably those with the largest terri-

tories in this situation. Territorial borders are usually delimited by ledge edges,

fissures, and the like. The highest part of all territories is the focal point, allowing

the occupier a vantage site for surveying its domain. The vertical area is greater

Table 2

Particulars of the spatial organization of Agama atra (Aa) and Cordylus cordylus cordylus (Cc)

in the vicinity of De Kelders, Cape. All areas shown are in square metres. “T’ denotes territory

and ‘HR’ denotes home range in the table. Only the N of Territory and/or home range holding

adults with a minimum of 10 recaptures is shown (compare with Table 1).

Adults per Type of social

Species 3d 2° hectare Koppie organization

Coastal cliffs

N=67 N=68

Aa . «= 78,0-188;5,@2:5) +22;5-91,5 G60) 125,0 — dt

N=75 N=76

Cc. . . 65,6-124,5 (90,0) 33,2-100,0 (80,0) 165,0 — Hf

Inland rock outcrops

N=20 IN=—20

Aa. . . 98,8-250,0 (120,0) 32,5-105,0 (77,0) 51,0 — i%

N=46 N=45

Cc. . . 70,0-130,0 (110,0) 60,7—100,1 (82,6) 104,0 — T

Koppies situated in dunes. Adult gd and

9 share a territory and/or home range

with their juveniles.

(only) ee) ee

” 4,0-20,5 (14,1) “i

Ne O56. 0466 Gomi, oe s HR

4,0-22,0 (1327) T

8 8) apie eae z HR

4,0-16,0 (8,5) T

26 | 72, 40-000 66) 2 0 ee HR

10 12 4,0-16,0 (8,9) 244,2 C Tier If HR

24 24 4,0-10,0 (7,0) 288,0 C Tier III HR

20 18 4,0-12,0 C719) 285,0 © Tier TV HR

4 6 4,0-12,0 (7,6) 260,0 C Tier V HR

Isolated rocks situated in sandy areas.

Adult 33 and 99 share a territory with

Ge their juveniles.

(only) 33 29

nN—7 ad 84,0-180,0* (108,9) 4,2 gi

* Does not reflect a litigated area of 18,0 m® between two Cordylus groups, nor changes in

territory sizes after the removal of Group | (see Table 3).

AGAMA ATRA AND C. C. CORDYLUS IN THE VICINITY OF DE KELDERS, C.P. 5

than the horizontal, allowing more effective surveillance of a larger territorial

area than could be realized if the territories were arranged along a contour.

While such an arrangement is favoured by the topography, it allows more

agamids to maintain effectively larger territories than could be realized on flat

areas. Unfortunately, maps drawn in plan view do not accurately reflect those

areas in steep slope.

Territories extend seawards almost to the water’s edge, the Agama

including various intertidal arthropods in its diet. All male territories are larger

than those of the females, and each male has a group of two or three females.

The smaller territories of the females are wholly or mostly within that of the

male. Members of each group have their preferred resting-spots. Females within

the territories of despotic males seem to be more aggressive in defending their

respective territories from other females of their male’s territorial group, but

markedly so against females of another male (see also ‘Social Interactions’)

especially those of the non-despot males. The development of despotism in

such areas may indicate that the population pressure is reaching the limits for

a territorial organization. If the population pressure increases, so much time

may be spent in territorial defence as to be detrimental to the well-being of the

species. Creation of a hierarchical structure or despotism would reduce or

eliminate antagonistic behaviour. With greater population pressure, a social

organization of ‘free-run’ could be expected. Another alternative would be the

reduction of territories to a certain size and the stabilization of population size.

Territories of A. atra are largest in areas with lower population densities.

The nature of the topography seems to allow the coastal agamids to establish

and maintain larger territories than the high population density would allow

on areas of a flatter or more uniform topography.

A study of an intertidal population of Agama atra, Mabuia capensis and

M. homalocephala inhabiting scattered rock along the Rooi Els River mouth

and beach showed that agamids are limited to such rocks as occur, and many

occupy larger territories than is reported here. This is especially true of those

occupying scattered rocks right at the water’s edge and isolated from other

suitable habitat by sand or dense brush. In this situation there is no despotism.

Cordylus cordylus has a higher population density on coastal cliffs and

territories of both sexes are smaller than those on inland rock outcrops. In all

situations female territories are smaller but rarely overlap those of more than

one male (see also ‘Social Interactions’). The sexes do not form groups in the

manner of A. atra. C. cerdylus digs a burrow, usually located in a redoubt, as

in the soil deep in a rocky crevice. In their burrows they are virtually secure

from any danger, since they inflate the body, grasp the sides of the burrow with

their clawed feet, and present the spiny tail to the intruder. The delimitation of

C. cordylus territories closely follows that described above for A. atra in these

situations.

In dune and sandy areas C. cordylus occurs in isolated population subunits,

closely connected to koppies or areas of scattered rock, The social organization

6 ANNALS OF THE SOUTH AFRICAN MUSEUM

here is quite complex and obviously closely regulated to the rocky habitat they

prefer. Each territory or home range is occupied by an adult male and female

along with their juveniles of varied age groups. In the territories the adults of

both sexes fiercely defend their area against intrusion by any lizard of any age

group. In the presence of adults, the juveniles sometimes assist in the defence

of the area. In the home ranges, each group has its own foraging area but tran-

sients are tolerated. Burrows of each group are always defended. In home ranges

and territories the retreats are located in rocky crevices or burrows and are

occupied by the entire group. Such burrows have a narrow entry way and very

large, often branching terminal enlargements. Thus, the C. cordylus ‘family

group’, where it occurs, is a tighter organization than that of A. atra.

On small koppies (‘A’ and ‘B’ in Table 2, and see also pp. 16-17) C.

cordylus has a territory and home range with the territory on the periphery. The

territory includes some of the rocky area, but mostly it is a skirt of sand of

varying width around the koppie. The lizards’ burrows are located in the rocky

area. One group in koppie ‘B’ had only a territory. Two groups in koppie ‘C’

and one group in koppie “B’ had only home ranges. Those maintaining only

home ranges were located more or less in the centre of the koppies, being com-

pletely surrounded by other cordylids. The borders in koppie ‘C’ closely follow

the demarcation by large blocks of rock, but in the more scattered koppie ‘B’

the borders are more arbitrary.

The spatial organization of C. cordylus on koppie ‘C’ closely follows the

step-like arrangement of koppie ‘C’. Thus, they can be arranged in tiers from

the periphery (Tier.I) of the koppie, to each successively higher terrace, inwards

to the centre (Tiers II-V). On this koppie only the 24 groups of Tier I maintain

a territory and home range, being territorial on the periphery, with the home

range on the rocky area. The territories consist only of a sandy area of varying

width, mostly narrow and especially so on the western edge of the koppie

where a large dune closely abuts. The population density increases inwards and

the size of the home ranges diminishes. Slight deviations from this trend are

due to larger rock blocks at the centre, possibly influencing border demarcation

and the density of lizards to occupy such home ranges. Borders of home ranges

closely follow that of the jumbled large rocks. That is, a home range is limited

to one large rock or associated rocks on each tier.

Population density varies inversely with territory size. Such territories

occur on sandy areas with scattered large rocks, and on sandy hummocks with

rocky core and brush cover. Generally speaking, those territories on sandy

areas with scattered large rocks were larger than those situated on the hummocks.

These cordylids maintain only rigidly defended territories. The centre of activity

is on the scattered rocks, especially those in which their burrows are located.

They do forage over the sand between the rocks within their territory. Five of

the seven such C. cordylus groups had borders abutting on others. The two

C. cordylus groups with the largest territories (Group |: 180,0 m?, and Group 2:

116,0 m2, respectively) often disputed a litigated border area of 18,0 m? along

AGAMA ATRA AND C. C. CORDYLUS IN THE VICINITY OF DE KELDERS, C.P. 7

a dry wash on their borders. The reason for the dispute is not clear, since the

dry wash is mostly hard sand, having no rocks of any size to battle over. Since

its ownership alternately shifted and its integrity apart was sometimes mutually

observed, I consider it a ‘buffer zone’. C. cordylus Group 3 occupied a 100 m2

territory situated in a sandy area with rocks but had no common border with

any other C. cordylus group. Situated on the hummocks were Group 4 with

104 m? (partly bordering on Groups | and 5), Group 5 with 94 m? (bordering

on the territories of 1 and 4), Groups 6 and 7 with 84 m? each, of which only

Group 6 bordered partly on Group 5.

The removal of both adult cordylids, plus some juveniles of Group 1,

resulted in an interesting adjustment for the surviving juveniles and reappor-

tioning of this territory amongst the surrounding groups. This reapportioning

can be divided into three stages over 120 days, details of which are given in

Table 3. In the first stage Group 2 took over the litigated area and 72 m? of

the territory of Group 1, including most of Group 1’s retreat rock, its 1 m

high pinnacle and another rock. Group 4 took over 16 m? of the territory of

Group 1, and while mostly borderland it included a part of Group 1’s retreat

rock, but not the burrow or pinnacle. Group 5 took over 56 m? of the territory

of Group 1, consisting mostly of sand and one rock, quite isolated from its

original holdings. This arrangement lasted for about a month, and over the

subsequent three weeks further adjustments occurred, largely in the favour of

Group 2.

It is somewhat interesting that Group 2 maintained the integrity of its

Table 3

Territorial changes of Cordylus cordylus cordylus groups (Nos 1-7) inhabiting isolated rocks

in sandy areas. Each group composed of one adult male and one adult female and a number

of juveniles (see Table 1). After the removal of the adults of Group 1, the acquisition is shown

of Group 1 territory by the other groups, with their new total territories in parentheses. All

areas shown are in square metres.

November March

30 days later

120 days later

1 1804 some removed —- 128

2 1162 +90 (106)> +20 (226)» 134»

3 100 no change no change no change

4 104 +16 (120) +8 (112) 104

5 95 = Oncol) +44 (139) 95

6 84 no change no change no change

ff 84 no change no change no change

= Does not include litigated area of 18,0 m?.

= Includes litigated area of 18,0 m?.

= In stage 2, Group 2 added Group 1 territory taken by Groups 4 and 5, their reduced

acquisition of Group 1 territory is shown.

ad — In stage 3 the surviving Group 1 juveniles had evicted Groups 2, 4 and 5 from the

former territory of their adults.

8 ANNALS OF THE SOUTH AFRICAN MUSEUM

swollen territory, even though it seemed difficult to defend. That is, its original

central rock with the retreat burrow was located more in the western extreme

of its territory; quite far removed from the eastern limits of its annexations of

Group | land, located closer to the territories of Groups 4 and 5. Also, the

territory of Group 2 bordered on two territorial groups in Tier I of koppie ‘C’,

neither of which was observed to intrude into Group 2 territory.

More interesting was the surviving Group | juveniles, who in the absence

of their parents and some of their peers, did not make any obvious attempts to

maintain any territorial integrity of the whole or part of the Group | territory.

It could be assumed that they could not defend it in the manner or to the extent

that the adults did, but their defending the central rock (about 16 m?) and retreat

burrow would seem barely plausible. Group | juveniles were apparently taken

over, integrated with, or just ignored by the successors to their parents’ territory.

During and after the stabilization of the situation, Group | juveniles continued

to occupy the area, though they seemed limited to an area of about 20 m?,

mostly concentrated to the central rock. They were not challenged, but seemed

submissive in the presence of C. cordylus from the other groups which had

annexed parts of Group | territory. They did continue to utilize their own

retreat, which, on occasion was also occupied by C. cordylus of Group 2. The

latter, however, did not occupy the Group | retreat on a permanent basis and

maintained their own retreat. It hardly seemed conducive to lizard occupancy

to disturb the retreats too often.

In March 1972 the area was reinvestigated, with the interesting discovery

that the newly mature older Group | juveniles had acquired territorial desires

and had reclaimed and were successively defending most of their old Group |

territory (128 m?), except an area of mostly sand with two small outlying rocks,

which was little used even when their parents were present. Groups 2, 4 and 5

individuals had been totally evicted from Group | territory, but Group 2 inte-

grated the former litigated area into their territory. Group 2 territory was not

what it was when first studied.

Examination of C. cordylus territorial organization in other koppies,

hummocks and sandy areas with scattered rocks follow the same pattern

described above for that investigated in detail in the particular study site. While

no detailed investigation was undertaken of C. cordylus territories in the rocky,

sandy beach areas in the supratidal area, it appears to follow that of the dune

sandy areas with scattered rocks.

Juvenile cordylids on coastal cliffs and inland rock outcrops have ‘free

run’ of adult territories, but gradually disperse as they mature and take up

territories where they can defend an area. In both Agama atra and C. cordylus

there appears to be a ‘phantom population’, mostly of recently matured adults,

that does not defend a territory, possibly because it cannot find a place to defend

due to those already ensconced. In rocky situations the ‘phantom population’

has a reasonable chance of surviving until vacancies occur. On isolated rocks

in sandy areas juvenile cordylids disperse as they mature and try to set up a

AGAMA ATRA AND C. C. CORDYLUS IN THE VICINITY OF DE KELDERS, C.P. 9

territory of their own, which seems to be a difficult task as almost all available

sites are occupied. Since maturing juvenile males are evicted if they show interest

in the female (their mother) of the adult male (their father), they are forced

into a hostile environment (sand) to disperse to any area that can afford them

shelter to survive. Many succumb to predators. In this habitat the ‘phantom

population’ is almost theoretical. Maturing males of these isolated rocks are

virtually dependent on the death of their adults (particularly their father) if

they are to survive and establish their own territory. The pressure on maturing

juvenile females is less, but the adult female (their mother) will evict them if they

show interest in their father. For those on the large koppies this pressure on

newly matured cordylids is less intense.

The spatial relationships of American iguanids have been studied most,

and good reviews are those of Carpenter (1967), Rand (1967), Ruibal (1967)

and Mayhew (1968). Most lizards have a 50:50 sex ratio.

Of Old World lizards Harris (1964) reports a density of Agama agama

36 per acre (90,0 per hectare). Burrage (1973) found the population density of

the mesic Chamaeleo pumilus to vary from 12 adults per hectare in brushy areas

to 75-200 adults per hectare on reeds surrounding still bodies of water, with

males averaging 43°% of the population. However, population density in

arboreal forms (e.g. Anolis and Chamaeleo pumilus) is three-dimensional, not

in plan as with ground-dwelling species. The ubiquitous, ground-dwelling,

deserticulous C. namaquensis is most numerous in topographically varied areas,

with a mean annual density of 51,8 adults per hectare, with a sex ratio slightly

favouring females.

Of New World lizards, population density in some, e.g. Uta stansburiana

and related insular forms of Uta (Soulé 1964); U.s. hesperis, stejnegeri and elegans

(Burrage 1966); U. s. stansburiana and stejnegeri (Tinkle 1961, 1967; Tinkle,

McGregor & Dana 1962; Tinkle & Woodard 1967), varies from an average

inland density of approximately 25,5 territory-holding adult utas per acre

(approximately 63,8 per hectare) to littoral and insular populations two to three

times as dense. In some unusual conditions some mainland Uta stansburiana

populations may be as much as 700 per acre (1 750 per hectare) in Utah

(Knowlton, Fronk, & Maddock 1942); in California and Mexico (Burrage

1966); in Colorado (Ferguson, pers. commun.)). Burrage (1966) found that the.

population density of U. s. hesperis varies according to the topography and to a

lesser degree the plant density of the particular locales studied. U. s. hesperis

population densities vary from 20,0 to 750,0 (x = 179,5) adults per hectare in

areas altered (e.g. steeply graded, or terraced, a greatly reduced plant density of

usually uniform type, chiefly weeds in the early stages of plant succsssion) by

human activity. On the undisturbed backshore of topographically rugged or

varied beaches with a low plant density there are 55,0 to 62,5 (k = 59,0) adult

utas per hectare, 37,5 adult utas per hectare in beach canyons, and 30,0 to 47,5

(X = 32,5) adult uta per hectare in inland areas. Beach areas are topographically

varied and intricately eroded, allowing a higher population density than can

10 ANNALS OF THE SOUTH AFRICAN MUSEUM

be realized on flat terrain. Disturbed areas and beaches have scattered plants

in which are abundant prey concentrations. Tinkle (1967) found that U. s.

stejnegeri in Texas concentrates its activity around Neotoma (wood rat) nests—

a rich source of arthropod prey—which are sited in Prosopis (Mesquite) clumps.

Jorgensen & Tanner’s (1963) Nevada data suggest that the prey of the Ufa s.

stansburiana is more dispersed, and/or the topography more uniform, the plants

more dispersed, and so are the lizards.

Very few tropical lizards have been studied, but Heatwole (1967) mentions

Anolis limifrons having a population density of 400 per acre (approximately

1 000 per hectare) and refers to Turner & Guest finding densities of >600 per

acre (approximately >1 500 per hectare) for Anolis gunlachi, a medium-sized

anole of Puerto Rican montane rain forests. Cagle (1946) reports a density of

500 adult Hemidactylus (Gekkonidae) in a 2 400 ft? (approximately 223,2 m?)

area on Tinian, so assuming an even distribution this gives 10000 per acre

(25 000 per hectare). Hirth (1963) studied the ecology of a teiid (Ameiva

quadrilineata) and an iguanid Basiliscus vittatus on a Costa Rican beach. He

found the density of juveniles of the former species varies from 12,6 to 24,2

per hectare, and the adults from 17,1 to 39,1 per hectare, the combined numbers

suggesting a te1id population of 29,7 to 63,9 per hectare. Juveniles of B. vittatus

vary from 13,8 to 27,9 per hectare. The adults of this species are only 5,2 to 10,9

per hectare, owing to the fact that they did not especially frequent the study area

and do not compete with their juveniles. Actually, adult B. vittatus density

would be at least double. For B. basiliscus in the Panama Canal Zone, Barden

(1943) estimates a population density of one lizard per 15,2 m of shoreline.

Regarding spatial organization of lizards, some authors confuse the issue

by using ‘defended home range’ instead of ‘territory’, justifying such discon-

certing usage because the home range of a particular species is also its territory.

While a ‘defended home range’ is synonymous with ‘territory’, the same authors

omit the word ‘defended’ in their discussion, when it should be included. Since

in some parts of their distribution the same species can also possess a territory

(defended) and a home range (undefended), it is too cumbersome to speak both

of a ‘defended home range’ and a ‘home range’. Needless confusion results

from those who use only home range when obviously territory is the correct

term. Thus, I feel use of ‘defended home range’ should be dropped in favour of

‘territory’, and in no case should ‘home range’ be used for any area defended

by any species. Table 4 gives the mean territory and/or home range sizes of

some of the lizards reported in the literature.

Discussions of spatial occupancy in lizards are disconcerting, because of

varied techniques employed, consideration of widely separate populations in

different habitats, physical factors (varied geologic substrates, local topography),

and biotic factors (e.g. population density of the studied species, competition,

plant density). Few attempts have been made to study adjacent populations of

the same species in the same general area under slightly different conditions of

local topography (e.g. relief, slope angle), types of geologic substrates, plant

AGAMA ATRA AND C. C. CORDYLUS IN THE VICINITY OF DE KELDERS, C.P. 11

Table 4

Review of mean territory and home range sizes of some of the lizards reported in the literature.

Species

Amblyrhynchus

cristatus

Ameiva quadrilineata

Anolis sagrei

Basiliscus vittatus

Chamaeleo

namaquensis

Chamaeleo pardalis

Chamaeleo pumilus

Cnemidophorus

hyperythrus

Cnemidophorus tigris

Ctenosaura pectinata

Sceloporus olivaceus

Uta stansburiana

hesperis

Uta stansburiana

hesperis

Uta stansburiana

stansburiana

Uta stansburiana

Stejnegeri

Sex | Age

3d adult

ole) adult

3d juvenile

92 juvenile

3d adult

eye) adult

3d adult

6d juvenile

22 ‘juvenile

3d adult

ele) adult

35+ 22 juvenile

3d adult

ele} adult

eye) adult

3d adult

ele} adult

$3 +29 juvenile

3d adult

ele) adult

S6d+29 adult

33 juvenile

QQ juvenile

3d adult

ole) adult

33 juvenile

22 juvenile

3d adult

ole) adult

3d adult

oho) adult

3d adult

ele) adult

365+ 29 juvenile

feito adult

ele) adult

33d adult

eke) adult

Area occupied

Acres Metres?

= 1,0

— 16,4

— 1551

= Aili

— 1Be7/

— 36,0

— 14,9

== 12,4

— 12,0

— 122

— 2-5

— Ales

— 1 250,2

— 382,0

— 867,9

Several 10m?

— 600,0

0,07 283,3

0,10 404,7

0,09 364,2

050) 210235

0,24 O7le3

== 2

0,027 109,3

0,017 68,8

0,17 687,9

0,07 283,3

0,005 20,2

0,004 16,2

0,05 202,4

0,034 137,6

0,075 303,4

0,056 226,6

0,021 85,0

0,016 64,8

0,04 161,9

0,10 404,7

0,03 eA

0,11 444.2

0,03 121,4

Remarks

Only in

breeding

season

Only displays

Coastal

Inland

Coastal

Inland

California

Calif. coastal

Calif. inland

Intertidal

home range of

stranddwellers

Nevada

Texas

Source

Carpenter (1967)

Hirth (1963)

Evans (1938)

Hirth (1963)

Burrage (1973)

Bourgat (1968)

Burrage (1973)

Bostic (1964)

Jorgensen &

Tanner (1963)

Jorgensen &

Tanner (1963)

Jorgensen &

Tanner (1963)

Evans (1951)

Blair (1960)

Burrage (1966)

Jorgensen &

Tanner (1963)

Jorgensen &

Tanner (1963)

Jorgensen &

Tanner (1963)

Tinkle (1967)

12 ANNALS OF THE SOUTH AFRICAN MUSEUM

density, and lizard population density. Burrage (1966) conducted a five-year

investigation of spatial occupancy in Uta stansburiana hesperis populations in

southern California at selected, proximal study stations of varied location and

topography (e.g. coastal bluffs, sandy and rocky beaches, coastal and inland

canyons, rugged and flat topography), geologic substrates, plant density and

populaton density. He found widely different territory sizes and social organiza-

tions obtaining at each study station to a degree reported as ‘standard’ for this

species by Jorgensen & Tanner (1963) at one site in Nevada, and Tinkle,

McGregor & Dana (1962) in an area of Texas. Thus, environmental factors in

widely separate areas could account for an actual difference in territory size

(see Table 4) rather than computation error (Jorgensen & Tanner used density

probability function, obtaining larger territory sizes than Tinkle et a/. using the

minimum polygon method). Furthermore in Texas, utas concentrate their

activity around Neotoma nests in Prosopis clumps and, thus, have less need to

wander for food than the utas in Nevada, probably resulting in smaller Texas

Uta territories than those in Nevada.

Furthermore, as Tinkle (1967) observes, there are few studies based on

large numbers of captures. Also, despite the studies demonstrating territorial

behaviour in lizards (mostly temperate iguanids), there has been little speculation

as to its adaptive significance in reptiles, as has been done by Nice (1941),

Hinde (1956) and Carpenter (1958) for other animals, mostly birds. Rand (1967)

has made a step in this direction, based largely on his observations of the iguanid

Anolis lineatopus on Jamaica. Hypothetical values of territory can be: (1)

securing a requisite share of environmental resources, and/or (2) mating, and/or

(3) survival of offspring. Rand reports that the critical environmental resources

for A. lineatopus need be defended intra- and interspecifically. Food is the

limiting resource for other lizards, such as Sceloporus merriami (Milstead 1961)

and in such cases both sexes defend.

Defence only of the resting site is noted (Knowlton ef a/. 1942; Burrage

1966) in Uta stansburiana populations too large to permit or necessitate terri-

torial establishment; as areas with abundant and/or concentrated prey. U. s.

hesperis maintains territories in the backshore areas of beaches, but has only

home ranges intertidally, where the variety and numbers of food are greater.

There are paltry data on spatial organizations of Old World lizards. Harris

(1964) gives territory sizes of 600—900 ft? (58,8-84,6 m?) for male Agama agama

in population densities of 36 adults per acre (90 per ha). Harris noted that

dominant females never leave dominant males. Several adult females occurred

with each dominant male, one female being dominant and ‘true’ to the male.

Subdominant females will desert. All juvenile males leave the area they occupy

as hatchlings, but young females may remain.

Bourgat (1968) gives an area of ‘several 10 m®’ for the home range of

Chamaeleo pardalis on Reunion. Burrage (1973) found that C. pumilus has a

home range of 10 m? in plan, but since the space is three-dimensional, the actual

area covered is closer to 600 m?. C. pumilus only defends its night-time resting

AGAMA ATRA AND C. C. CORDYLUS IN THE VICINITY OF DE KELDERS, C.P. 13

perch, the security of which is important to this species. This observation is true

of other arboreal chamaeleons (Brain 1961; Bustard 1958, 1965, 1966, 1967;

Von Frisch 1962; Spence 1966).

In the ground-dwelling, deserticulous C. namaquensis, most territories

embracing different habitats are larger than those limited to one. The largest

male territory was 8 000 m?; the largest female territory was 1 632 m?. Territory

size in this species varies according to reproductive demands, with male terri-

tories increasing in area during courting and those of females during egg-laying.

Male territories, however, are always larger than those of females and were

surrounded by more smaller female territories than they bordered on those of

other males. As with other chamaeleons, C. namaquensis has a more or less

centrally located resting-area, usually in a redoubt. Unlike most lizards, C.

namaquensis territories do not have areas of greater or lesser usage. On waking

in the morning C. namaquensis patrols to the border limits and encompasses

its entire domain. With the passage of time, C. namaquensis patrols its territories

in decreasing circuits inwards to the rest area. The border configuration of

C. namaquensis territories is especially interesting, for finger-like projections of

the territory of one chamaeleon intrude into that of another. In areas of micro-

relief, as hummocks, such projections closely followed the high ground, but this

curious system prevailed even on monotonously flat areas, and especially on

dunes, where one would suspect territories to be circular or squarish, since

there was no micro-relief or landmarks to regulate border configuration. Main-

tenance of territories in C. namaquensis assured that any one male could meet

several females during the normal diel patrol of both sexes. Also, vigorous

territorial defence by the female C. namaquensis would prevent exhumation of

the defending chamaeleon’s buried eggs by other females seeking to build a

nest. Each female C. namaquensis constructs its nest only within the borders of

its territory.

In the case of Agama atra and Cordylus cordylus on coastal and inland

rocky outcrops, territorial establishment is best interpreted as enforcing a

spacing of the population for the most efficient use of available resources. Closer

to the sea, there is a reduction of available area, but an increase in the numbers

and variety of prey favours a trend to despotism in the slightly larger local

population of Agama. ;

Cordylus cordylus in the inner tiers of the rock koppies has populations too

large to establish or maintain territories successfully and shelter is not at a

premium and possibly also food. This is not so for those living on isolated rocks

in sandy areas. Here, suitable habitat and retreats, and possibly food, are at a

premium, and each family group vigorously defends its territory from intrusion

by others. This, however, should not obscure the possibility that the cordylids

are limited only to the rocky situations because suitable or sufficient supplies

of their food are.

I have fully discussed the relationship of juvenile cordylids to their adults.

Freshly-hatched agamids of both sexes initially site within the territories of

14 ANNALS OF THE SOUTH AFRICAN MUSEUM

their parents. A few weeks after hatching, juvenile males occupy a shifting

home range, gradually dispersing out and away from their parents. They are

not free from challenge by adults whose territories they transgress, but are

neither evicted. Juvenile females are not free from challenge from the adults,

but it seems more recognitory and they are not evicted. Harris (1964) made

similar observations for juvenile agamids.

Burrage (1973) reports that the juveniles of the ground-dwelling Chamaeleo

namaquensis occupy shifting home ranges (10-53 m?; x = 27,5) occurring

wholly or partly within the strongly defended territories of the adults (initially,

that of their parents), from whom they are free of challenge. As the juveniles

of C. namaquensis matured, they began defending their area as they stabilized

its location. By shifting an undefended space when juvenile and free of challenges,

they might find an unoccupied area they can defend in adulthood. Juveniles

always slept on a raised object, such as a twig, and not necessarily the same object

each night, and companionably together. Juveniles of C. pumilus have no regular

sleeping area in the prolonged temporal sense and a continuously shifting home

range. Resting C. pumilus juveniles are well spaced on twigs or grasses and are

hostile to any other chamaeleon being near them. Even new-born young execute

vigorous displays. Only captive C. namaquensis juveniles displayed and evinced

ownership in the presence of food. A sort of ‘dog-in-the-manger’ affair, where

one would be eating in the food dish and vigorously prevent any other chamae-

leon(s) from sharing the meal.

In most cases juvenile lizards have smaller territories or home ranges than

adults. The exceptions, as Hirth (1963) reports for Ameiva and Basiliscus, are

in Tinkle’s (1967) view due to calling a home range a composite of areas occupied

by a young animal as it disperses, and it is not the same as a stabilized home

range of an adult. This may be so, as if, for example, I had called as stable home

ranges the total area occupied by the shifting home ranges of C. namaquensis

juveniles as they dispersed, the area would surpass the territories of the adults.

That a juvenile could or would maintain a larger area than an adult seems quite

unlikely. However, I found that the Californian anguids Gerrhonotus multi-

carinatus webbi and G. multicarinatus nanus occupy a shifting home range as

juveniles and adults. Obviously, this problem needs more thorough investiga-

tion. Usually, the role of territories and home ranges in juveniles of a species

is overlooked, which is a pity, because understanding spatial relationships at

all ages in a given species population gives a far better understanding of the

nature of such arrangements to any species.

It is perhaps a mistake to say juveniles are non-territorial, and it would be

well to study when juveniles set up territories (see also Blair 1960) with increasing

maturity. Burrage (1966) found that juvenile Uta stansburiana hesperis had

free run of adult territories and was not challenged by the adults and did not

exhibit any spatial ownership to each other. Most juveniles of U. s. hesperis

lived on the domains of adults who were usually their parents; this juvenile to

adult association appears to be a loose family unit of sorts. This agrees with

AGAMA ATRA AND C. C. CORDYLUS IN THE VICINITY OF DE KELDERS, C.P. 15

Hunsaker’s (1962) report that Sceloporus, less than 50°%% of adult size, was free

from challenges. In locales with denser populations, as beach stations, juveniles

of U. s. hesperis were not usually found to occupy discrete territories, except

when a vacancy was created by the death of an adult. For example, at Pacific

Beach Bluffs (adult population density 130 per hectare), only four juvenile

females occupied territories ranging from 4,1 to 12,1 m* (kK = 8,1 m?). At

Alvarado Canyon, a less densely populated (adult population density 62,5 per

hectare) locale, 17 juvenile male territories (4,2—28,3 m?; x = 12,1 m?) and

13 juvenile female territories (4,1-16,2 m*; X = 8,1 m?) were present. While a

lesser population density favours establishment of juvenile territories in this

species, it does not, in itself, guarantee it. Juvenile territories were always located

in a redoubt, compact and easy to defend. At Lower Tourmaline Canyon, a

beach canyon, with many such redoubts (depression drainage pits associated

with the weathering of a particular geological formation), allowed many juveniles

to maintain territories, though the adult population density was 75 per hectare.

This station had the second greatest number of juvenile territories observed

for this species. Seven juvenile male territories ranged from 4,1 to 52,6 m?

(k = 28,3 m?) and five juvenile female territories from 4,1 to 24,3 m?

atm).

The territory of any given beach strand-dwelling adult male (N=410) and

adult female (N=450) U. s. hesperis consists of a small nucleus area

(gg X = 20,2 m?; 92 x = 12,9 m?), where the lizard has its burrow and con-

centrates its activity; surrounding this nucleus area is a larger defended area

(gd k = 178,1 m?; 99 X = 105,2 m?), with a mean intertidal ‘free’ zone (home

range) of an additional 85,0 m? in males; 72,8 m? in females. Non-strand utas

lacked the home range and are territorial, though it is subdivided as above. By

temporally studying establishment of spatial occupancy in this species, the

nucleus area of the adult territory is seen to be that area the occupying lizard

can initially successfully defend when a maturing, older juvenile. Over this

nucleus the maturing juvenile exercises ownership of and is active over an

increasingly wider area. It is more selective to success (of territorial establish-

ment) and survival if the lizard initially defends a small, compact area in a

redoubt (i.e. stone rubble, thorny xeric plants, or both, etc.) in which it has a

retreat, rather than attempts to dominate a larger adult-sized area lacking or.

devoid of shelter. Juvenile U. s. hesperis only begins defending when (1) there

is space available, and/or (2) such space is readily defended, as a small compact

redoubt, and/or (3) the overall, more importantly, the adult population density

is low. As it is more critical for adults in this species to maintain territories,

the establishment of territories in this species is a virtual ‘right’ of adulthood.

This is indicated by the dearth of juvenile U. s. hesperis territories, and the

limited conditions in which they occur. The fact that U. s. hesperis juveniles are

free of challenge tends to select against the establishment of juvenile territories.

In U. s. hesperis there is also a ‘phantom population’ as I have described for

Agama atra and Cordylus cordylus.

16 ANNALS OF THE SOUTH AFRICAN MUSEUM

While territorial formation is common in iguanids and some agamids

(Carpenter 1967), it does not occur in some, as the iguanid Phrynosoma (Lynn

1965). Iguanids, agamids and chamaeleonids display ritualistically in species-

specific patterns, which are a type of aggression and the chief determinants of

social structure. Such display apparently functions in territorial declaration,

which is a type of space dominance. Territoriality decreases intraspecific stress

by stabilizing the social organization of the population and dispersing the

species population, especially the males throughout the available suitable

habitat (Carpenter 1967).

DISPLAY PATTERNS

Both sexes of Agama atra and Cordylus cordylus display. A weak display

is given in a sitting position with the front of the body raised. In A. atra intense

displays are given with the body raised on all fours and with most of the tail

held off the ground. Two display types can be recognized. Courtship display

consists of three sharply peaking high-level bobs each lasting about 1,2 seconds,

followed by two very low-level bobs, each lasting about 0,5 of a second. The

entire set takes about 4,0 seconds. Two or three such sets are normally delivered,

with an interval of 2,0 to 6,0 seconds between each. The female responds to the

male’s display by presenting to him in a manner facilitating copulation, and

some females seem to instigate it. Territorial display consists of two steeply

peaking high-level bobs, as in courtship display, followed by two different bobs

executed from the intermediate to the high-level range. The first of these lasts

about 0,7 of a second, and the last for almost a second.

Courting and territorial display of Cordylus cordylus consists of a set of

three intermediate-level bobs, each of 1 second duration. However, the delivery

and body positioning determines whether the bob is for territorial or courting.

In territorial display the defender, from its vantage point, delivers head down to

the transgressor. The hind quarters are held to the substrate, but the fore part

of the body is alternately raised and lowered, thus giving the delivery the appear-

ance of ‘push-ups’. While Agama display also can resemble ‘push-ups’, it is not

so pronounced as with Cordylus. Three to four sets are normally given with a

5-second interval between each, though intensity depends on how deeply the

intruder has violated the territory of the defender. In courting the display is

given by both sexes facing each other laterally with the entire body and usually

the tail held free of the substrate. During such display both sexes may feign

biting the other before copulation commences and the number of sets given and

the interval between each varies with the energy of individual pairs.

SOCIAL INTERACTIONS

On the coastal cliffs and inland rocky outcrops, Cordylus cordylus and

Agama atra display is entirely intraspecific, and a C. cordylus territory may

‘include’ partly or wholly that of an Agama, even a despot. However, on the

isolated rocky situations in sandy areas, only geckos apparently co-inhabit with

AGAMA ATRA AND C. C. CORDYLUS IN THE VICINITY OF DE KELDERS, C.P. 13,

C. cordylus. It is difficult to know why Agama, and for that matter other lizards,

seem absent. It is possible that other lizards have not been able to cross the

sandy expanses to populate the rocky areas, but then C. cordylus with its

preference for rocks would seem as effectively excluded from this habitat as

A. atra and other lizards. Perhaps the high density of C. cordylus in such situa-

tions has effectively precluded the occupation of it by other saxicolous lizards.

A. atra females of one male’s group seldom engaged in territorial disputes

with each other, but frequently did so with females and occasionally even males

of other groups. However, it is the prime responsibility of each male to maintain

the territorial integrity of his group. When two females of different groups had

a border dispute the respective males charged down, and if both males stayed

on their own territories, the dispute was quelled. If, however, one or both males

strayed over the border, much frenzied displaying resulted and on rare occasions

vicious ‘free-for-all’ fighting. Incursions by members of despot groups are

tolerated by non-despots, though their incursions are initially weakly challenged,

followed by a submissive posture. The integrity of the retreat of any individual

group is, however, viciously defended from but usually respected by an intruding

despot group member. Despot group members were involved in fewer disputes,

proportionately, among themselves than were non-despots.

The observed social interactions of 57 groups were studied in detail and

are given in Table 5. While most matings are intra-group, despot males occa-

sionally courted females of surrounding non-despot males. Mating between

despot and non-despot group members occurred rarely, usually involving sub-

dominant despot females and non-despot males, when the despot male was not

looking.

Both sexes of Cordylus cordylus on coastal cliffs and on inland rock outcrops

maintain individually discrete territories, though those of males are larger and

overlap those of the females. There is no group formation as is the case of the

agamids. Thus, both sexes defend their territories against and by cordylids of

either sex. During the reproductive season, while challenging intruding members

of the other sex, each sex is more tolerant of such intrusion. On the other hand,

they defend more aggressively against members of their own sex.

C. cordylus occurs in groups on isolated koppies and scattered rocks in

sandy areas. In those maintaining only home ranges, there is a definite home -

range, and each group tolerates transient incursion by the surrounding groups.

It should be noted that such incursions were observed only very rarely. Each

group’s retreat is, however, strongly defended. In those groups with a territory

and home range a group would tolerate incursion of the area called home range,

but not the territory.

Those maintaining territories, with or without home ranges, would brook

no incursion of their territories. Both adult males and females defend singly or

together, occasionally with some assistance from the juveniles, particularly the

older ones.

Display patterns and social structure of lizards are amply reviewed by

ANNALS OF THE SOUTH AFRICAN MUSEUM

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AGAMA ATRA AND C. C. CORDYLUS IN THE VICINITY OF DE KELDERS, C.P. 19

Carpenter (1962, 1967), Evans (1951), Hunsaker (1962), Hunsaker & Burrage

(1969), and Ruibal (1967), and are virtually limited to American iguanids,

though there have been some studies on agamids (Carpenter unpubl.:; Cowles

1956; Greenberg 1957; Hairston 1957; Harris 1964; Mayhew 1963; Schmidt

1935). It appears that display patterns and social organization are similar

between agamids, and their close relatives the iguanids. Chamaeleonid —an

Ascalabotan family more closely related to the agamids—display has been

studied by Brain (1961), Burrage (1973), Bustard (1958, 1965, 1966, 1967),

Schmidt & Inger (1965), Spence (1966) and Von Frisch (1962).

For territorial, social and spatial behaviour, Carpenter (1967) differentiates

between ‘assertion display’ and ‘full or challenge display’. The former is a

‘showing the flag’, a weak or low intensity performance warning other lizards

to ‘keep their distance’. The challenge display is a high-intensity performance by

very aggressive adults. As Carpenter observes, even freshly hatched and new-

born young execute head-bobs and push-ups and I have discussed this on

pages 13-16.

The following terms are applied by Carpenter to lizard behaviour patterns:

individual space, territory, despotism, hierarchy, activity range (home range),

and various levels of social organization, noting that those iguanids with the

most elaborate aggression displays have the most complex social organizations.

As Carpenter uses the term, social organization ‘. . . results from the inter-

actions of members of the same species which influence or tend to establish

patterns of space occupancy and inter-individual response’. These patterns may

be rigid or transient; can seasonally vary.

Lizard display is considered to be intraspecific, but most lizards deliver at

least the assertion display to all intruders. Depending on highly competitive

areas and very critical resources, the more aggressive challenge display is only

intraspecific. Courtship display favours a response only from members of the

same species, especially of the opposite sex, for which Noble & Bradley (1933)

consider it selects.

Fighting in wild populations of lizards is rare, but has been recorded for

the iguanids Sceloporus grammicus (Evans 1946) and Uta stansburiana (Burrage

1966; Tinkle 1967). Some aspects of social behaviour may be different in

captivity, with an increase or decrease in antagonistic behaviour (Burrage 1973, -

as regards chamaeleons). In a multi-species assemblage of captive iguanids,

Hunsaker & Burrage (1969) found a shift from territories to a social hierarchy,

with a development of despotism, as a result of population pressure and reduc-

tion of the available area; which hierarchy is established by increasingly vicious

fighting, rather than the normal displays, as population pressure increases,

and/or the available area is reduced. Similar social hierarchies, dominated by a

despot male, have been recorded by Evans (1951) in wild populations of the

normally territorial Crenosaura pectinata, where the population was crowded

around a rich source of food. They were crowded because their natura] habitat

had been burnt over. Their adaptation to living close to the food supply involved

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

a reduction of available space for refuge and basking and a subsequent shift

from the territorial system. Burrage (1966) in California, Ferguson (pers.

commun.) in Colorado, and Knowlton et al. (1942) in Nevada, noted a some-

what similar arrangement in inland situations with a high Uta stansburiana

population density and/or small area, whereby this normally territorial species

merely defended only the burrow and had a small home range. Soule (1964)

noted this same phenomenon in high density insular and littoral populations of

Uta, as Burrage (1966) also observed. Burrage (1966) notes that strand-dwelling

U. s. hesperis possesses hotly defended territories on the backshore and home

ranges seawards to at least the mean high tide limit. Clarke (1965) studied several

iguanids, demonstrating a formation of dominant-subdominant hierarchies.

Harris (1964) gives a detailed account of social behaviour in Agama agama,

also noting formation of social hierarchies in high population densities. Harris

notes the establishment of territorial borders in this species (see remarks on the

litigated area between Cordylus Groups 1 and 2 on page 6), where two males

often fought over a path between their boundaries. Fighting decreased as the

border became established and eventually display along the path replaced

combat, with rarer attempts at violation. Establishment of social hierarchies in

high population densities limits the frequency of antagonistic behaviour that

would result in attempts at territory formation. Also, in high population den-

sities, the territory size would be reduced and the increasing rate would be high,

necessitating almost continual defence on the part of the respective defenders,

with insufficient or no time for other activities. It seems, however, that on

topographically varied areas territorial establishment and high population

densities are possible (Burrage 1966, and pages 4-5).

As Carpenter (1967) notes, it appears that most iguanids are territorial, but

details of territories are not known for the vast majority of these. The data

suggest territorial formation in agamids and some chamaeleonids, but few of

these groups, especially chamaeleons, have been studied to the degree that

iguanids have.

Our knowledge of social behaviour in reptiles is poor, and sometimes

confusing, which is perhaps a pity since we could then better understand social

behaviour patterns in the higher vertebrates.

SUMMARY

The spatial and social organizations are described of Agama atra and

Cordylus cordylus cordylus inhabiting coastal cliffs, inland rock outcrops and

dune and sandy situations. A total of 1 030 A. atra and | 459 C. cordylus were

marked. The population of A. atra is denser along the coastal cliffs and least

dense on inland rock outcrops. A. atra is absent in suitable habitats surrounded

by very dense scrub or large expanses of sand. C. cordylus occurs in all the

habitats studied, as long as there are some rocks, which are the focal point of

its activity and in which these lizards have their retreats. C. cordylus is densest

AGAMA ATRA AND C. C. CORDYLUS IN THE VICINITY OF DE KELDERS, C.P. 21

on the terraces of large rock koppies surrounded by sand and least dense in

areas of isolated rocks among sand.

Territories and home ranges were computed only on the basis of 10 or

more recaptures. A. atra only possesses territories. Adult population density

varies inversely with territory size. Male territories are always the largest and

contain several smaller female territories. The dominant female is always ‘loyal’

to the male. Towards the edge of the water there is a tendency to despotism, the

territories of despots always being the largest. Juveniles are generally not

challenged. Male juveniles leave the area they occupy as hatchlings, but females

may remain. Despot groups were involved in proportionately less territorial

challenges than non-despot groups. Male despots had more sexual contact with

non-despot females than despot females had with non-despot males.

On the periphery of the larger koppies family groups of C. cordylus main-

tain a territory and a home range on the koppie itself. The larger the area and

the lower the population density, the larger are the areas of the home range and

the territories. In any situation and regardless of the spatial organization, the

retreat burrow is defended. In sandy situations on isolated rocks only territorial

organization occurs, with each territory being occupied by an adult male and

female along with their juveniles of varied age groups. All members of each

group defend the territory. The largest Cordylus territories occur on isolated

rocks among sand. An experimental removal of a group of cordylids is given,

showing territorial adjustments over a long period. Juvenile cordylids quit or

are evicted from the territory of the ‘family group’ as they mature, or in the

event of the death of their parents succeed to the adults’ territory.

REFERENCES

Acocks, J. P. H. 1953. Veld types of South Africa. Mem. bot. Surv. S. Afr. 28: 1-192.

BARDEN, A. 1943. Notes on the basilisk at Barro Colorado Island, Canal Zone. Ecology 24:

407-408.

Biair, W. F. 1960. The rusty lizard. A population study. Austin: University of Texas Press.

BourGat, R. 1968. Etude des variations annuelles de la densité de population de Chamaeleo

pardalis Cuv., 1892 dans son biotope de I’[le de la Réunion. Vie Milieu (C) 19: 227-231.

Bostic, D. L. 1964. The ecology and behaviour of Cnemidophorus hyperythrus beldingi Cope

(Sauria: Teiidae). Unpubl. M.Sc. thesis, San Diego (California) State College.

BRAIN, C. K. 1961. Chamaeleo dilepis—a study on its biology and behaviour. J. herpetol. Ass.

Rhod. 15: 15-20.

BuRRAGE, B. R. 1966. The natural history of the western ground uta, Uta stansburiana hesperis

Richardson (Sauria: Iguanidae). Unpubl. M.Sc. thesis San Diego (California) State

College.

BurRRAGE, B. R. 1973. Comparative ecology and behaviour of Chamaeleo pumilus pumilus

(Gmelin) and C. namaquensis A. Smith (Sauria: Chamaeleonidae). Ann. S. Afr. Mus. 61:

1-158.

BUSTARD, H. R. 1958. Use of horns by Chamaeleo jacksoni. Br. J. Herpet. 2: 105-107.

BusTARD, H. R. 1965. Observations on the life history and behavior of Chamaeleo hohnelii

(Steindachner). Copeia 1965: 401-410.

BUSTARD, H. R. 1966. Observations on the life history and behavior of Chamaeleo bitaeniatus

Fischer. Herpetologica 22: 13-23.

BUSTARD, H. R. 1967. The comparative behavior of chameleons: fight behavior in Chamaeleo

gracilis Hallowell. Herpetologica 23: 44—-S0.

ANNALS OF THE SOUTH AFRICAN MUSEUM

CAGLE, F. R. 1946. A lizard population on Tinian. Copeia 1946: 4-9.

CARPENTER, C. C. 1962. A comparison of the patterns of display of Urosaurus, Uta and

Streptosaurus. Herpetologica 18: 145-152.

CARPENTER, C. C. 1967. Aggression and social structure in iguanid lizards. Jn: MILSTEAD, W. W.,

ed. Lizard ecology; a symposium: 87-105. Columbia: University of Missouri Press.

CARPENTER, C. R. 1958. Territoriality: a review of concepts and problems. Jn: Rog, A. &

Simpson, G. G. eds. Behavior and evolution: 224-250. New Haven: Yale University Press.

CLARKE, R. F. 1965. An ethological study of the iguanid lizard genera Callisaurus, Cophosaurus

and Holbrookia. Emporia St. Res. Stud. 13: 1-66.

Cow Les, R. B. 1956. Notes on the natural history of a South African agamid lizard. Herpetolo-

gica 12: 297-302.

Evans, L. T. 1938. Cuban field studies on territoriality of the lizard, Anolis sagrei. J. comp.

Psychol, 25: 97-125.

Evans, L. T. 1946. Social behavior of the lizard, Sceloporus grammicus microlepidotus. Anat.

Rec. 94: 53-54.

Evans, L. T. 1951. Field study of the social behavior of the black lizard, Ctenosaura pectinata.

Am. Mus. Novit. 1493: 1-26.

Firzsimons, V. F. 1943. The lizards of South Africa. Trans. Mus. Mem. 1: 1-528.

FrRIscu, O. von. 1962. Zur Biologie des Zwergchamaleons (Microsaurus pumilus). Z. Tierpsychol.

19: 276-289.

GREENBERG, B. 1957. Behavior of iguanid and agamid lizards. Yb. Am. phil. Soc. 1957: 251-253.

Hairston, N. G. 1957. Observations on the behavior of Draco volans in the Philippines.

Copeia 1957: 262-265.

Harris, V. A. 1964. The life of the rainbow lizard. London: Hutchinson.

HEATWOLE, H. F. 1967. Jn: MILSTEAD, W. W., ed. Lizard ecology; a symposium: 62. Columbia:

University of Missouri Press.

HINpDE, R. A. 1956. The biological significance of the territories of birds. /bis 98: 340-369.

HirtuH, H. R. 1963. Ecology of two lizards on a tropical beach. Ecol. Monogr. 33: 83-112.

HunsSAKER, D. 1962. Ethological isolating mechanisms in the Sceloporus torquatus group of

lizards. Evolution 16: 62-74.

HunsSAKER, D. & BURRAGE, B. R. 1969. The significance of interspecific social dominance in

iguanid lizards. Am. Midl. Nat. 81: 500-511.

JORGENSEN, C. D. & TANNER, W. W. 1963. The application of the density probability function

to determine the home ranges of Uta stansburiana stansburiana and Cnemidophorus tigris

tigris. Herpetologica 19: 105-115.

KNOWLTON, G. F., FRONK, W. D. & MAppbock, D. R. 1942. Range lizards as insect predators.

J. econ, Ent. 35: 942.

Lynn, R. T. 1965. A comparative study of display behavior in Phrynosoma (Iguanidae).

SWest. Nat. 10: 25-30.

MAYHEW, W. W. 1963. Observations on captive Amphibolurus pictus, an Australian agamid

lizard. Herpetologica 19: 81-88.

MAYHEW, W. W. 1968. Biology of desert amphibians and reptiles. Jn; BRown, G. W., ed.

Desert biology: special topics on the physical and biological aspects of arid regions. 1:

195-356. New York, London: Academic Press.

MILSTEAD, W. W. 1961. Competitive relations in lizard populations. Jn: BLAtr, W. F., ed.

Vertebrate speciation: 460-489. Austin: University of Texas Press.

Nice, M. M. 1941. The role of territory in bird life. Am. Midl. Nat. 26: 441-487.

Nose, C. K. & BRADLEY, H. T. 1933. The mating behavior of lizards; its bearing on the

theory of sexual selection. Ann. N.Y. Acad. Sci. 35: 25-100.

RAND, A. S. 1967. The adaptive significance of territoriality in iguanid lizards. Jn: MILSTEAD,

W. W. ed. Lizard ecology: a symposium: 106-115. Columbia: University of Missouri Press.

RuIBAL, R. 1967. Evolution and behavior in West Indian anoles. Jn: MILSTEAD, W. W.., ed.

Lizard ecology: a symposium: 116-140. Columbia: University of Missouri Press.

ScHMIDT, K. P. 1935. Notes on the breeding behavior of lizards. Publs Field Mus. Nat. Hist.

(Zool.) 20: 71-76.

SCHMIDT, K. P. & INGER, R. F. 1965. Living reptiles of the world. London: Hamilton.

AGAMA ATRA AND C. C. CORDYLUS IN THE VICINITY OF DE KELDERS, C.P. 23

SouLE, M. 1964. Evolution and population phenetics of the side-blotched lizards (Uta

stansburiana and its relatives) on the islands in the Gulf of California, Mexico. Unpubl.

Ph.D. dissertation, Stanford University.

SPENCE, J. M. 1966. Observations on the Damara Chameleon Microsaura damarana Boulenger.

Ann. Cape prov. Mus. 5: 145-148.

TINKLE, D. W. 1961. Population structure and reproduction in the lizard Uta stansburiana

stejnegeri. Am. Midl. Nat. 66: 206-234.

TINKLE, D. W. 1967. The life and demography of the side-blotched lizard, Uta stansburiana.

Misc. Publs Mus. Zool. Univ. Mich. 132: 1-182.

TINKLE, D. W., McGrecor, D. & DANA, S. 1962. Home range ecology of Uta stansburiana

stejnegeri. Ecology 43: 223-229.

TINKLE, D. W. & WooparpD, D. W. 1967. Relative movements of lizards in natural populations

as determined from recapture radii. Ecology 48: 166-168.

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REFERENCES

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in text; full references at the end of the article, arranged alphabetically by names, chronologi-

cally within each name, with suffixes a, b, etc. to the year for more than one paper by the same

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Examples (note capitalization and punctuation)

BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FIscHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris

88: 100-140.

FIscHER, P.-H., DUvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des

littorines. Archs Zool. exp. gén. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee

region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320.

Konan, A. J. 1960b. Spawning behaviour, egg masses and larval development in Conus from the

Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn: SCHULTZE, L.

Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und

zentralen Siid-Afrika. 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16:

269-270.

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Bryan R. Burrage

POPULATION STRUCTURE IN

AGAMA ATRA AND CORDYLUS CORDYLUS CORDYLUS

IN THE VICINITY OF DE KELDERR

aoe 7 2 OF

_ VOLUME 66 PART 2 OCTOBER 1974

ANNALS

“OF THE SOUTH AFRICAN

CAPE TOWN

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 66 °#£4Band

October 1974 Oktober

Part Z Deel

A NEW SOUTH AFRICAN REPRESENTATIVE

OF THE SOUTH WEST AFRICAN GENUS NAMIBIM YDAS

HESSE (DIPTERA: MYDAIDAB),

WITH SOME ECOLOGICAL NOTES ON THE HABITS

OF THE SPECIES

Au J. HESSE

Cape Town Kaapstad

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

A NEW SOUTH AFRICAN REPRESENTATIVE OF THE SOUTH WEST

AFRICAN GENUS NAMIBIM YDAS HESSE (DIPTERA: MYDAIDAB),

WITH SOME ECOLOGICAL NOTES ON THE HABITS OF THE SPECIES

A. J. HESSE

South African Museum, Cape Town

(With 1 figure)

[MS. accepted 8 October 1973]

CONTENTS

PAGE

Introduction é : : 4 : : : : : 25

Description : : : : ; : : , ‘ 26

Ecological notes : ; A ; ‘ : : : : 31

Supplementary notes to original generic description . 3 32

Distribution of the genus : A : : : : : 33

Summary 2 : : : 5 : : : : , 33

Acknowledgements : : : ‘ : : : 34

References . : : : ; : : : ; 34

INTRODUCTION

Since the publication of my paper (Hesse 1972) on new Mydaidae from

the Namib Desert and south-western Africa a new representative of the genus

Namibimydas, described therein, was obtained from the west coastal region of

South Africa.

As the genus Namibimydas is remarkable in many respects and as another

representative of it has now been discovered far south of the Namib Desert,

but also along the sandy coastal belt, a description of this new species is

imperative.

This discovery of another species, represented by several specimens of both

sexes, also enables me to supplement the original description of the genus which ~

was based on only two old, somewhat damaged, specimens in the Stuttgart

Museum.

Moreover the discoverer of the interesting new species also made some field

observations on the habits of this mydaid which are as remarkable as the insects

themselves and which, in view of our extreme ignorance of the ecology of

Mydaidae, deserve to be recorded.

It all began early in January 1973 when Mr A. J. Prins, our Assistant

Entomologist, was on vacation at Paternoster on the west coast between Sal-

danha Bay and Stompneus Bay and, when unprepared for the occasion, he first

noticed a 9 of these mydaids ovipositing in the sand.

To obtain specimens for the Museum Mr Prins, accompanied by Mr V.

Ann, S. Afr. Mus. 66 (2), 1974: 25-34, 1 fig.

26 ANNALS OF THE SOUTH AFRICAN MUSEUM

Branco, our artist, visited the same locality again early in February 1973. As in

the case of most South African species and the Mydaidae in general, they also

found that the desired insects were by no means plentiful and they were able to

obtain only 8 specimens during the two days they spent there.

DESCRIPTION

Subfamily Syllegomydainae

Tribe Halterorchini

Genus Namibimydas Hesse

Namibimydas prinsi sp. nov.

This new species which I have great pleasure in dedicating to Mr A. J.

Prins, who first discovered it, is characterized as follows:

Body-colour with the head and body mainly dark or blackish, appearing

darker or more black in $34, sometimes more greyish black in 99; pleural and

sternal parts and venter in some 99 (tenerals) appearing pallid; sides of mesono-

tum, postalar calli, scutellum and metanotum in some 99 (probably teneral)

also sometimes paler, more obscurely pallid to dirty yellowish; lower part of

buccal cavity (more extensive in 99), anterior spiracles and notopleural suture,

sclerites below wing-bases, posterior thoracic spiracles or part below halteres,

hind margins of tergites 3-7 in gg and 2-7 in 99, especially on sides, the sides

of tergites obscurely in teneral 99, lateral and posterior margins of hypopygial

tergite 9 of 33, processes and broad sides of navicular sternite 9 in gg, and the

acanthophorites and spines in 99 yellow or yellowish; bullae, even in 39, rather

small, dull blackish red to black, elongate-oval, far apart along hind margins

of tergite 2, very much smaller in 99; antennae mainly dark blackish brown to

black, only the articulations between the joints and the terminal tubercles usually

paler, more pallid or yellowish (the entire antennae in teneral 99 appearing

subpallid or dirty yellowish); proboscis blackish brown to black, paler in

tenerals; legs in 3 with the coxae blackish brown to black, the middle and hind

ones, especially latter, more yellowish brown towards apex below, the front

femora mainly blackish brown, only the lower apical half yellowish, the middle

and hind femora blackish brown above, yellowish below, the bases of hind

ones also tending to be more yellow, with all the tibiae narrowly darkened above

to a variable extent, the hind ones more evidently so apically, the tarsi mainly

yellowish, but also slightly darkened above on apical parts of the joints, the

last 2 or 3 joints sometimes more extensively darkened, the claws black in

apical halves; legs in 99 more extensively pallid or yellowish, more indistinctly

or scarcely darkened on femora and tibiae above, the legs entirely pallid or

yellowish in teneral forms.

Integument of head and body mainly dull, covered with greyish white

tomentum to a variable extent; part of median ocellar ridge, antennal joint 3,

head below on each side of buccal cavity, and the proboscis more or less shining;

A NEW SOUTH AFRICAN REPRESENTATIVE OF THE GENUS NAMIBIMYDAS HESSE 27

thorax above mainly dull, leathery, with setiferous puncturation; pleurae also

mainly dull, the middle part of sternopleuron, not covered with tomentum,

shining to a variable extent; metanotum dull, leathery; abdomen dul! above,

setiferously punctured, the minute punctures lodging the hairs, the extreme sides

of tergites tending to be more shiny, especially in 99; the sides of tergites 2—7

in 99, especially intramarginally, rugose, transversely striate in 34, tergite 7 in

0° transversely coarsely striated or grooved discally, and to a certain extent also

on tergite 6 laterally and posteriorly; hood-like tergite 8 in 99 punctured, less

coarsely transversely striate, also more shining; sternum shining; venter in both

sexes shining or subshining, transversely striated in 99, less distinctly so in ¢3;

hypopygium in §¢ mainly dull, but basal junction between tergite and sternite 9,

to a certain extent sides of 9, and keel of 9 shining; acanthophorites and

their spines in 9° also shining; legs, including coxae, more or less shining,

the femora finely transversely striate, and the coxa, also more or less striate

laterally.

Vestiture on head, body and legs mainly snow white in both sexes, longer

and denser in §¢ than in 99; that on head long, dense and conspicuous in 34,

slightly shorter in 99; postvertical bristles white, slender, hair-like, difficult to

see among the dense occipital hairs, more discernible in 99; hairs on occipital

part laterally behind eyes shorter than rest of cephalic hairs, shorter in 99;

palps with a tuft of long hairs and base of proboscis below also with some long

hairs; head in front and sides of occiput dull, covered with fine greyish tomen-

tum, denser and more evident along eye-margins; hairs on thorax above in

front longer and denser than on rest of thorax, shorter in 99; those on disc of

mesonotum shorter and sparser, shorter in 99, leaving a bare streak on each

side in posterior half in both sexes, with a tuft of longish hairs medially in

front of scutellum, and also with longish ones along notopleural part, above

wing-bases, and on postalar calli; metanotum mainly bare on greater middle

part, but with long hairs on sides; pleurae thinly covered with greyish tomen-

tum, dull, except for the more shiny sutural and sternopleural parts, with an

upper posterior tuft of long hairs on mesopleuron, long hairs along its posterior

margin, long ones on pteropleuron and dense ones on metapleuron anterior

to halteres and posterior thoracic spiracles, the hypopleuron mostly bare except .

for some long hairs along its posterior coxal border; abdomen in 34 with fairly

dense and long snow white hairs, those on tergites 1 and 2, especially on sides,

slightly longer and denser, but those on sides of 3—5 in tufts, scarcely shorter,

and those on sides of rest of tergites also long, but less in tufts, those on hypopy-

gium, where present, also long and conspicuous, the hairs on venter in dd

slightly sparser than on abdomen above, but also long, with the hairs on abdo-

men above in g4, when viewed from above, more or less concentrated medially

on tergites 3-7 in form of rosette-like tufts; hairs on abdomen above in 9°

long and dense only on tergite 1, especially on sides, with only a small tuft of

shortish ones on sides of tergites 2 and 3, the rest of surface above on tergites 2-6

with minute, decumbent hairs in fine setiferous punctures and which in certain

28 ANNALS OF THE SOUTH AFRICAN MUSEUM

lights have a sericeous yellowish tint, with the hairs on tergite 7 and the posterior

genital tergite 8 longer than the minute ones on preceding tergites, more seri-

ceous white and directed forwards, with the abdomen above in 99 also densely

covered with greyish white or slaty grey tomentum on tergites 3—7 (this is some-

times denuded to a variable extent), the hairs on acanthophorites and anal

lobes yellowish to golden yellowish, with the hairs on venter in 99 minute,

much sparser than on tergites, tinted slightly sericeous yellowish in certain

lights, those on sternite 7 also sparser, but longer than on preceding sternites

and directed forwards as on tergite 7; hairs on legs in 3, especially on femora,

denser, longer and more bushy than in 99 in which sex the longer ones are

distinctly less dense, especially on hind legs.

Head much broader than across broadest part of thorax, quite 3,83-4 mm

in $d and 3,68-4,04 mm in 99, though on the whole the head of 99 appears

relatively narrower; interocular space on vertex markedly broad as originally

described for the genus, on the average slightly narrower in §¢ (2,32-2,68 mm)

than in 99 (2,4—-2,72 mm), vertex not much sunk in, the central ocellar ridge also

ends on vertex in a fairly deep sulcus as described for the type-species of the

genus, the boss-like part on each side however less conspicuously prominent;

antennae (cf. Fig. 1A) distinctly much shorter than interocular space, joint 1

thickened, about 14 to nearly 3 times length of 2, joint 3 stout, columnar, about,

to even slightly more than, twice length of 1 and 2 combined, much shorter

than club, its apical part broadened, the club itself nearly or about 13 times

as long as joint 3, subspindle-shaped, especially in side view, thickest just before

to about the middle, covered with a faint, but distinct, brassy pruinescence or

tomentum, except on sensory area, with a belt of minute, spine-like hairs around

the thickest middle part and across this part also a few longer, gleaming hairs

mainly across the inner, upper and outer sides, especially in 92, with the dis-

tinctly-demarcated sensory area, in more or less apical half below, large, elongate-

ovate and bordered, ending apically in the crater-like terminal prominence

lodging a minute stylet; proboscis (Fig. 1B) relatively long, slender, somewhat

variable in length, about 2—2,6 mm (the latter when fully extended), on the

whole slightly shorter than antennae, with a few, scarcely discernible, hairs

along lower part on each side, at least in 9, its labella short, oval, the apex

bluntly rounded, sometimes appearing obliquely subtruncated; palps small,

subglobular.

Wings comparatively short, not reaching tip of abdomen, entirely and

markedly transparent, with a faint milky white tint in certain lights; veins white

and transparent, only the extreme base of wings, costal vein and to a certain

extent second vein slightly yellowish; venation like that of Afroleptomydas;

first posterior cell obtusangular apically, either angular and shortly stalked

apically or narrowly opening apically on second vein; second submarginal cell

with an appendix basally; discoidal cell acute and shortly stalked apically;

axillary lobe and alula moderately developed; halteres whitish.

Legs rather short, slender, the femora in 34 slightly thicker than in 99,

A NEW SOUTH AFRICAN REPRESENTATIVE OF THE GENUS NAMIBIMYDAS HESSE 29

the hind ones in 99 longer, but in g¢ very slightly more thickened, the legs in

3d distinctly more densely hairy; femora in 33 with only dense long and short

hairs below, no distinct spines being discernible, even on hind ones below;

femora in 99 also with long and short hairs, more above than below, but much

fewer and less dense than in 33, the front and middle ones without any detectable

spines below, the hind ones less hairy, mainly short-haired above, with com-

paratively only a few long ones basally above and, in addition to a row of long

bristly hairs along inner side below, with more or less two irregular rows of

comparatively short, slender, white, bristle-like spines, as well as irregularly

disposed, shorter, spine-like setae or seta-like spinelets below; tibiae curved in

both sexes, the hind ones so near apex, hairy in both sexes, but more densely

and to a certain extent longer so in 3d, especially the hind ones in latter sex

with dense long and shorter hairs, with the front and middle tibiae in $3 armed

below with fine hair-like or bristle-like spicules, these in 2° very slightly thicker,

and hind tibiae in 29 with stouter and more distinct spicules; apical spicules or

spurs of tibiae below well developed, fairly long, golden yellow, equally deve-

loped in both sexes; tarsi with the spicules below, especially on hind ones, also

well developed and golden yellow; claws in 39 longer, more developed than in

29; pulvilli in 99 shorter and less developed than in 33.

Fig. 1. Namibimydas prinsi sp. nov.

A. Side view of left antenna of 2 allotype. B. Side view of proboscis of 2 allotype. C. Posterior

view of hypopygium of ¢ (vestiture partially omitted to show some structures more clearly).

D. Right side view of the partially extruded hypopygium of a d. (AE = the characteristically

long, curved aedeagus lodged in the keel part of sternite 9; AL = anal lobes; Ap = appendage

of concealed or semi-concealed segment 8; Ep = epimere; IX.S. = large boat- or shell-shaped

stermite 9; IX.T. = lid-like tergite 9, immovably attached to sternite 9; Pr = processes of

sternite 9; VII.T. = tergite 7 partially or entirely covering tergite 8.)

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

Hypopygial complex of 3 (Fig. 1C—D) with tergite 8 sometimes exposed as a

short half-moon-shaped posterior part, depending on the upward or downward

movements of the hypopygium, its projecting lateral lobe on each side of base

of sternite 9 lobe-like; sternite 7 with only its hindermost part exposed, being

almost entirely pushed forwards by base of sternite 9, its exposure or conceal-

ment being due to the upward, downward and forward movements of the

hypopygium; hypopygium itself large and conspicuous as described for the

genus (Hesse 1972; 160), in form of an oval box-like structure in side view,

tergite 9 and anal lobes constituting a sort of fixed lid, the tergite itself about

1,6-1,8 mm long, convex, deeply and angularly incised posteriorly in which

the anal lobes are situated, basally slightly transversely depressed above and

on sides basally firmly and immovably united by a shining, weal-like, strongly-

chitinized connection on each side dorsally to sternite 9, and which also

constitutes the base of the prong on each side of the latter sternite; sternite 9

large, navicular, about 2,72—2,8 mm long, compressed from side to side, the

more strongly-chitinized keel of the ‘boat’ being narrower than the sides above,

posteriorly the sternite is deeply and angularly incised V-like; upper chitinized

and ridge-like side or ‘gunwale’ of this boat-like sternite is continued pos-

teriorly as a lateral flange to form the posteriorly projecting, flattened, down-

wardly-directed, tongue-shaped process or prong on each side, the upper

posterior part of the ‘gunwale’, which also constitutes the upper part or corner

of the V-shaped incision on each side, is prominent, rounded, strongly chitinized,

shiny, and connected with the lateral prong, the less chitinized, paler sides of

the sternite with long white hairs which are directed downwards posteriorly on

each side of incised part; keel of sternite 9 is flattened, strongly chitinized, brown,

and lodges the long, double-tubular aedeagus; aedeagus itself remarkably long,

in form of a long, curved, double-tubular organ, composed of two parallel

adherent tubes, thicker basally than posteriorly and lodged in the keel-part

of sternite 9, bending forwards posteriorly just inside the posterior V-shaped

incised part of the sternite and there becoming separated as two separate,

thinner, aedeagal branches, each ending in a gonopore; epimere, as far as this

can be seen, in form of a flattened bluntly-forked or bifid structure anterior to

the aedeagal branches of aedeagus and pointing slightly backwards.

The process of copulation, though not observed by the collectors, probably

takes place end to end as in Asilidae. In this case both the aedeagal tubes and the

forked epimere are probably pushed backwards or posteriorly through the

posterior V-shaped incision of sternite 9 to engage the genital organs of the 9

in which they are also aided by the posterior lateral processes of the sternite

which probably act as hooking structures.

Genital segments of 2 structurally more or less as described and figured for

the genus (Hesse 1972: 161, 164); hood-like tergite 8 with its hind margin

slightly produced medially, there rounded and carinate, the disc transversely

coarsely striate and punctured, more coarsely punctured intramarginally

posteriorly; acanthophorites each with about 7 flattened, apically slightly

A NEW SOUTH AFRICAN REPRESENTATIVE OF THE GENUS NAMIBIMYDAS HESSE 3]

broadened, yellowish spines, the first one at the base being more slender and

rod-like; anal lobes with dense, short, stiff hairs or setae; sternite 8 half hidden,

its hind margin emarginate or indented and its sides slightly rugose.

From 1 3g holotype, 1 2 allotype, 3 3 paratypes and 3 9 paratypes in the

South African Museum.

Distribution

South-western Cape: Paternost2r (A. J. Prins, 13 February 1973) (3 holo-

type, allotype, 2 paratype, and 3 3 paratypes); Paternoster (V. Branco,

13 February 1973) (2 2 paratypes).

Compared with the type-species gaerdesi Hesse (Hesse 1972: 161) from

the Namib Desert, this new species may at once be distinguished by its much

smaller size, relatively shorter, more slender legs, comparatively denser and

entirely white vestiture in 93, distinctly very much shorter or minute hairs on

abdomen above in 99, hairs on metanotum confined to sides, joint 3 of antennae

proportionally longer relative to the other joints, much shorter and relatively

less slender proboscis which is slightly shorter, not longer, than antennae and

with a more bluntly-rounded apex of labella, the finer and more numerous spines

and spinelets on hind femora below, especially in 99, comparatively shorter

apical spurs on tibiae, the more boat-shaped, not Argonauta-shell-shaped,

sternite 9 of the ¢ hypopygium, the broader, more lobe-shaped lateral appen-

dages of the half-concealed tergite 8, the more rounded upper corners of pos-

terior V-shaped incision of sternite 9 which is not fused with lower part of

posterior lateral lobes of tergite 9, the fewer (only about 6 or 7, not 8) spines on

acanthophorites of 99, and the less sharply or deeply indented hind margin of

the half-concealed sternite 8 in 99.

ECOLOGICAL NOTES

Habitat

According to the observations of Mr Prins this mydaid is by no means

plentiful and occurs only along the littoral sandy zone on the coast near the sea,

and is practically confined to the superlittoral part of it not reached by high tide.

The region frequented is between the limit of the vegetation and the high tide

mark, a white sandy environment strewn with broken shells, fragments of sea-

weed, especially those of Ecklonia, and also other debris washed up by the high

tide or blown there by wind.

To this type of environment the mydaid is eminently adapted procrypti-

cally. Its dense snow white vestiture, transparent wings, greyish tomentum, and

bare blackish patches on the abdomen of the 99, render the species almost

invisible against the background, and more so if individuals rest on the sand.

Movement and flight

According to Mr Prins this insect appears to be remarkably agile in moving

to and fro and sideways on the sand, being able to crawl backwards as easily

32 ANNALS OF THE SOUTH AFRICAN MUSEUM

as forwards. In flight it appears to be as agile and even faster, darting back-

wards, forwards and sideways, within limits, with equal ease. These movements

render it a difficult insect to catch. Moreover it appears to be very sensitive

to vibrations in the sand caused by approaching footsteps or the slightest thump

on the sand.

Oviposition

By sheer coincidence Mr Prins observed the procedure adopted by

the 9 in the deposition of her eggs. During his first visit he was resting on the

sand when, a little distance away from him, a 9 alighted and began to insert the

tip of her abdomen in the sand. This she wriggled in until the entire abdomen

disappeared in the loose sand. Not content with the depth reached she wriggled

still further in until the entire thorax, wings and legs disappeared. Eventually

the only parts of her anatomy still exposed were the pair of protruding antennae

and front half of the eyes, frons, and clypeal part.

Equally rapidly she wriggled out of the sand again. Mr Prins then very

carefully scooped out the patch of sand, containing some whitish eggs, and

placed the sample in a glass tube with the intention of experimenting with it

later on. Unfortunately some unauthorized visitor at the hotel, who wanted a

glass tube for some other purpose, threw the sand away. Since then and during

subsequent visits to the same area all efforts on the part of Mr Prins to observe

another ovipositing female and to collect her eggs have been in vain.

As far as I am aware this is the first record of a dipterous insect ovipositing

in this remarkable way of penetrating and almost disappearing backwards into

a sandy medium. This method of oviposition appears to be still another adaptive

response of this mydaid to a loose sandy environment.

Enemies

It was observed that a great enemy of this mydaid is a sand-loving and

dune-frequenting species of lizard which, according to the two collectors,

snapped up several specimens which they themselves tried to obtain.

SUPPLEMENTARY NOTES TO ORIGINAL GENERIC DESCRIPTION

As the original generic description of Namibimydas (Hesse 1972: 158-161)

was based on only two old specimens (¢ and 9) of the type-species gaerdesi, the

addition of 8 specimens of the new representative of the same genus enables me

to supplement, redefine or correct the original description of some of its struc-

tures as follows:

Head with the broad interocular space on vertex proportionally either

more or less equally broad in both sexes or slightly broader in 99 than in 3@;

club of antennae longer than joint 3, elongate pyriform to spindle-shaped; the

labella of the elongate and slender proboscis either spear-blade- or lance-blade-

Shaped and sharply pointed or subtruncately rounded apically.

A NEW SOUTH AFRICAN REPRESENTATIVE OF THE GENUS NAMIBIMYDAS HESSE 33

Abdomen with tergite 8 in g¢ either entirely concealed or only partly

exposed above base of tergite 9, saddle-shaped; sternite 7 in Sg either displaced

forwards and hidden under 6 or its posterior part exposed as a short cover over

extreme base of sternite 9.

Hypopygium of 35 remarkably and conspicuously large, in form of an oval,

box-like structure, with tergite 9 and anal lobes constituting the lid, but the

tergite itself immovably united on each side basally to sternite 9; latter sternite

enormously developed and also as described for this species, boat-like, shell-like,

keeled below, with a projecting process or prong on each side postero-dorsally,

posteriorly vertically deeply incised (V-shaped), the upper angles or corners of

the incision (or posterior angles of sides of sternite 9) prominent, rounded, or

broadly projecting upwards under the apical part of posterior lateral lobes of

lid-like tergite 9 (this latter condition described by me for the type-species

gaerdesi (Hesse 1972: 162, fig. 6 right) as an obvolvent structure connected or

fused to tergite 9 may probably not be united to tergite 9 at all, for it could not

be seen as an unconnected structure in the old dried ¢ holotype); aedeagus

(which also could not be seen properly in the unique ¢ of the type-species) is, as

described for this new species, remarkably long, in form of a long, curved,

strongly-chitinized, double-tubular organ, composed of two parallel, adherent

tubes, lodged in the keel part of sternite 9 and which posteriorly become sepa-

rated into two separate aedeagal branches which point forwards; epimere

flattened and forked.

Legs with either a double row of distinct spines on slight tubercles, or with

more numerous seta-like spinelets, on hind femora below.

DISTRIBUTION OF THE GENUS

In view of the fact that most of the known genera of Mydaidae in South

Africa occur as separate, more or less geographically-restricted species and not

as very widely distributed forms, the genus Namibimydas, which now occurs as

two separate species, at Walvis Bay and Paternoster respectively, may also be

represented by still other unrecorded species along the coastal, littoral, sandy

belt, stretching from the south-western Cape to Angola. Judging from the |

habitat of the new species and that of the type-species it appears that this genus

will be found to be restricted to the comparatively narrow, sandy or dune-sand

environment along the west coast between the high tide mark and the landward

limit of the non-marine vegetation. ;

SUMMARY

A new species prinsi of the genus Namibimydas Hesse, originally described

from the Namib Desert, is described from Paternoster on the west coast between

Saldanha Bay and Stompneus. Some ecological notes on the habits of the new

species are added, and supplementary notes to the original generic description

34 ANNALS OF THE SOUTH AFRICAN MUSEUM

are appended. A text-figure is given to illustrate the more important generic

and specific characters of the new mydaid.

ACKNOWLEDGEMENTS

My thanks are due to the Assistant Entomologist Mr A. J. Prins and one

of the Museum’s artists, Mr V. Branco, who collected the specimens of the new

species at Paternoster. To Mr Prins I am indebted for the interesting observations

he made on the habits of this species.

REFERENCES

Bezzi, M. 1924. The South African Mydaidae (Diptera) as represented in the South African

Museum. Ann. S. Afr. Mus. 19: 191-232.

Hesse, A. J. 1969. The Mydaidae (Diptera) of southern Africa. Ann. S. Afr. Mus. 54: 1-388.

Hesse, A. J. 1972. New Mydaidae (Diptera) from the Namib Desert and south-western Africa.

Ann. S. Afr. Mus. 60: 109-171.

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BuLLouGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris

88: 100-140.

FiscHER, P.-H., DuvAL, M. & RarFFy, A. 1933. Etudes sur les échanges respiratoires des

littorines. Archs Zool. exp. gén. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee

region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320.

Konan, A. J. 196056. Spawning behaviour, egg masses and larval development in Conus from the

Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51. :

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L.

Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und

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A. J. Hesse

A NEW SOUTH AFRICAN REPRESENTATIVE

OF THE SOUTH WEST AFRICAN GENUS NAMIBIMYDAS

HESSE (DIPTERA: MYDAIDAB),

WITH SOME ECOLOGICAL NOTES ON THE

HABITS OF THE SPECIES

| VOLUME 66 PART 3 OCTOBER 1974

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 66 Band

October 1974 Oktober

Part 3 Deel

hE CRANIAL MORPHOLOGY

OF THE LOWER TRIASSIC DICYNODONT

MYOSAURUS GRACILIS

MICHAEL A. CEUVER

Cape Town Kaapstad

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

THE CRANIAL MORPHOLOGY OF THE LOWER

TRIASSIC DICYNODONT MYOSAURUS GRACILIS

MICHAEL A. CLUVER

South African Museum, Cape Town

(With 6 figures)

[MS. accepted 15 October 1973}

CONTENTS

PAGE

Introduction . ; ; ‘ é : res)

Material and techniques . : ‘ : a 36

Classification . ; : : ; ; ne EST

Cranial morphology of Myosaurus gracilis . ae 7

Jaw function in Myosaurus : : : 5 et

Comparison with other dicynodonts . : . 49

Summary : : : ’ : : Li Oe

Acknowledgements . : : : ‘ Sy

References : : : ; Y ; DS)

Abbreviations ; Q ; : Shea

INTRODUCTION

The dicynodont Myosaurus gracilis Haughton is known from several small

skulls, some with associated postcranial remains, all from a single locality on

the commonage of Harrismith, O.F.S., South Africa, where the remains of a

Lystrosaurus zone (Basal Triassic) fauna occur in abundance. The genus is

remarkable in that, until recently (Hotton 1974), it represented the only small-

sized dicynodont form known to have survived the Permian-Triassic transition

as recorded in the South African Beaufort succession. With an average skull

length of 40 mm, Myosaurus is dwarfed by other South African Triassic

dicynodont genera, such as its contemporary Lystrosaurus (typical skull length

100 mm) and the Cynognathus zone Kannemeyeria (typical skull length 400 mm).

In view of its almost isolated stratigraphic position, the cranial morphology

and possible relationships of Myosaurus would appear to be matters of some

interest, but after Haughton’s (1917) description, no further preparation of the

South African Museum type material was undertaken, with the result that, with

the exception of the skull roof and occiput, the skull structure remains largely

unknown. With the emphasis in dicynodont classification now increasingly

involving areas other than the skull roof (Toerien 1953; Cluver 1970; Hotton

& Cluver in press) it seemed that a detailed description of the material, including

the palate, basicranium, braincase and lower jaw, would be necessary before

the relationships of the genus could be adequately considered.

Ann. S. Afr. Mus. 66 (4), 1974: 35-54, 6 figs.

36 ANNALS OF THE SOUTH AFRICAN MUSEUM

In his original description, Haughton (1917) considered that Myosaurus

was allied to Emydops, but differed in being free of molar or canine teeth;

Van Hoepen (1934) came to a similar conclusion. Broom (1932) mentioned

resemblances to Cistecephalus, from which, however, the genus differed in its

small size and retention of the preparietal bone. The only other reference to the

relationships of Myosaurus was made by Broom (1941) when describing

Myosauroides minnaari, which he thought differed from Myosaurus mainly

through its lack of a preparietal. Myosauroides, from the Cistecephalus zone,

could, according to Broom, be placed ‘. . . somewhere between Myosaurus and

Cistecephalus’ (p. 210).

MATERIAL AND TECHNIQUES

In addition to the two co-type skulls (S.A.M. Nos 3526 and 3526a) in the

South African Museum, five specimens were obtained on loan from the Bernard

Price Institute for Palaeontological Research, University of the Witwatersrand,

Johannesburg; two of these latter specimens (B.P.I. Nos 2701b and 4269) have

associated lower jaws. Disarticulated postcranial material is associated with the

two type skulls, while B.P.I. No. 4269 has the articulated anterior third of the

postcranial skeleton attached to the skull.

The bulk of the preparation was confined to the two type skulls, which

are well preserved and virtually undistorted and which together complement

each other to provide a complete picture of the skull morphology. These two

specimens were completely cleared of matrix by means of automatic mallet and

needle, and all areas, including the interior of the braincase, were made available

for examination. Specimen 3526a had suffered a fracture across the back of the

skull base during the original preparation, and it was thus also possible to clean

out the inside of the left otic capsule. Preparation of the B.P.I. specimens was

not carried to the same extent, and was mainly limited to exposure of the palate

and, in two cases, the lower jaw.

Illustrations of the skull are orthoprojections; in the case of dorsal and

ventral views the projections are on to a plane including the base of the foramen

magnum and the ventral, inner surface of the nostril. The illustrations are

mainly composite reconstructions of both co-type skulls, but, thanks to the

almost total lack of distortion in the material, the reconstructions accurately

represent the actual condition of the skulls.

In addition to these specimens of Myosaurus, the type (R.C. 54) of

M yosauroides minnaari Broom was obtained on loan from the Rubidge Collec-

tion, and the palate and mandible prepared. A second, better preserved specimen

of this species (A.M.N.H. 8209), loaned from the American Museum of Natural

History, New York, yielded additional information after preparation. Several

specimens of Cistecephalus in the collection of the South African Museum as

well as the type and paratype (BMNH R4956, R4957) of Emydops longus in the

British Museum (Natural History) London, were also used for comparison

with both Myosaurus and Myosauroides.

CRANIAL MORPHOLOGY OF TRIASSIC DICYNODONT M YOSAURUS GRACILIS SH

CLASSIFICATION

For present purposes, the dicynodont classification of Haughton & Brink

(1954), modified after Romer (1966) (see also Cluver 1974) can be most con-

veniently used to state the present systematic position of Myosaurus gracilis:

Class: Reptilia

Order: Therapsida

Suborder: Anomodontia (sensu Romer 1966)

Infraorder: Dicynodontia (sensu Romer 1966)

Family: Endothiodontidae (sensu Haughton & Brink 1954)

Subfamily: Pristerodontinae

Genus: Myosaurus Haughton

Diagnosis: Skull small, with relatively large orbits and weak snout. Intertem-

poral portion of skull roof wide. Postfrontals absent, pineal foramen relatively

far back in skull roof. Squamosal flared sharply laterally above quadrate.

Frontals and prefrontals make near-contact in midline ventrally to olfactory

passage. Secondary palate long, no anterior palatal ridges. Palatine small,

meeting premaxilla. Long vomerine septum dividing internal nares.

Myosaurus gracilis Haughton (1917)

Diagnosis: as for genus.

Co-types: Two skulls in nodule (S.A.M. Nos 3526 and 3526a) from Harrismith,

O.F.S., South Africa.

Horizon/age: Lystrosaurus zone, Lower Triassic.

CRANIAL MORPHOLOGY OF MYOSAURUS GRACILIS

Skull roof

The nostrils are nearly terminal, and the anterior portion of the premaxilla,

dividing the nostrils, is accordingly reduced to a delicate splint of bone, missing

in most specimens. A short internasal process of the premaxilla extends back

between the nasals, which may form slight bosses above and behind the nostrils.

Behind the nostrils, and in front of the anterior border of the orbits, there is a

slight narrowing or waisting of the snout, adding to the generally weak appear-

ance of this anterior part of the skull. The septomaxilla is small, and confined

to the interior of the nostril, where it forms part of the posterior wall. The

maxilla rises fairly steeply behind the nostril, but is separated from the pre-

frontal by a narrow but persistent process of the lacrimal (Fig. 1B). The pre-

frontal is not strongly developed on the outer surface of the skull, and no boss

is formed; the large orbit is thus almost circular in antero-lateral view.

The frontals (Fig. 1A) are important elements of the skull roof, meeting

the nasals far forwards in an almost transverse suture, and extending back to a

point level with the middle of the temporal fenestra. The pineal foramen lies

38 ANNALS OF THE SOUTH AFRICAN MUSEUM

e€O0c

den

ang

Fig. 1. Myosaurus gracilis x 4. A-B. Skull in dorsal and lateral view

respectively. C. Mandible in lateral view.

CRANIAL MORPHOLOGY OF TRIASSIC DICYNODONT MYOSAURUS GRACILIS 39

relatively far back in the skull, and there is a prominent, diamond-shaped

preparietal.

Apart from the posterior portions of the frontals, the broad intertemporal

region is composed of the parietals and postorbitals, a postfrontal being alto-

gether absent (Fig. 1A). Each parietal is wide posteriorly, and bifurcates ante-

can.proc

Pc.cr.

pal

lat. pal. for.

=4 LCA

bas

boc

Fig. 2. Myosaurus gracilis x =. A. Skull in ventral view. B. Mandible in dorsal

view.

riorly to surround the rear of the frontal of its side. Laterally it meets the post-

orbital, which lies as a thin horizontal sheet over the upper part of the temporal

opening. This part of the postorbital is essentially a lateral continuation of the

parietal sheet, and therefore a bone of the dorsal skull roof. The lateral edge

of the post-orbital is thickened and forms a stout dorsal border for the temporal

fenestra.

40 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 3. Myosaurus gracilis. A. Palate in ventral view, x 2. B. Snout in posterior

view, X =. C. Occiput, x %. Left stapes removed.

Occiput

The interparietal (Fig. 3C) lies wholly in the occiput, and does not overlap

on to the skull roof, from which the occiput is clearly demarcated (Fig. 1A).

A median crest divides the interparietal into two halves. Although not clear in

all specimens, tabulars are present and lie as squarish plates on each side of

the interparietal. The supraoccipital is clearly distinguishable from the sur-

rounding dermal bones, and is on each side bounded by the crescent-shaped

opening of a nutrient channel (Fig. 3C), similar to that found in Lystrosaurus

(Cluver 1971). Each opening lies between the lateral edge of the supraoccipital

and the medial edge of the squamosal and is closed off ventrally by a short

inner lappet of the squamosal which makes a brief contact with the ventro-

lateral corner of the supraoccipital. This process of the squamosal is pierced

by a small foramen, which opens into the crescent-shaped space; directly below

CRANIAL MORPHOLOGY OF TRIASSIC DICYNODONT MYOSAURUS GRACILIS 4]

lies the large posttemporal fenestra. It seems (Cox 1959; Cluver 1971) that all

of these channels served for the forward drainage of blood from the neck

musculature into the cranial cavity, prior to its discharge into the v. capitis

lateralis and v. jugularis (see below).

The sutures between supraoccipital, exoccipital and opisthotic are clearly

visible in occipital view. The exoccipital, carrying a process for articulation with

the proatlas, is seen to have a limited lateral extent, so that the opisthotic meets

the supraoccipital in a suture leading towards the posttemporal fenestra. The

opisthotic is a stout bone, abutting against the squamosal laterally and notched

for the passage of blood-vessels on the other three sides. Dorsally it closes off

the posttemporal fenestra, through which passed the v. capitis dorsalis (Cox

1959); medially it forms the lateral wall of the jugular foramen and ventrally it

is cut away for the course of the v. capitis lateralis.

The area of the braincase between the posttemporal fenestra and the jugular

foramen is swollen, and stands out as a smooth bulge on the occiput. The

exoccipitals meet in the midline ventrally, and form the posterior floor of the

foramen magnum; the two exoccipital condyles overhang the basioccipital

condyle to a quite considerable extent (Fig. 2A).

The squamosal forms the lateral border of the occipital plate in the usual

way, but ventrolaterally it is drawn out as a prominent flange (Fig. 3C, lat. fl.)

which extends out from directly above the quadrate. This is an unusual feature,

not seen in other dicynodonts, and is most likely related to development of the

lateral external jaw adductor muscle mass, for which the anterior face of this

portion of the squamosal serves as an area of origin.

The quadrate (Figs 1B, 2A, 3C) is basically a vertical disc with a sharply

offset lateral articular shelf, and the articular surfaces are at sharp angles to

each other. The ventral, highly convex edge of the disc articulates with the

medial articular process of the lower jaw, while the lateral, more gently convex

lateral condyle meets the main articular surface of the articular bone. The two

articular surfaces of the quadrate are separated by a sharply defined vertical

step.

The quadratojugal is fused with the dorsal surface of the lateral quadrate

condyle, and meets the squamosal dorsally in a highly interdigitating suture. A

large quadrate foramen (Fig. 3C) lies between the quadratojugal and quadrate,

and opens low down on the occiput between the rear of the lateral quadrate

condyle and the ventral edge of the descending sheet of the squamosal.

The stapes (Fig. 2A) is a simple rod, inflated both proximally, to form a

fairly distinct footplate, as well as distally, where it lies up against the inner side

of the medial quadrate condyle.

Skull base and palate

In ventral view (Figs 2A, 3A), the skull displays several features of impor-

tance. The large fenestrae ovales are rimmed by the high, thin-walled basiocci-

pital-basisphenoidal tubera. The porous nature of the bone surrounding each

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 4. Myosaurus gracilis. Stereophotographs of: A. Braincase in ventral view

(S.A.M. No. 3526). B. Interior of braincase showing left prootic and floccular

fossa (S.A.M. No. 3256a). C. Palate (S.A.M. No. 3526a).

CRANIAL MORPHOLOGY OF TRIASSIC DICYNODONT MYOSAURUS GRACILIS 43

fenestra suggests the former presence of cartilage, in which the functional

fenestra ovalis probably lay.

The pterygoids meet in the midline beneath the basisphenoid, anterior to

the foramina for the internal carotid arteries, and a fairly strong median crest

is formed. The slender quadrate ramus of each pterygoid extends back and

laterally, terminating in a recess in the medial surface of the quadrate of its

side (Fig. 3C). In front of their meeting in the midline, the pterygoids diverge

laterally and form the outer borders of the long, vaulted choana. Anteriorly

they meet the ectopterygoids and palatines of each side and terminate as fairly

prominent, vertically expanded blades. Between each anterior pterygoid blade

and the palatine is a sizeable, slit-like lateral palatal foramen; a freestanding

ventral process of the palatine forms a part of the medial border of this foramen.

The palatal portion of the palatine is limited to a small, thin sheet, per-

forated by a large foramen (Figs 2A, 3A, pal. for.). This foramen, unusual in

dicynodonts, opens into the interior of the choana above the horizontal portion

of the palatine, where a second foramen between the palatine, jugal and ecto-

pterygoid leads into the antero-ventral corner of the orbit in front of the upper

opening of the lateral palatal foramen (Fig. 3B, for.). The palatal surface of the

palatine is smooth, and its generally delicate construction, coupled with its

apparent association with a blood-vessel or nerve, makes it doubtful whether

it supported a layer of horn, as in other dicynodonts. The significance of this is

more fully discussed below.

In the roof of the choanal vault the ventral edge of the parasphenoidal

rostrum is visible in the long and narrow interpterygoidal vacuity. The vacuity

is bounded anteriorly by the vomers, in normal dicynodont fashion; in front of

this the two halves of the vomer are fused into a thin vertical septum which

extends forwards to meet the premaxilla and divide the anterior part of the

choana into two passages.

The premaxilla forms most of the secondary palate. It meets the palatine

posteriorly and the maxilla is confined to an altogether lateral position. Anterior

palatal ridges are lacking, but a median premaxillary ridge is a distinct feature.

In section the ridge is T-shaped, with an inflated palatal portion supported by a

thin vertical girder. On each side of the girder and parallel with it, the pre- -

maxilla is drawn out as a longitudinal crest, so that the median ridge is in effect

flanked by a pair of partially enclosed channels (Fig. 3A, gr.). Anteriorly, where

the median ridge merges with the general palatal surface, the grooves appear

to diverge laterally without, apparently, penetrating the premaxilla, while

posteriorly they lead towards an anterior notch in the choana, between pre-

maxilla, palatine and vomer.

Similar premaxillary grooves have been described in other dicynodonts

and it has been suggested (Cluver 1971) that they marked the passage of nutrient

blood-vessels and/or nerves. On the other hand, the anterior choanal notch of

dicynodonts, which the grooves approach in Myosaurus, has been indicated as

the most likely point of emergence of the duct of Jacobson’s organ (Cluver 1971),

44 ANNALS OF THE SOUTH AFRICAN MUSEUM

and the condition in Myosaurus could perhaps be interpreted as indication of a

doubling back, in an anterior direction, of the duct to an opening in the primitive

forward position. Equally possible, however, is an association between the pre-

maxillary grooves and the palatine foramina, and with the lack of conclusive

evidence the question must remain unresolved.

The palate is bounded laterally and anteriorly by a modest rim, which is

drawn down to a small but clear maxillary caniniform process. The medial

surface of this rim has a shallow depression in front of the caniniform process,

but this is not the deeply incised palatal notch of ‘Dicynodon’ testudirostris

(Cluver 1970; Hotton & Cluver in press), nor is any caniniform flange present.

Between the rear of the caniniform process and the expanded pterygoidal sheet

on the choanal border, the palatal rim is at first a fairly sharp postcaniniform

crest, and then becomes a low and smoothly rounded ridge, formed by the

maxilla and the small, tightly interdigitated ectopterygoid. As with the palatine,

the palatal surface and rim is nearly devoid of nutritive foramina and channels,

and the pitting and rugoseness of other dicynodont palates is absent.

Braincase, basicranial axis and orbit

The otic capsule is the most prominent part of the preserved bony braincase.

The opisthotic and exoccipital portion is inflated posteriorly above the jugular

foramen, as stated above, while in front the prootic is laterally displaced to

accommodate what appears to have been a relatively large brain. That the

structures of the brain were closely apposed to the walls of the braincase is

further suggested by the presence of a very deep floccular (sub-arcuate) fossa

in the rear of the prootic (Fig. 4B). This fossa is remarkable in that it com-

municates, by means of a fairly large opening, with the vestibule of the inner

ear (see below). On its anterior, outer face the prootic is deeply incised between

the posttemporal fenestra and the posterodorsal corner of the braincase, and

below its meeting with the parietal the prootic is cut away to form a very distinct

notch leading into the interior of the braincase. A very clearly demarcated

channel for the v. capitis dorsalis (Cox 1959; Cluver 1971) is thus formed. As

the posttemporal fenestra opens more or less directly into the ventral part of

this passage, and as the prootic shows no comparable groove below the fenestra,

it appears that most of the blood from the occiput passed into the v. cephalica

posterior within the braincase (Dendy 1909; Swain 1968); a limited portion

only would probably have flowed down through a sinus below the posttemporal

fenestra to meet the v. capitis lateralis.

Within the braincase, the pair of posterior cephalic veins would have been

joined by the small occipital veins passing forwards between the supraoccipital

and interparietal (see above) before descending to issue from the skull as the

jugular veins. There is a deep hollow in the inner surface of the parietal in the

postero-dorsal corner of the braincase above the foramen magnum, and it

seems likely that in life this was filled by a dorsal cartilaginous extension of the

supraoccipital, similar to that of Lystrosaurus (Cluver 1971); the pair of occipital

veins would probably have entered the cranial cavity at this level.

CRANIAL MORPHOLOGY OF TRIASSIC DICYNODONT MYOSAURUS GRACILIS 45

Several inner ear structures can be made out. The fenestra ovalis is rela-

tively very large (Fig. 4A), but was probably furnished with a cartilaginous rim

during life. From the fenestra ovalis a short, wide chamber leads to the vestibule.

This chamber, which may be called the perilymphatic chamber, is medially

expanded below the vestibule of the inner ear, and is unlike the simple auditory

tube of other dicynodonts, which links the fenestra ovalis with the deep struc-

tures of the inner ear. It appears that in Myosaurus a fairly large footplate, per-

haps partly cartilaginous, was associated with a considerably expanded peri-

lymphatic sac or cistern. Inside the braincase the internal auditory meatus is

confluent with the inner opening of the jugular canal, so that at this point the

otic capsule is separated from the skull base by a wide space, leading to the

jugular canal behind, the fenestra ovalis ventrally and the vestibule and succulo-

cochlear recess medially. A relatively large recess, which housed the ampullae

of the horizontal and anterior vertical semicircular canals, is visible, and it

appears that the utriculus lay in a dorsal cavity fairly clearly demarcated from

the wider saccular space below.

The floccular fossa opens into the utricular cavity, as stated above. This

appears to be a unique condition. Olson (1944) states that the anterior vertical

semicircular canal was in many cases confluent with the floccular fossa in his

sectioned dicynodonts, but in Myosaurus this is not the case, and the canal in

question arises far forwards of the inner opening of the fossa. It seems very

unlikely that a direct communication between the interior of the otic capsule

and the cranial cavity ever existed, and capsule and fossa were almost certainly

membraneously separated during life.

Close in front of the internal auditory meatus lies a foramen, in the position

of the internal opening of the VIIth (facial) nerve (Olson 1944; Cluver 1971).

A short and narrow canal leads into the foramen from the anterior, saccular

part of the vestibule, below the anterior ampullary recess, so that there appears

to be a connection between the otic capsule and the facial canal. Comparison

with a Lystrosaurus braincase (S.A.M. No. 11180; see also Cluver 1971) shows

that in this genus a deep and narrow cleft leads from below the front of the

anterior ampullary recess to the region of the facial nerve foramen, while what

appears to be a similar cleft can be seen in some of Olson’s (1944) sectioned

dicynodonts. Closure and canalization of such a cleft up to the level of the

facial foramen would lead to a condition similar to that seen in Myosaurus,

but the function of the cleft or canal remains obscure. While in a form such as

Lystrosaurus the cleft could have supported a portion of the perilymphatic

duct system, in Myosaurus the canal apparently enclosed a nerve, blood-vessel

or duct, which then, presumably, either emerged into the anterior part of the

braincase alongside the root of the facial nerve, or passed down the facial fora-

men to leave the braincase with the cranial nerve. Serial grinding of a Myosaurus

braincase would seem to offer the only means of resolving the question.

No fenestra rotunda appears to have been present, and the perilymphatic

duct probably terminated in the jugular foramen, as suggested by Olson (1944).

46 ANNALS OF THE SOUTH AFRICAN MUSEUM

The floor of the braincase, formed by the basioccipital and basisphenoid,

is smooth and flat, and a dorsum sellae is altogether absent. The paired internal

carotid arteries enter the interior of the skull through a wide common opening,

which probably housed a ventral part of the hypophysis as well. A true fossa

hypophyseos is absent. Alongside this opening the ossified bases of the pilae

antoticae of the prootics extend up as a pair of relatively long and stout pillars,

inclined slightly forwards. Immediately lateral to each pillar lies the slender

epipterygoid, rising from its footplate to meet a broad, descending flange of the

parietal. This sheet of the parietal, together with the epipterygoid and ossified

pila antotica, forms an incomplete sidewall to the brain cavity at the hypophysis-

pineal organ level.

Immediately in front of the inner opening of the internal carotid artery

lies a low bony septum, identified as a separate presphenoid element by Cluver

(1971) but regarded as an anterior basisphenoidal extension by some authors

(Broom 1926; Cox 1959). In Myosaurus the presphenoid (Figs 1B, 3B) is

clasped by the trough-like cultriform process of the parasphenoid and, as

shown in Lystrosaurus (Cluver 1971), was probably connected to a more exten-

sive cartilaginous interorbital septum at its abrupt anterior truncation. Laterally

and ventrally the parasphenoid is firmly held by the pterygoids up to the

latter’s meeting with the palatines. At this point a slit-like foramen is formed on

each side between the pterygoid and the side of the parasphenoid-presphenoid

septum (Fig. 1B, VII pal.), and presumably served for the exit of the ramus

palatinus of the VIIth (facial) nerve.

Anterodorsally the neurocranium is represented by the ossified orbito-

sphenoid-mesethmoid complex (Fig. 1B, os). This complex is best seen in S.A.M.

No. 3526a, where it has been displaced into a position suitable for detailed

preparation. The orbitosphenoid is a wide trough, with a ventral notch poste-

riorly for the exit of the pair of optic nerves, and bears a prominent ventral keel

which is deepest anteriorly. This part of the complex, lying up against the ventral

surface of the frontals, presumably housed part of the olfactory lobes. The

mesethmoid, a short but deep septum clearly distinguishable from the orbito-

sphenoid, divides the pair of olfactory channels as they pass forwards into the

snout.

Only a short space separates the mesethmoid from the anterior, inner wall

of the orbit. Myosaurus is unusual among dicynodonts in that the frontal,

prefrontal and lacrimal, which normally surround a wide, anterior orbitonasal

opening leading into the snout, have undergone an extensive medial develop-

ment, the frontals in fact almost meeting in the midline below the olfactory

opening (Fig. 3B). The interior of the snout is thus, with the exception of the

olfactory opening, almost completely closed from the orbit; a small, almost

circular space between the prefrontals probably represents an orbitonasal

opening reduced from the usual dicynodont condition (see e.g. Lystrosaurus,

Cluver 1971). Ventrally the palatines meet the prefrontals near the midline,

thereby excluding the lacrimal from the border of the orbitonasal opening.

CRANIAL MORPHOLOGY OF TRIASSIC DICYNODONT MYOSAURUS GRACILIS 47

Below and behind the orbitonasal opening the palatines lie closely apposed to the

parasphenoidal rostrum, so that the midline region is completely floored by bone.

Lower jaw

The symphyseal region, usually powerfully developed in dicynodonts, is

relatively weak in Myosaurus (Figs 1C, 2B). There are no dentary tables and the

very slightly concave dorsal dentary surface is scooped out in front to form a

deep cleft behind the anterior shovel-shaped tip. This anterior edge is not well

preserved in any specimen, but indications are that it was at least fairly sharp.

The weak development of the dentaries is reflected in the considerable exposure

of the splenials in anterior view.

A fairly prominent lateral dentary shelf is formed in front of and above the

mandibular fenestra, while the rear half of the dentary’s dorsal surface, behind

the slightly grooved anterior portion, is narrow and smoothly rounded. The

anterior surface of the dentary, below the shovel edge, is heavily pitted by

nutritive foramina, and indications are that a horny sheath covered this front

part of the lower jaw. No such pitting can be detected on the dorsal edges of the

dentary, where clear evidence of a horn covering is usually found in dicynodonts.

The jaw is deepest in the angular region, even when the extensive reflected

lamina of the angular is not taken into consideration. In the rear of the jaw the

articular is seen to be drawn down ventrally into a long and powerful retro-

articular process, but little else, especially of the inner surfaces, can be made out.

JAW FUNCTION IN MYOSAURUS

The jaw musculature of dicynodonts has been reconstructed by Crompton

& Hotton (1967). From subsequent comparisons (Cluver 1970, 1971) it appears

that the area of greatest variation in this system lies in the nature of the insertion

of the external adductor muscles on the lower jaw. Two divisions of the external

adductor musculature were present in dicynodonts, these being a medial external

adductor arising from the bones surrounding the temporal fenestra, and a

lateral external adductor arising from the anterior surface of the descending

squamosal sheet and the ventral surface of the squamosal’s zygomatic bar.

Crompton & Hotton (1967) suggest that in dicynodonts such as Lystrosaurus

and Emydops the medial mass inserted into a groove in the dorsal surface of the

dentary behind the dentary tables, while the lateral mass was attached to a

lateral dentary shelf lying above and partly in front of the mandibular fenestra.

However, it appears that in certain genera (Cluver 1970, 1971) the groove and

shelf are absent or present only in modified form, and it has been suggested

(Cluver 1974) that an additional slip of the external adductor muscle inserted

around the dorso-lateral margin of the adductor fossa, on the inner side of the

jaw ramus.

In Myosaurus no dorsai dentary groove is found, but a prominent lateral

dentary shelf is present, and the dentary is powerfully buttressed over the

48 ANNALS OF THE SOUTH AFRICAN MUSEUM

mandibular fenestra. The lateral external adductor muscle was therefore pro-

vided with a well-defined area of attachment, but the limits of the insertion

area of the medial group are less clear; this muscle probably attached on the

inner surface and the rounded dorsal edge of the dentary, behind its horn-

covered anterior portion.

The possible range of jaw movements in small edentulous dicynodonts

such as Cistecephalus have been recently discussed (Cluver 1974) and it has

been shown that the typically wide cistecephalid palate, narrow symphyseal

region and relatively flat quadrate and articular articulation surfaces could

indicate lateral movement across the palate during the anteroposterior cycle

of jaw movements as described in dicynodonts by Crompton & Hotton (1967).

In this respect, however, Myosaurus appears to have retained the primitive

range of movements: the articular surfaces of the quadrate are at sharp angles

to each other and quadrate-articular rotation appears to have been highly

unlikely. In addition, the long and narrow secondary palate, with rim and canini-

form processes hardly wider than the symphyseal area of the lower jaw, suggests

that upper and lower apposing surfaces would have met during strictly antero-

posterior sliding movements.

The nature of these occluding surfaces, however, is unusual. As shown

above, the dentaries are weakly developed anteriorly, and there are no dentary

tables such as appose the palatine and maxilla in Lystrosaurus and Dicynodon

(Cluver 1970, 1971). At the same time, a battery of mandibular teeth, which

bite against the palatine in forms such as Emydops, is absent, and the apparent

lack of a horn covering in the palatine region, as well as the delicate structure

of that bone, appears to be a consequence of this.

The only clear evidence of horn is found on the anterior surface of the

fused dentaries, and points to the presence of a horny sheath over the sharp-

edged, square tip of the symphysis. Behind this the lateral edge of the dentary,

bordering the slightly concave dorsal surface, is fairly sharp and may have

carried a posterior extension of the beak. While the shovel-like tip would have

acted against the anterior premaxillary surface, as suggested by the absence of

anterior palatal ridges, the more lateral parts of the lower beak would have

had some slicing effect against the palatal rim, which, in turn, was probably

furnished with a corresponding horny layer. It seems unlikely, in view of the

smooth bone surfaces involved, that either of these lateral horn coverings were

substantial or extended as far medially as the median palatal ridge with its pair

of flanking channels.

The dietary preferences of an animal such as Myosaurus can only be

speculated upon. Certainly, the lack of well-developed shearing and crushing

areas in the mouth suggests a food source very different to the presumably

fairly coarse vegetation for which the contemporary Lystrosaurus was equipped.

Cox (1972) has recently suggested that the small cistecephalid Kawingasaurus

fed upon soft-bodied, and presumably slow-moving, invertebrates, and a

roughly similar diet in the case of Myosaurus cannot be ruled out.

CRANIAL MORPHOLOGY OF TRIASSIC DICYNODONT MYOSAURUS GRACILIS 49

med. pal. r

pal.

Fig. 5. A. Emydops longus paratype B.M.(N.H.) R4957. Palate, x 2. B-C.

Myosauroides minnaari type R.C.54, x 2. Anterior palate and dorsal view of

mandible respectively.

COMPARISON WITH OTHER DICYNODONTS

At present, three genera especially appear to have significant resemblances,

and possible affinities, with Myosaurus. These are Emydops, Cistecephalus and

Myosauroides.

The Upper Permian Emydops (Fig. SA) has been mentioned as a possible

Myosaurus ancestor by Haughton (1917) and, although the genus has many

clearly primitive characters, it does include features which deserve consideration.

These are a definite palatine-premaxilla contact, a long narrow interpterygoidal

vacuity, and a thin, steep vomerine septum dividing the internal nares. The

palatine, in contrast to Myosaurus, is still well developed and received the bite

of the lower jaw teeth during mastication, but in the upper jaw teeth are vestigial

and apparently of no great functional significance. In the skull roof the propor-

tions of intertemporal to interorbital width are close to those of Myosaurus,

but the postorbital is not flared laterally above the temporal fenestra and instead

forms the vertical border of the intertemporal bar. A large postfrontal, lost in

the more progressive smaller genera, is still present, as is the preparietal. The

lower jaw has a row of functional teeth on the medial edge of the dentary, but

anteriorly no dentary table is formed and a shovel-like tip, basically similar to

that of Myosaurus, is formed.

50 ANNALS OF THE SOUTH AFRICAN MUSEUM

Clearly, Emydops itself should not be designated a direct ancestor of

Myosaurus, but it does none the less possess several characters expected in

such an ancestor, and it seems very likely that a more detailed knowledge of

Emydops and allied forms would reveal a more definite picture of the origins

of later groups such as Myosaurus.

Fig. 6. Cistecephalus sp. S.A.M. No. 10665 x #. A. Ventral

view of skull. B. Dorsal view of mandible.

The genus Cistecephalus has recently been reviewed by Keyser (1973) and

discussed by Cluver (1974), and the skull morphology is now well known

(Fig. 6). While there are obvious adaptations to a specialized way of life, several

resemblances to Myosaurus exist. A postfrontal is absent, as in Myosaurus,

but the exceptionally wide intertemporal sheet lacks a preparietal. Ventrally

CRANIAL MORPHOLOGY OF TRIASSIC CIDYNODONT M YOSAURUS GRACILIS 5]

the secondary palate has very much the same organization as Myosaurus, and

anterior palatal ridges are absent and the maxilla is confined to a lateral posi-

tion. The palatines are small and, as in Myosaurus, each is pierced by a foramen.

In the lower jaw the dentaries are drawn up anteriorly to a transverse blade,

fairly sharp, while laterally a prominent dentary shelf is formed.

Here, however, the similarities cease. In Cistecephalus the interpterygoidal

vacuity has been obliterated by reinforcement of the basicranial girder, and the

vomer is a deep and wide plate with its ventral edge level with the ventral

borders of the pterygoids. Anteriorly, two pairs of pockets, one lateral and one

medial to the palatal rim, lie at the level of the weak caniniform process; these

pit-like depressions seem to be unique to Cistecephalus. A wide orbitonasal

opening leads into the interior of the snout, while the lacrimal foramen (if

present) is unusually placed near the border of this opening. The lower jaw,

again, is more powerfully developed in Cistecephalus than in Myosaurus, and

the deep symphyseal region is characteristically scooped out behind the square,

shovel-like tip; the dorsal surface of each dentary is widely expanded behind

this, and very different to the narrow, weakly developed dentary ramus of

Myosaurus. When, in addition to these factors, the perforated stapes and

secondary widening of the skull roof are taken into consideration, it becomes

clear that descent of a form such as Myosaurus from this genus is highly improb-

able, even although certain apparently significant resemblances between the

two genera exist.

Myosauroides, the third genus which can be conveniently compared with

Myosaurus, shows resemblances to both that genus and Cistecephalus (Fig. 5B,

C). The intertemporal region is of the same width as in Myosaurus and also

lacks a postfrontal, while a ventral flange of the frontal extends medially in a

manner reminiscent of Myosaurus. Anterior palatal ridges are lacking, the

palatine is weak, and the interpterygoidal vacuity and delicate vomerine septum

are almost identical to the Myosaurus condition. The jaw symphysis has the same

sharply squared-off tip, and the stapes is the usual thin, unperforated rod.

Myosauroides has a fairly prominent caniniform process and a higher palatal

rim than Myosaurus, but this rim is continued posteriorly from the caniniform

process for a short way as a postcaniniform crest, similar to that of Myosaurus.

Differences from Myosaurus, which are at the same time resemblances to-

Cistecephalus, are the broad, grooved dorsal dentary surfaces, the lack of a

preparietal, and the generally still wide orbitonasal passage.

Of the three genera, Myosauroides appears to be the closest to Myosaurus,

and it is quite conceivable that, with the exception of the preparietal, charac-

teristics of Myosaurus such as the highly reduced orbitonasal passage and the

weak dentary ramus could have been derived from the Myosauroides condition.

In terms of jaw function, too, Myosaurus and Myosauroides appear to be

broadly similarly adapted, and there is no evidence in Myosauroides of lateral

jaw displacement during mastication such as is found in Cistecephalus (see

above). The quadrate-articular surfaces are tightly interlocked and the secondary

m3 ANNALS OF THE SOUTH AFRICAN MUSEUM

palate is not significantly wider than the symphyseal region of the lower jaw.

However, the lack of a preparietal precludes Myosauroides minnaari itself from

direct ancestry of Myosaurus.

Viewed generally, the three genera Myosaurus, Myosauroides and Cistece-

phalus appear to be united by fairly clear common characteristics, with Myosaurus

and Myosauroides more closely related to each other than either is to the strongly

divergent Cistecephalus. If these genera are indeed part of a natural group, this

is not reflected in any present classification, where Cistecephalus and Myosaurus,

for instance, are usually placed in separate families (Haughton & Brink 1954;

Boonstra 1972; Cluver 1974). In view of the fact, however, that a common

origin of the group cannot as yet be convincingly established, and as our

knowledge of the many Permian pristerodontinid genera is still very incomplete,

it seems unwise to attempt any large-scale classificatory reorganization of the

endothiodontids at present.

SUMMARY

The skull of Myosaurus gracilis shows several departures from the usual

dicynodont condition, especially in the otic capsule, snout, secondary palate

and mandible. Perhaps in keeping with its lone survival into the early Triassic,

most of these modifications are connected with mastication, and seem to

reflect a dietary change. Myosaurus shares such features as a broad inter-

temporal region, lack of a postfrontal, the small palatine and laterally confined

maxilla, as well as the absence of anterior palatal ridges and the presence of a

square-tipped, shovel-shaped jaw symphysis, with Upper Permian genera such

as Myosauroides and Cistecephalus. While there is a possibility of these three

genera belonging to a single natural group, their relationships with the numerous

and generally poorly-known Permian endothiodontids remain obscure, and a

reorganization of the classification of the Endothiodontidae has not been

attempted. Among themselves, Myosaurus and Myosauroides show the closest

relationship, while the more aberrant Cistecephalus is strongly specialized in

respect of, especially, the skull roof, basicranial axis and palate-mandible

relationships.

ACKNOWLEDGEMENTS

I am indebted to Dr J. W. Kitching of the Bernard Price Institute for

Palaeontological Research, Johannesburg, for the loan of specimens of

Myosaurus, and to Mr Richard Rubidge of Wellwood, Graaff-Reinet, for

permission to prepare and study the type of Myosauroides minnaari. A second

specimen of the latter species was lent to me through the kindness of Dr Eugene

S. Gaffney of the American Museum of Natural History, New York, while the

type and paratype of Emydops longus were prepared and studied at the British

CRANIAL MORPHOLOGY OF TRIASSIC DICYNODONT MYOSAURUS GRACILIS 53

Museum (Natural History) through the kind permission of Dr A. J. Charig.

Visits to these two overseas institutions were made possible through study

grants from the South African Council for Scientific and Industrial Research,

and the Trustees of the South African Museum.

The beautifully prepared type skulls of Myosaurus gracilis are testimony

to the exceptional skill of Mrs Ione Rudner of the South African Museum.

Mr N. J. Eden was responsible for the photography, and Mr V. Branco

prepared the final drawings.

REFERENCES

Boonstra, L. D. 1972. Discard the names Theriodontia and Anomodontia: a new classifica-

tion of the Therapsida. Ann. S. Afr. Mus. 59: 315-338.

Broom, R. 1926. On the mammalian presphenoid and mesethmoid bones. Proc. zool. Soc.

Lond. 1926: 257-264.

Broom, R. 1932. The mammal-like reptiles of South Africa and the origin of mammals. London:

Witherby.

Broom, R. 1941. Some new Karoo reptiles with notes on a few others. Ann. Transy. Mus.

20: 194-213.

CLUVER, M. A. 1970. The palate and mandible in some specimens of Dicynodon testudirostris

Broom & Haughton (Reptilia, Therapsida). Ann. S. Afr. Mus. 56: 133-153.

CLuver, M. A. 1971. The cranial morphology of the dicynodont genus Lystrosaurus. Ann. S.

Afr. Mus. 56: 155-274.

CLuver, M. A. 1974. The skull and mandible of a new cistecephalid dicynodont. Ann. S. Afr.

Mus. 64: 137-155.

Cox, C. B. 1959. On the anatomy of a new dicynodont genus with evidence of the position of

the tympanum. Proc. zool. Soc. Lond. 132: 321-367.

Cox, C. B. 1972. A new digging dicynodont from the Upper Permian of Tanzania. Jn: JoysEy,

K. A. & Kemp, T. S. eds. Studies in vertebrate evolution: 173-189. Edinburgh: Oliver &

Boyd.

Crompton, A. W. & HotTToN, N. 1967. Functional morphology of the masticatory apparatus

of two dicynodonts (Reptilia, Therapsida). Postilla 109: 1-51.

Denby, A. 1909. The intracranial vascular system of Sphenodon. Phil. Trans. R. Soc. (B)

200: 403-426.

HauGuton, S. H. 1917. Descriptive catalogue of the Anomodontia, with special reference to

the examples in the South African Museum (Part 1). Ann. S. Afr. Mus. 12: 127-174.

HAUGHTON, S. H. & Brink, A. S. 1954. A bibliographic list of Reptilia from the Karroo beds .

of Africa. Palaeont. afr. 2: 1-187.

Keyser, A. W. 1973. A preliminary study of the type area of the Cistecephalus zone of the

Beaufort Series, and a revision of the anomodont family Cistecephalidae. Mem. geol.

Surv. S. Afr. 62: 1-71.

Otson, E. C. 1944. The origin of mammals based on the cranial morphology of the therapsid

suborders. Spec. Pap. geol. Soc. Am. 55: 1-136.

Romer, A. S. 1966. Vertebrate paleontology. Chicago: University of Chicago Press.

SWAIN, R. 1968. The pineal vascular system in Lacerta muralis, with notes on the venous

system of other lizards. J. Zool. 154: 487-493.

TOERIEN, M. J. 1953. The evolution of the palate in South African Anomodontia and its

classificatory significance. Palaeont. afr. 1: 49-117.

VAN Hoepen, E. C. N. 1934. Oor die indeling van die Dicynodontidae na aanleiding van nuwe

vorme. Paleont. Navors. nas. Mus. Bloemfontein 11: 67-101.

ang.

art.

bas.

boc.

can. proc.

den.

den. gr.

ect.

e0c.

ept:

for.

for. mag.

fir

gr.

1.C.a.

ip.

ipt. vac.

jug.

lac.

lat. fl.

lat. pal. for.

max.

med. pal. r.

nas.

no.

nut. ch.

op.

OSs.

pa.

pal.

ANNALS OF THE SOUTH AFRICAN MUSEUM

ABBREVIATIONS

angular pal. for.

articular pas.

basisphenoid Pe: cE:

basioccipital pfr.

caniniform process pm.

dentary po.

dentary gr. pp.

ectopterygoid prs.

exoccipital pt.

epipterygoid Pet

foramen q.

foramen magnum q. for

frontal lor

groove Sa.

internal carotid artery opening soc.

interparietal sq.

interpterygoidal vacuity st.

jugular t.

lacrimal tab.

lateral flange of squamosal A.M.N.H.

lateral palatal foramen

maxilla B.M.(N.H.)

medial palatal ridge on

premaxilla B.P.I.

nasal

notch

nutrient channel R.C.

opisthotic

orbitosphenoid S.A.M.

parietal

palatine

palatine foramen

parasphenoid

postcaniniform crest

prefrontal

premaxilla

postorbital

preparietal

presphenoid

pterygoid

posttemporal fenestra

quadrate

quadrate foramen

quadratojugal

surangular

supraoccipital

squamosal

stapes

tooth

tabular

American Museum of Natural

History, New York

British Museum (Natural

History), London

Bernard Price Institute for

Palaeontological Research,

Johannesburg

Rubidge Collection, Graaff-

Reinet

South African Museum, Cape

Town

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For journal articles give title of article, title of journal in italics (abbreviated according to the

World list of scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses,

volume number, part number (only if independently paged) in parentheses, pagination.

Examples (note capitalization and punctuation)

BuLtLoucu, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FIscHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris

88: 100-140.

FISCHER, P.-H., DUvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des

littorines. Archs Zool. exp. gén. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee

region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19605. Spawning behaviour, egg masses and larval development in Conus from the

Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51. 2

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn: SCHULTZE, L.

Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und

zentralen Siid-Afrika. 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16:

269-270.

ZOOLOGICAL NOMENCLATURE

To be governed by the rulings of the latest International code of zoological nomenclature

issued by the International Trust for Zoological Nomenclature (particularly articles 22 and

51). The Harvard system of reference to be used in the synonymy lists, with the full

references incorporated in the list at the end of the article, and not given in contracted form

in the synonymy list.

Example

Scalaria coronata Lamarck, 1816: pl. 451, figs 5 a, b; Liste: 11. Turton, 1932: 80.

Michael A. Cluver

THE CRANIAL MORPHOLOGY

OF THE LOWER TRIASSIC DICYNODONT

MYOSAURUS GRACILIS

50O/.E8

VOLUME 66 PART 4 NOVEMBER 1974

ANNALS

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 66 Band

November 1974 November

Part 4 Deel

feEE SPECIMENS OF DECAPODA (CRUSTACEA)

PNoInE COLLECTIONS OF IHE

SOUTH AFRICAN MUSEUM

BRIAN KENSLEY

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

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TYPE SPECIMENS OF DECAPODA (CRUSTACEA)

IN THE COLLECTIONS OF THE SOUTH AFRICAN MUSEUM

BRIAN KENSLEY

South African Museum, Cape Town

[MS accepted 15 October 1973]

CONTENTS

PAGE

Introduction : : : ; : ‘ : 55

Historical discussion ; g : " : 55)

List of type specimens. : 2 : : 56

References ; : ; : ‘ ; ; 76

Index : ; . ; E y : ; 78

INTRODUCTION

According to the recommendation 72D of the International Code of

Zoological Nomenclature (1964), institutions holding type material should

publish lists of their type holdings. This is the first of the crustacean type lists

of the holdings of the South African Museum, and has been consciously

modelled on a publication of the British Museum (Natural History) (Thurston

& Allen 1969), as it was felt that the format of the latter was particularly concise

and lucid.

The name under which a species was first described is placed in square

brackets if this name has since been submerged. The current name is then placed

in round brackets, following the original reference.

If a valid species has been transferred from the original to another genus,

the current name and reference is followed by the original name in round

brackets.

Where one of several specimens of a species from the same locality was

sent to T. R. R. Stebbing, who subsequently described it as a new species,

the specimens which he did not see are designated as topotypes.

Where a species was described from more than one specimen (even if from

a single male and a single female) without a holotype being chosen, the specimens

are designated as syntypes.

HISTORICAL DISCUSSION

Much the largest number of undescribed decapods were collected by the

S.S. Pieter Faure. This was a steam-trawler purchased by the Government of

the Cape Colony in 1897, to assist with the first Marine Biological Survey. The

survey covered the area from St. Helena Bay in the west to East London and,

later, from Cape Vidal in Zululand to the south coast of Natal.

5/5)

Ann S. Afr. Mus. 66 (4), 1974: 55-80.

56 ANNALS OF THE SOUTH AFRICAN MUSEUM

The S.S. Pickle, a vessel of the Government of the Union of South Africa

conducted a marine survey of Mocambique in 1925-6, the results of which were

reported on by K. H. Barnard in 1926.

Some undescribed material was obtained by the R/V Africana II, largest

of the Division of Sea Fisheries research ships.

Material from the south-west Indian Ocean was obtained on the seventh

cruise of the United States National Science Foundation R/V Anton Bruun,

while participating in the International Indian Ocean Expedition in 1964.

The zoology departments of two South African universities have also

contributed to the list of new decapod species. These are the University of the

Witwatersrand, and the University of Cape Town. The latter has covered much

of the coastline of South Africa during surveys of the shore and estuarine fauna,

while the operations of the University of Cape Town research ship John D.

Gilchrist have also yielded undescribed species.

Individual collectors have also contributed to the fauna list. These include

J. D. Gilchrist, Government Marine Biologist during the early part of this

century, K. H. Barnard, of the South African Museum, and H. W. Bell-Marley

of Natal.

CRUSTACEA

DECAPODA

REPTANTIA

BRACHYURA

Family Majidae

Subfamily Inachinae

ACHAEUS Leach

Achaeus barnardi Griffin, 1968: 81.

Holotype: S.A.M. A1392 (male; chelipeds and pereiopods missing; in

alcohol).

Paratypes: S.A.M. A1392 (2 males, | ovigerous female; most pereiopods

detached; in alcohol).

Locality: Off Cape Morgan, Cape Province, 36 fathoms.

Collected by: S.S. Pieter Faure.

Achaeus spinosissimus Griffin, 1968: 76.

Holotype: S.A.M. A8309 (male; most pereiopods detached; in alcohol).

Locality: Off Hood Point, Cape Province, 49 fathoms.

Collected by: S.S. Pieter Faure.

TYPE SPECIMENS OF DECAPODA (CRUSTACEA) =

PLATYMAIA Miers

Platymaia turbynei Stebbing, 1902: 3.

Syntypes: S.A.M. A1358-A1362 (15 females, 23 males; numerous loose

appendages; in alcohol).

Locality: Off Durban, Natal, 440 fathoms.

Collected by: S.S. Pieter Faure.

DEHAANIUS Macleay

Dehaanius undulatus Barnard, 1947: 361.

Syntypes: S.A.M. A3131 (2 females; 1 perfect, 1 with all pereiopods

detached; dry). S.A.M. A6353 (1 male; carapace broken, 1 cheliped

present; dry). S.A.M. A6376 (1 female; all pereiopods detached; dry).

Localities: Durban, Natal; Delagoa Bay, Mocambique.

Collected by: H. W. Bell-Marley, K. H. Barnard.

Subfamily Acanthonychinae

MENAETHIOPS Alcock

Menaethiops delagoae Barnard, 1955: 13.

Holotype: S.A.M. A10906 (male; dry).

Locality: Delagoa Bay, Mocambique.

Collected by: University of the Witwatersrand.

Menaethiops natalensis Barnard, 1955: 13.

Syntypes: S.A.M. A10904 (male; dry). S.A.M. A10905 (1 male, 2 females;

dry).

Localities: S.A.M. Al10904 Umpangazi, Natal. S.A.M. A10905 Delagoa

Bay, Mocambique.

Collected by: University of Cape Town, University of the Witwatersrand.

Subfamily Pisinae

EURYNOME Leach

Eurynome elegans Stebbing, 1921: 454.

Holotype: S.A.M. A1610 (female; most appendages detached; dry).

Locality: Cape Vidal, Natal NNEAN, 15 km, 80 fathoms.

Collected by: S.S. Pieter Faure. .

Family Hymenosomatidae

RHYNCHOPLAX Stimpson

Rhynchoplax bovis Barnard, 1947: 362.

Syntypes: S.A.M. A8454 (male; pereiopods missing; in alcohol). S.A.M.

A8336 (3 females; most appendages detached; in alcohol).

58 ANNALS OF THE SOUTH AFRICAN MUSEUM

Localities: S.A.M. A8454, 25 km west of East London, Cape Province,

37 fathoms. S.A.M. A8336, Port Elizabeth, Cape Province.

Collected by: S.S. Pieter Faure.

Family Retroplumidae

RETROPLUMA Gill

Retropluma planiforma Kensley, 1969: 158.

Holotype: S.A.M. A12644 (female; 3 pereiopods missing; in alcohol).

Paratypes: S.A.M. A12643, A12645 (2 males, 3 females; several appendages

missing or detached; in alcohol).

Locality: 29.378, 31.33E, 175-200 metres.

Collected by: R/V Anton Bruun, 7th Cruise.

Family Palicidae

PALICUS Philippi

Palicus sexlobata Kensley, 1969: 156.

Holotype: S.A.M. A12642 (male; 1 chela, 2 pereiopods detached, rest

missing; in alcohol).

Locality: 24.46S, 35.18E, 110 metres.

Collected by: R/V Anton Bruun, 7th Cruise.

Family Ocypodidae

CLEISTOSTOMA de Haan

Cleistostoma algoense Barnard, 1954: 122.

Syntypes: S.A.M. A850 (1 ovigerous female; carapace, most pereiopods

detached; in alcohol). S.A.M. A8493 (1 ovigerous female; 3 males; all

pereiopods detached; dry).

Locality: Zwartkops Estuary, near Port Elizabeth, Cape Province.

Collected by: University of Cape Town.

PARACLEISTOSTOMA de Man

Paracleistostoma fossula Barnard, 1955: 24.

Holotype: S.A.M. A10778 (ovigerous female; all pereiopods detached;

in alcohol).

Locality: Delagoa Bay, Mocambique.

Collected by: University of the Witwatersrand.

TYPE SPECIMENS OF DECAPODA (CRUSTACEA) 59

Family Grapsidae

ILYOGRAPSUS Barnard

[lyograpsus rhizophorae] Barnard, 1955: 26. (Transferred to J. paludicola

(Rathbun) by Crosnier, 1965.)

Holotype: S.A.M. A10913 (male; all pereiopods detached; in alcohol).

Locality: Inhambane, Mocambique.

Collected by: University of Cape Town.

Family Portunidae

LUPOCYCLUS Adams & White

Lupocyclus tugelae Barnard, 1950: 148.

Holotype: S.A.M. A8320 (female; 4 detached pereiopods; dry).

Locality: Off Tugela River mouth, Natal, 36 fathoms.

Collected by: S.S. Pieter Faure.

THALAMITA Latreille

Thalamita delagoae Barnard, 1950: 178.

Holotype: S.A.M. A10870 (male; all pereiopods detached; dry).

Locality: Delagoa Bay, Mocambique.

Collected by: University of the Witwatersrand.

[Thalamita inhacae| Barnard, 1950: 179. (Transferred to T. bouvieri Nobili, by

Crosnier, 1962.)

Holotype: S.A.M.A10871 (female; 3 pereiopods missing, rest detached; dry).

Locality: Inhaca Island, Delagoa Bay, Mocambique.

Collected by: University of the Witwatersrand.

XAIVA Macleay

Xaiva mcleayi (Barnard), 1947: 363 (as Portumnus mcleayi).

Syntypes: S.A.M. A631, A632 (3 males, 6 females; in alcohol).

Localities: Algoa Bay, Cape Province, to Port Shepstone, Natal.

Collected by: S.S. Pieter Faure.

Family Xanthidae

Section HYPEROMERISTA

DAIROIDES Stebbing

Dairoides margaritatus Stebbing, 1920: 234.

Syntypes: S.A.M. A1606 (2 males; 1 dry, carapace fragmented, | in alcohol).

60 ANNALS OF THE SOUTH AFRICAN MUSEUM

Locality: Durnford Point, Natal NM?W, 19 km, 90 fathoms.

Collected by: S.S. Pieter Faure.

LYBIA Milne Edwards

Lybia plumosa Barnard, 1947: 364.

Holotype: S.A.M. A10847 (female; all pereiopods detached; dry).

Locality: Umtwalumi, Natal.

Collected by: University of Cape Town.

Family Goneplacidae

HEXAPUS de Haan

Hexapus stebbingi Barnard, 1947: 366.

Syntypes: S.A.M. A434-7, A847-8, A1611 (5 ovigerous females, 13 females,

7 males; in alcohol).

Localities: S.A.M. A434-7, Sebastian Bay, Cape Province, 20-34 fathoms.

A847, 33.52S, 25.43E, 164 fathoms. A848, Riet Point, Cape Province,

NE by E, 3 km, 23 fathoms. Al611, Cape St. Blaize, Cape Province,

SW by W4W, 10 km, 15 fathoms.

Collected by: S.S. Pieter Faure.

OMMATOCARCINUS White

Ommatocarcinus pulcher Barnard, 1950: 286.

Holotype: S.A.M. A8322 (male; 6 pereiopods detached; in alcohol).

Locality: Natal coast, 28 fathoms.

Collected by: H. W. Bell-Marley.

THAUMASTOPLAX Miers

Thaumastoplax spiralis Barnard, 1950: 301.

Syntypes: S.A.M. A8487 (3 males, | female; in alcohol).

Locality: Off St. Helena Bay, Cape Province.

Collected by: S.S. Pieter Faure.

XENOPHTHALMODES Richters

Xenophthalmodes brachyphallus Barnard, 1955: 34.

Syntypes: S.A.M. A10825 (1 male, 1 female; dry).

Locality: Inhambane, Mocambique.

Collected by: University of Cape Town.

TYPE SPECIMENS OF DECAPODA (CRUSTACEA) 61

Family Dromiidae

CRYPTODROMIOPSIS Borradaile

Cryptodromiopsis bituberculata (Stebbing), 1920: 254 (as Eudromia bituberculata).

Holotype: S.A.M. A858 (female; carapace detached; in alcohol).

Locality: Off Rooi Els River, False Bay, Cape Province, 18 fathoms.

Collected by S.S. Pieter Faure.

Cryptodromiopsis lepidota Barnard, 1947: 369.

Holotype: S.A.M. A814 (female; 2 pereiopods detached; dry).

Locality: Off Hood Point Lighthouse, near East London, Cape Province,

49 fathoms.

Collected by: S.S. Pieter Faure.

DROMIDIA Stimpson

Dromidia aegibotus Barnard, 1947: 366.

Syntypes: S.A.M. A769 (1 male, 1 female; in alcohol). S.A.M. A770

(1 male, 1 female; in alcohol). S.A.M. A778 (1 male; in alcohol).

Localities: S.A.M. A769, False Is., Algoa Bay, Cape Province, 38 fathoms.

S.A.M. A770, False Bay, Cape Province. S.A.M. A778, Rocky Bank,

False Bay, Cape Province, 17-27 fathoms.

Collected by: S.S. Pieter Faure.

Dromidia dissothrix Barnard, 1947: 367.

Holotype: S.A.M. A879 (1 female; 7 pereiopods detached; dry).

Locality: Off Hout Bay, Cape Province.

Collected by S.S. Pieter Faure.

DROMIDIOPSIS Borradaile

Dromidiopsis cornuta Barnard, 1947: 367.

Syntypes: S.A.M. A856 (male; all pereiopods detached; dry). S.A.M.

A8237 (female; all pereiopods detached; dry).

Localities: S.A.M. A856, Cape St. Blaize, Cape Province, NE2?N, 16 km,

39 fathoms. S.A.M. A8237, False Bay, Cape Province, 23 fathoms.

Collected by: S.S. Pieter Faure.

EUDROMIDIA Barnard

Eudromidia hendersoni (Stebbing), 1921: 462 (as Eudromia hendersoni).

Holotype: S.A.M. A813 (female; carapace and pereiopods detached; dry).

Locality: False Bay, Cape Province, 19 fathoms.

Collected by: S.S. Pieter Faure.

62 ANNALS OF THE SOUTH AFRICAN MUSEUM

PETALOMERA Stimpson

Petalomera laevis Kensley, 1970: 111.

Holotype: S.A.M. A12824 (Ovigerous female; in alcohol).

Paratype: S.A.M. A12825 (male; in alcohol).

Locality: Jangamo, Mocgambique.

Collected by: University of Cape Town.

Family Dynomenidae

[MAXILLOTHRIX] Stebbing

[Maxillothrix actaeiformis] Stebbing, 1921: 457. (Transferred to Dynomene

pilumnoides Alcock, by Barnard, 1947.)

Syntypes: S.A.M. A839 (1 male, 3 females; most appendages detached; in

alcohol).

Locality: Umhlangakulu River, Natal, NW by N, 12 km, 50 fathoms.

Collected by: S.S. Pieter Faure.

Family Homolidae

HOMOLA Leach

Subgenus PAROMOLA Wood-Mason

Homola (Paromola) alcocki (Stebbing), 1920: 255 (as Latreillopsis alcocki).

Holotype: S.A.M. A1450 (female; carapace detached; in alcohol).

Locality: Algoa Bay, Cape Province, 40 fathoms.

Collected by: S.S. Pieter Faure.

Family Leucosiidae

EBALIA Leach

Ebalia agglomus Barnard, 1955: 39.

Holotype: S.A.M. A10739 (male; carapace fragmented; pereiopods

excluding chelipeds missing; dry).

Locality: Delagoa Bay, Mocambique.

Collected by: University of the Witwatersrand.

Ebalia (Lithadia) barnardi Stebbing, 1920: 248.

Holotype: S.A.M. A504 (female; all pereiopods detached; dry).

Locality: Umhloti River, Natal, NW by W3W, 4 km, 25 fathoms.

Collected by: S.S. Pieter Faure.

TYPE SPECIMENS OF DECAPODA (CRUSTACEA) 63

Ebalia glomus Stebbing, 1921: 460.

Syntypes: S.A.M. A503 (1 male, 1 female; all pereiopods detached; dry).

Locality: Umhloti River, Natal, NNW, 2 km, 27 fathoms.

Collected by: S.S. Pieter Faure.

[Nursia postulans| Stebbing, 1921: 461. (Transferred to Ebalia tuberculosa

(M. Edw.) by Barnard, 1950.)

Holotype: S.A.M. A502 (1 male; carapace and pereiopods detached; dry).

Locality: Cape Natal, W by N2N, 17 km, 185 fathoms.

Collected by: S.S. Pieter Faure.

HETERONUCIA Alcock

Heteronucia angulata Barnard, 1947: 372.

Holotype: S.A.M. A8210 (1 female; all pereiopods detached; dry).

Locality: Delagoa Bay, Mocgambique.

Collected by: University of the Witwatersrand.

Family Tymolidae

CORYCODUS Milne Edwards

Corycodus disjunctipes (Stebbing), 1910: 340 (as Nasinatalis disjunctipes).

Syntypes: S.A.M. A516, A517, (3 females; dry, in same container as

S.A.M. A1456 & A1609).

Locality: Cape Natal, NW4W, 9 km, 62 fathoms.

Collected by: S.S. Pieter Faure.

CYMONOMUS Milne Edwards

Cymonomus trifurcus Stebbing, 1920: 245.

Holotype: S.A.M. A884 (1 male; carapace and all appendages detached;

dry).

Locality: Buffalo River, Cape Province, NW4W, 30 km, 300 fathoms.

Collected by: S.S. Pieter Faure.

ETHUSA Roux

Ethusa sinespina Kensley, 1969: 161.

Holotype: S.A.M. A12648 (1 ovigerous female; pereiopods detached;

in alcohol).

Paratype: S.A.M. A12649 (1 female; 7 pereiopods missing; in alcohol).

Localities: A12648, 29.42S, 31.38E, 350 metres. A12649, 29.35S, 31.42E,

138 metres.

Collected by: R/V Anton Bruun, 7th Cruise.

64 ANNALS OF THE SOUTH AFRICAN MUSEUM

XEINOSTOMA Stebbing

Xeinostoma eucheir Stebbing, 1920: 243.

Syntypes: S.A.M. A1608 (1 male, 1 female; several pereiopods detached;

dry).

Locality: Cape Vidal, Natal, NNEN, 15 km, 90 fathoms.

Collected by: S.S. Pieter Faure.

ANOMURA

Family Lithodidae

NEOLITHODES Milne Edwards & Bouvier

Neolithodes asperrimus Barnard, 1947: 374.

Syntypes: S.A.M. A6371 (1 male; dry, 1 female; in alcohol).

Locality: Off Saldanha Bay, Cape Province, 300 fathoms.

Collected by: Trawler Ben Holden, Cape Town.

Neolithodes capensis Stebbing, 1905: 70.

Topotype: S.A.M. A8230 (1 ovigerous female; in alcohol).

Locality: Off Cape Point Cape Province, N 70° E, 64 km, 800 fathoms.

Collected by: ?

Family Pomatochelidae

POMATOCHELES Miers

Pomatocheles balssi Stebbing, 1914: 3.

Holotype: S.A.M. A1571 (1 male; carapace and appendages detached; dry).

Locality: Cove Rock, Cape Province, NW3W, 21 km, 80-130 fathoms.

Collected by: S.S. Pieter Faure.

Family Paguridae

ANAPAGURUS Henderson

Anapagurus hendersoni Barnard, 1947: 377.

Syntypes: S.A.M. A1500-03, (7 males, 1 female, 5 ovigerous females;

in alcohol). S.A.M. A8225-6 (1 male; in alcohol).

Localities: S.A.M. A1500, Bird Island passage, Cape Province, 10 fathoms.

A1501, Cape Natal, W by N, 10 km, 48 fathoms. A1502, Umhloti River,

Natal, NW3W, 24 km, 100 fathoms. A1503, Cape Natal, W by N, 10 km,

54 fathoms. A8225, East London, Cape Province, NW3N, 32 km,

400 fathoms.

Collected by: S.S. Pieter Faure.

TYPE SPECIMENS OF DECAPODA (CRUSTACEA) 65

CANCELLUS Milne Edwards

Cancellus makrothrix Stebbing, 1924: 6.

Topotype: S.A.M. A1541 (1 male; in alcohol).

Locality: Bird Island passage, Cape Province, 10 fathoms.

Collected by: S.S. Pieter Faure.

DIOGENES Dana

Diogenes extricatus Stebbing, 1910: 355.

Holotype: S.A.M. A1491 (male; in alcohol).

Locality: Off Seal Island, Mossel Bay, Cape Province.

Collected by: S.S. Pieter Faure.

PAGURISTES Dana

Paguristes barnardi Forest, 1954: 208.

Syntypes: S.A.M. A12779 (3 males; in alcohol). S.A.M. A12780 (2 males,

1 ovigerous female; in alcohol).

Localities: S.A.M. A12779, Agulhas, Cape Province. S.A.M. A12780,

Arniston, Cape Province.

Collected by: University of Cape Town.

Paguristes engyops Barnard, 1947: 375.

Syntypes: S.A.M. A3255 (3 males, 1 female, 3 ovigerous females; in

alcohol).

Locality: Buffels Bay, Cape Province.

Collected by: K. H. Barnard.

Paguristes macrotrichus Forest, 1954: 211.

Holotype: S.A.M. A851 (female; in alcohol).

Locality: Umhlangakulu River, Natal, NW by N, 12 km, 50 fathoms,

Collected by: S.S. Pieter Faure.

[Paguristes rosaceus| Barnard, 1947: 375. (Transferred to Paguristes gamianus-

H. Milne Edwards by Forest, 1954.)

Syntypes: S.A.M. A8452 (3 ovigerous females; in alcohol).

Locality: Off Keurbooms River, Cape Province.

Collected by: K. H. Barnard.

PARAPAGURUS Smith

[Parapagurus bouvieri] Stebbing, 1910: 357. (Transferred to Parapagurus pilosi-

manus Smith, see Barnard, 1950.)

Syntype: S.A.M. A1524 (female; in alcohol).

66 ANNALS OF THE SOUTH AFRICAN MUSEUM

Locality: Buffalo River, Cape Province, NW4W, 30 km, 300 fathoms.

Collected by: S.S. Pieter Faure.

Parapagurus kilburni Kensley, 1973: 285.

Holotype: S.A.M. A13185 (male; in alcohol).

Paratypes: S.A.M. A13186 (4 males; in alcohol).

Locality: Off Durban, 270 metres.

Collected by: R. N. Kilburn, Natal Museum.

PYLOPAGURUS Milne Edwards & Bouvier

Pylopagurus liochele Barnard, 1947: 376.

Syntypes: S.A.M. A1543 (female; abdomen missing; in alcohol). S.A.M.

A4038 (male; abdomen and pereiopods detached; in alcohol).

Localities: A1543, Bird Island passage, Cape Province, 10 fathoms. A4038,

Cape Seal, Cape Province WS, 9 km, 37 fathoms.

Collected by: S.S. Pieter Faure.

Family Porcellanidae

PISIDIA Leach

Pisidia delagoae (Barnard), 1955: 40 (as Porcellana delagoae, see Haig, 1965).

Holotype: S.A.M. A10703 (1 ovigerous female; pereiopods detached; in

alcohol).

Locality: Delagoa Bay, Mocambique.

Collected by: University of the Witwatersrand.

Family Uroptychidae

HAPALOPTYX Stebbing

Hapaloptyx difficilis Stebbing, 1920: 263.

Holotype: S.A.M. A1440 (1 ovigerous female; carapace and appendages

detached; dry; 3 slides).

Locality: Scottburgh light house, Natal, NW by W, 13 km, 92 fathoms.

Collected by: S.S. Pieter Faure.

Family Galatheidae

MUNIDOPSIS Whiteaves

Munidopsis barnardi Kensley, 1968: 290.

Holotype: S.A.M. A12636 (female; in alcohol).

Paratypes: S.A.M. A10497, A10508 (4 males; in alcohol). S.A.M. A10465.

(2 males; in alcohol). S.A.M. A10485 (1 male, 1 female; in alcohol).

TYPE SPECIMENS OF DECAPODA (CRUSTACEA) 67

Locality: Off Cape Point, Cape Province, 2 708-3 038 metres.

Collected by: R/V Africana IT.

Munidopsis chacei Kensley, 1968: 288.

Holotype: S.A.M. A10470 (ovigerous female; in alcohol).

Locality: Off Cape Point, Cape Province, 2 745 metres.

Collected by: R/V Africana IT.

Family Axiidae

METICONAXIUS de Man

Meticonaxius longispina (Stebbing), 1920: 265 (as Axius longispina).

Holotype: S.A.M. A957 (1 slide plus fragments in alcohol).

Locality: Cape Morgan, Cape Province, NNW, 11 km, 52 fathoms.

Collected by: S.S. Pieter Faure.

CALOCARIS Bell

Calocaris (Calocaris) barnardi Stebbing, 1914: 9.

Syntypes: S.A.M. A1549 (2 females; dry, 1 female; in alcohol).

Locality: Cape Castle, E4N, 21 km, 120 fathoms.

Collected by: S.S. Pieter Faure.

Family Callianassidae

CALLIANASSA Leach

Callianassa adamas Kensley, 1974: 266.

Holotype: S.A.M. A12103 (male; in alcohol).

Allotype: S.A.M. A12103 (female; in alcohol).

Paratype: S.A.M. A10985 (male; in alcohol).

Localities: S.A.M. A12103, Orange River mouth, 10-35 metres. S.A.M.

A10985, Olifants River mouth.

Collected by: Diamond dredge Emerson—K.

Callianassa gilchristi Barnard, 1947: 379.

Syntypes: S.A.M. A942 (female; in alcohol). S.A.M. A6807 (2 males;

several pereiopods detached; in alcohol).

Locality: False Bay, Cape Province.

Collected by: S.S. Pieter Faure.

Callianassa kraussi Stebbing, 1900: 39.

Syntypes: S.A.M. A941 (4 specimens; dry, in same bottle as several from

other localities).

68 ANNALS OF THE SOUTH AFRICAN MUSEUM

Locality: Gordon’s Bay, Cape Province.

Collected by: S.S. Pieter Faure.

Callianassa natalensis Barnard, 1947: 379.

Holotype: S.A.M. A8339 (female; 7 pereiopods detached; in alcohol).

Locality: Natal coast, from stomach of rock cod.

Collected by: H. W. Bell-Marley.

Callianassa subterranea australis Kensley, 1974: 271.

Holotype: S.A.M. A13531 (male; in alcohol).

Allotype: S.A.M. A13532 (female; in alcohol).

Paratypes: S.A.M. A13533-4 (1 male, 1 female; in alcohol). S.A.M. A12103

(1 female; in alcohol).

Localities: S.A.M. A13531—4, Liideritzbucht, South West Africa. S.A.M.

A12103, Orange River mouth.

Collected by: University of Cape Town. Diamond dredge Emerson—K.

Family Upogebiidae

UPOGEBIA Leach

Upogebia assisi Barnard, 1947: 381.

Holotype: S.A.M. A4348 (1 female; pereiopods detached; in alcohol).

Locality: St. Francis Bay, Cape Province.

Collected by: S.S. Pieter Faure.

PALINURA

Family Palinuridae

PROJASUS George & Grindley

Projasus parkeri (Stebbing), 1902: 38 (as Jasus parkeri).

Holotype: S.A.M. A993 (male; in alcohol).

Topotypes: S.A.M. A994-6. (2 females, 2 ovigerous females, 3 males).

Locality: Buffalo River, Cape Province, N, 24 km, 310 fathoms.

Collected by: S.S. Pieter Faure.

PALINURUS Fabricius

Palinurus gilchristi Stebbing, 1900: 31.

Syntype: S.A.M. A970 (male; 8 periopods missing; in alcohol).

Topotype: S.A.M. A971 (male; 1 pereiopod missing; in alcohol).

Locality: False Bay, Cape Province.

Collected by: S.S. Pieter Faure.

TYPE SPECIMENS OF DECAPODA (CRUSTACEA) 69

Palinurus delagoae Barnard, 1926. (See Berry & Plante 1973.)

Neotype: S.A.M. A13179 (male; in alcohol).

Locality: Off Tongaat, Natal.

Collected by: P. Berry, Oceanographic Research Institute, Durban.

PALINUSTUS Milne Edwards

Palinustus mossambicus Barnard, 1926: 126.

Holotype: S.A.M. A10684 (male; 5 pereiopods missing; in alcohol).

Locality: Mocambique, 25.0S, 33.10E, 406 metres.

Collected by: S.S. Pickle.

Family Eryonidae

POLYCHELES Heller

Polycheles demani Stebbing, 1917: 28.

Paratype: S.A.M. A4343 (male; in alcohol).

Topotypes: S.A.M. A1004-10, off Cape Point, (2 males, 6 females; in

alcohol).

Locality: Cape Point, Cape Province, NE by ESE 67 km, | 645 metres.

Collected by: S.S. Pieter Faure.

WILLEMOESIA Grote

Willemoesia bonaspei Kensley, 1968: 294.

Holotype: S.A.M. A10543 (ovigerous female; chelipeds missing; in alcohol).

Paratypes: S.A.M. A10473 (2 males; in alcohol). S.A.M. A10509 (1 female,

2 males; in alcohol). S.A.M. A12637 (2 males; in alcohol).

Locality: Off Cape Point, Cape Province, 2 708-3 038 metres.

Collected by: R/V Africana IT.

NATANTIA

PENAEIDEA

Family Penaeidae

Subfamily Penaeinae

PENAEUS Fabricius

[Penaeus caeruleus] Stebbing, 1905: 77. (Transferred to Penaeus monodon

Fabricius, see Barnard, 1950.)

Syntypes: S.A.M. 1039 (3 specimens in bottle with A1040, A1041; in

alcohol).

Locality: Nahoon River, East London, Cape Province.

Collected by: S.S. Pieter Faure.

70 ANNALS OF THE SOUTH AFRICAN MUSEUM

PENAEOPSIS Bate

[Penaeopsis spinulicauda] Stebbing, 1914: 17. (Transferred to Metapenaeus

monoceros (Fabricius), see Barnard 1950.)

Syntypes: S.A.M. A2231 (2 specimens, in same bottle as A1203; in alcohol).

Locality: Durban Bay, Natal.

Collected by: K. H. Barnard.

Subfamily Solenocerinae

SOLENOCERA Lucas

Solenocera africanum Stebbing, 1917: 32.

Syntypes: S.A.M. A1213 (1 female, few appendages of male; in alcohol).

Locality: Sebastian Bluff (Agulhas Bank), Cape Province, NW2W, 13 km,

34 fathoms.

Collected by: S.S. Pieter Faure.

Solenocera algoense Barnard, 1947: 383.

Holotype: S.A.M. A1211 (male; in alcohol).

Locality: Nanquas Peak, Cape Province, N, 12 km, 50 fathoms.

Collected by: S.S. Pieter Faure.

Solenocera comatum Stebbing, 1915: 67.

Syntypes: S.A.M. A1217 (1 male; in alcohol). S.A.M. A1218 (1 female;

in alcohol).

Locality: Off East London, Cape Province, 43-50 fathoms.

Collected by: S.S. Pieter Faure.

HYMENOPENAEUS Smith

Hymenopenaeus (Haliporoides) triarthrus (Stebbing), 1914: 21 (as Haliporoides

triarthrus).

Syntypes: S.A.M. A3940 (3 specimens in bottle with several others; in

alcohol).

Locality: East London, Cape Province, NW4N, 29 km, 250-300 fathoms.

Collected by: S.S. Pieter Faure.

Subfamily Aristaeinae

PLESIOPENAEUS Bate

Plesiopenaeus nitidus Barnard, 1947: 383.

Syntypes: S.A.M. A1193, A1260 (3 females, 3 males; in alcohol).

Locality: Off Cape Point, Cape Province, 500-630 fathoms.

Collected by: S.S. Pieter Faure.

TYPE SPECIMENS OF DECAPODA (CRUSTACEA) 71

Family Sergestidae

SERGESTES Milne Edwards

[Sergestes gloriosus] Stebbing, 1905: 84. (Transferred to Sergestes prehensilis

Bate by Yaldwyn 1957.)

Syntypes: S.A.M. A1050 (4 specimens, along with several specimens from

the same and other localities; in alcohol).

Locality: Sandy Point, Cape Province, NW by W, 27 km, 300 fathoms.

Collected by: S.S. Pieter Faure.

ACETES Milne Edwards

Acetes natalensis Barnard, 1955: 43.

Syntypes: S.A.M. A11976 (2 males, 3 females; in alcohol, | slide of male

petasma).

Locality: Durban Bay, Natal.

Collected by: University of Cape Town.

CARIDEA

Family Stylodactylidae

STYLODACTYLUS Milne Edwards

Stylodactylus stebbingi Hayashi & Miyake, 1968: 595.

Holotype: S.A.M. A1285 (male; in alcohol).

Paratypes: S.A.M. A1284-6 (11 ovigerous females, 1 female, 2 males;

in alcohol).

Localities: S.A.M. A1284—5, Buffalo River, Cape Province, NW4W, 30 km,

300 fathoms. S.A.M. A1286 South Head, Cape Province, E by S&S,

24 km, 190 fathoms.

Collected by: S.S. Pieter Faure.

Family Oplophoridae

NOTOSTOMUS Milne Edwards

Notostomus auriculatus Kemp, ined. (see Barnard 1950).

Holotype: S.A.M. A1189 (female; in alcohol).

Locality: Off Cape Point, Cape Province, 800 fathoms.

Collected by: S.S. Pieter Faure.

Family Hippolytidae

EUALUS Thallwitz

Eualus ctenifera (Barnard), 1950: 696 (as Spirontocaris ctenifera).

Syntypes: S.A.M. A6809 (1 ovigerous female; in alcohol). S.A.M. A8402

72 ANNALS OF THE SOUTH AFRICAN MUSEUM

(2 ovigerous females, 2 females, 2 males; in alcohol). S.A.M. A8413

(1 ovigerous female; in alcohol).

Localities: S.A.M. A6809, northern Natal coast, 70 fathoms. S.A.M.

A8402, 33.49S, 25.56E S.A.M. A8413, Durban, Natal.

Collected by: H. W. Bell-Marley, and S.S. Pieter Faure.

Eualus pax (Stebbing), 1915: 91 (as Spirontocaris pax).

Allotype: S.A.M. A3944 (ovigerous female; in alcohol).

Locality: Buffels Bay, Cape Province, 30 fathoms.

Collected by: S.S. Pieter Faure.

HIPPOLYTE Leach

Hippolyte palliola Kensley, 1970: 183

Holotype: S.A.M. A12710 (male; in alcohol).

Allotype: S.A.M. A12711 (ovigerous female; in alcohol).

Paratypes: S.A.M. A12712 (11 males; in alcohol).

Locality: 8 km south of Kunene River mouth, S.W.A.

Collected by: South African Museum and State Museum, Windhoek.

HIPPOLYSMATA Stimpson

Hippolysmata (Exhippolysmata) tugelae Stebbing, 1915: 94.

Syntype: S.A.M. A1274 (1 specimen, in same container as topotypes; in

alcohol).

Topotypes: S.A.M. A1274 (21 specimens; in alcohol).

Locality: Off south head of Tugela River, Natal, 12 fathoms.

Collected by: S.S. Pieter Faure.

LEBBEUS White

Lebbeus saldanhae (Barnard), 1947: 385 (as Spirontocaris saldanha).

Holotype: S.A.M. A8446 (female; in alcohol).

Locality: Off Constable Hill, Cape Province, 32 km, 145 fathoms.

Collected by: S.S. Pieter Faure.

LEONTOCARIS Stebbing

Leontocaris paulsoni Stebbing, 1905: 99.

Syntypes: S.A.M. A1166 (2 specimens in same container as S.A.M.

A1167-72; in alcohol).

Locality: Off Lions Head, Cape Province, 37 km.

Collected by: S.S. Pieter Faure.

TYPE SPECIMENS OF DECAPODA (CRUSTACEA) 73

Family Processidae

PROCESSA Leach

Processa austroafricana Barnard, 1947: 386.

Syntypes: S.A.M. A1097—-1104 (many specimens in alcohol).

Localities: A1097, Cape St. Blaize, Cape Province, 40 fathoms. A1098,

Algoa Bay, Cape Province, 10-16 fathoms. A1099, off Knysna Heads,

Cape Province. A1100, Cape St. Blaize, Cape Province, 40 fathoms.

A1101, off Struys Point, Cape Province, 42 fathoms. A1102, False Island,

Cape Province, 49 fathoms. A1103, off Great Fish Point lighthouse,

Cape Province, 40 fathoms. A1104, off Bird Island lighthouse, Cape

Province, 57 fathoms.

Collected by: S.S. Pieter Faure.

Family Glyphocrangonidae

GLYPHOCRANGON Milne Edwards

Glyphocrangon dentatus Barnard, 1947: 387.

Topotype: S.A.M. A8403 (male; dry).

Locality: Mogambique.

Collected by: S.S. Pickle.

Family Ogyrididae

OGYRIDES Stebbing

Ogyrides saldanhae Barnard, 1947: 387.

Syntypes: S.A.M. A1298 (70 specimens; in alcohol).

Locality: Saldanha Bay, Cape Province, 10 fathoms.

Collected by: S.S. Pieter Faure.

Family Alpheidae

ALPHEUS Fabricius

Alpheus bullatus Barnard, 1955: 45.

Syntype: S.A.M. A10624 (male; large chela detached; in alcohol).

Locality: Delagoa Bay, Mocambique.

Collected by: University of the Witwatersrand.

Alpheus dissodontonotus Stebbing, 1915: 83.

Holotype: S.A.M. A1561 (ovigerous female; carapace and pereiopods

detached; in alcohol).

Locality: 33.50S, 25.46E, 20 fathoms.

Collected by: S.S. Pieter Faure.

74 ANNALS OF THE SOUTH AFRICAN MUSEUM

Alpheus luciae Barnard, 1947: 389.

Holotype: S.A.M. A8423 (1 male; chelipeds detached; in alcohol).

Locality: St. Lucia Bay, Natal.

Collected by: H. W. Bell-Marley.

Alpheus nonalter Kensley, 1969: 172.

Holotype: S.A.M. A12650 (ovigerous female; in alcohol).

Allotype: S.A.M. A12651 (male; in alcohol).

Locality: 29.37S, 31.33E, 175-200 metres.

Collected by: R/V Anton Bruun, 7th Cruise.

Alpheus waltervadi Kensley, 1969: 175.

Holotype: S.A.M. A12646 (ovigerous female; in alcohol).

Paratype: S.A.M. A12647 (ovigerous female; in alcohol).

Locality: Walter’s Shoal, 33.13S, 43.51E, 38-46 metres.

Collected by: R/V Anton Bruun, 7th Cruise.

BETAEUS Dana

Betaeus jucundus Barnard, 1947: 388.

Holotype: S.A.M. A7807 (ovigerous female; in alcohol).

Locality: Keurbooms River, Cape Province.

Collected by: K. H. Barnard.

SALMONEUS Holthuis

Salmoneus rostratus Barnard, 1962: 240.

Syntypes: S.A.M. A11086 (2 ovigerous females; in alcohol).

Locality: Nosy Bé, Madagascar.

Collected by: University of the Witwatersrand.

SYNALPHEUS Bate

Synalpheus anisocheir Stebbing, 1915: 86.

Holotype: S.A.M. A1555 (ovigerous female; few appendages; in alcohol).

Locality: Gordon’s Bay, Cape Province.

Collected by: J. D. Gilchrist.

Family Palaemonidae

Subfamily Palaemoninae

LEANDER Desmarest

[Leander gilchristi] Stebbing, 1915: 75. (Transferred to Palaemon pacificus

(Stimpson) by Barnard 1950, 1955.)

Holotype: S.A.M. A3943 (ovigerous female; in alcohol).

TYPE SPECIMENS OF DECAPODA (CRUSTACEA) gis

Locality: East London, Cape Province.

Collected by J. D. Gilchrist.

[Leander peringueyi| Stebbing, 1915: 75. (Transferred to Palaemon pacificus

(Stimpson) by Barnard 1950, 1955).

Holotype: S.A.M. A1276 (ovigerous female; in alcohol).

EKocality: 33.49S, 25.56E.

Collected by: S.S. Pieter Faure.

PALAEMON Fabricius

[Palaemon delagoae| Stebbing, 1915: 74. (Transferred to Macrobrachium equidens

(Dana) by Holthuis 1950.)

Holotype: S.A.M. A2196 (male; chelipeds detached; in alcohol).

Locality: Delagoa Bay, Mocambique.

Collected by: A. W. Bayly.

Subfamily Pontoniinae

ISCHNOPONTONIA Bruce

Ischnopontonia lophos (Barnard), 1962: 242 (as Philarius lophos).

Holotype: S.A.M. A10652 (ovigerous female; chelipeds detached; in

alcohol).

Allotype: S.A.M. A12296 (male; carapace and chelipeds detached; in

alcohol).

Locality: Inhaca Island, Delagoa Bay, Mocambique.

Collected by: University of the Witwatersrand.

PERICLIMENAEUS Botradaile

Periclimenaeus uropodialis Barnard, 1958: 18.

Holotype: S.A.M. A10650 (females; carapace and several pereiopods

detached; in alcohol).

Locality: Delagoa Bay, Mocambique.

Collected by: University of the Witwatersrand.

PERICLIMENES Costa

Periclimenes (Periclimenes) delagoae Barnard, 1958: 14.

Holotype: S.A.M. A10648 (ovigerous female; carapace and most appen-

dages detached; in alcohol).

Locality: Delagoa Bay, Mocambique.

Collected by: University of the Witwatersrand.

76 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Crangonidae

METACRANGON Zarenkov

Metacrangon jacqueti bellmarleyi (Stebbing, 1914: 29) (as Sclerocrangon

bellmarleyi).

Syntypes: S.A.M. A1077 (male; in alcohol). S.A.M. A1564 (female; in

alcohol).

Locality: Cape Natal, N by E, 38 km, 440 fathoms.

Collected by: S.S. Pieter Faure.

PONTOPHILUS Leach

Pontophilus megalocheir (Stebbing), 1915: 71 (as Philocheras megalocheir).

Holotype: S.A.M. A1317 (female; carapace and most appendages detached;

in alcohol).

Locality: Off Cove Rock, East London, Cape Province, 25 fathoms.

Collected by: S.S. Pieter Faure.

REFERENCES

BARNARD, K. H. 1926. Report on a collection of Crustacea from Portuguese East Africa.

Trans. R. Soc. S. Afr. 13: 119-130.

BARNARD, K. H. 1947. Descriptions of new species of South African decapod Crustacea, with

notes on synonymy and new records. Ann. Mag. nat. Hist. (11) 13: 361-392.

BARNARD, K. H. 1950. Descriptive catalogue of South African decapod Crustacea (crabs and

shrimps). Ann. S. Afr. Mus. 38: 1-824.

BARNARD, K. H. 1954. New records and new species of Crustacea from South Africa. Annis

Mus. r. Congo belge Sér. 4to (N.S. Sci. zool.) 1: 120-131.

BARNARD, K. H. 1955. Additions to the fauna-list of South African Crustacea and Pycnogonida.

Ann. S. Afr. Mus. 43: 1-122.

BARNARD, K. H. 1958. Further additions to the crustacean fauna list of Portuguese East Africa.

Mems Mus. Dr Alvaro de Castro 4: 1-23.

BARNARD, K. H. 1962. New records of marine Crustacea from the East African region. Crusta-

ceana 3: 239-245.

BERRY, P. F. & PLANTE, R. 1973. Revision of the spiny lobster genus Palinurus, in the south-

west Indian Ocean. Trans. R. Soc. S. Afr. 40: 373-380.

Forest, J. 1954. Les Paguristes des cétes occidentales et méridionales d’Afrique. Ann. S. Afr.

Mus. 41: 159-213.

GriFFIN, D. J. G. 1968. Two new species of Achaeus (Crustacea, Decapoda, Majidae) from

South Africa. Ann. S. Afr. Mus. 52: 75-87.

HaiGc, J. 1965. Sur une collection de crustacés porcellanes (Anomura: Porcellanidae) de

Madagascar et des Comores. Cah. O.R.S.T.O.M. (Oceanogr.) 3 (4): 39-50.

Ho.tuuls, L. B. 1950. The Palaemonidae collected by the Siboga and Snellius Expeditions with

remarks on other species. 1. Subfamily Palaemoninae. Siboga Exped. monogr. 39a*:

1-268.

HAYASHI, K.-I. & MAYAKE, S. 1968. Notes on the family Stylodactylidae with the description

of a new genus Neostylodactylus. J. Fac. Agric. Kyushu Univ. 14: 583-611.

KENSLEY, B. F. 1968. Deep sea decapod Crustacea from west of Cape Point, South Africa.

Ann. S. Afr. Mus. 50: 283-323.

TYPE SPECIMENS OF DECAPODA (CRUSTACEA) fl

KENSLEY, B. F. 1969. Decapod Crustacea from the south-west Indian Ocean. Ann. S. Afr. Mus.

52: 149-181.

KENSLEY, B. F. 1970a. Some decapod Crustacea from northern South West Africa, including

a new species of Hippolyte. Cimbebasia (A) 1: 179-188.

KENSLEY, B. F. 19706. A small collection of decapod Crustacea from Mocgambique. Ann. S.

Afr. Mus. 57: 103-122.

KeENSLEY, B. F. 1974. The genus Callianassa (Crustacea, Decapoda, Thalassinidea) from the

west coast of South Africa with a key to the South African species. Ann. S. Afr. Mus. 62:

265-278.

STEBBING, T. R. R. 1900. South African Crustacea. Mar. Invest. S. Afr. 1: 14-66.

STEBBING, T. R. R. 1902. South African Crustacea, Part II. Mar. Invest. S. Afr. 2: 1-92.

STEBBING, T. R. R. 1905. South African Crustacea. Part III. Mar. Invest. S. Afr. 4: 21-120.

STEBBING, T. R. R. 1910. General catalogue of South African Crustacea. Ann. S. Afr. Mus. 6:

281-593.

STEBBING, T. R. R. 1914. South African Crustacea. Part VII. Ann. S. Afr. Mus. 15: 1-55.

STEBBING, T. R. R. 1915. South African Crustacea. Part VIII. Ann. S. Afr. Mus. 15: 57-104.

STEBBING, T. R. R. 1917. South African Crustacea. Part IX. Ann. S. Afr. Mus. 17: 23-46.

STEBBING, T. R. R. 1920. South African Crustacea. Part X. Ann. S. Afr. Mus. 17: 231-272.

STEBBING, T. R. R. 1921. South African Crustacea. Part XI. Ann. S. Afr. Mus. 18: 453-468.

STEBBING, T. R. R. 1924. South African Crustacea. Part XII. Ann. S. Afr. Mus. 19: 235-248.

THURSTON, M. H. & ALLEN, E. 1969. Type material of the families of Lysianassidae, Stegoce-

phalidae, Ampeliscidae, and Haustoriidae (Crustacea: Amphipoda) in the collections

of the British Museum (Natural History). Bull. Br. Mus. nat. Hist. (Zool.) 17: 349-388.

YALDWYN, J. C. 1957. Deep water Crustacea of the genus Sergestes (Decapoda Natantia)

from Cook Strait, New Zealand. Zoology Publs Vict. Univ. Wellington 22: 1-27.

INDEX

(Synonyms in italics)

Acetes, 71

Achaeus, 56

actaeiformis (Maxillothrix), 62

adamas (Callianassa), 67

aegibotus (Dromidia), 61

africanum (Solenocera), 70

agglomus (Ebalia), 62

alcocki (Homola), 62

alcocki (Latreillopsis), 62

algoense (Cleistostoma), 58

algoense (Solenocera), 70

Alpheus, 73, 74

Anapagurus, 64

angulata (Heteronucia), 63

anisocheir (Synalpheus), 74

asperrimus (Neolithodes), 64

assisi (Upogebia), 68

auriculatus (Notostomus), 71

australis (Callianassa subterranea), 68

austroafricana (Processa), 73

Axius, 67

balssi (Pomatocheles), 64

barnardi (Achaeus), 56

barnardi (Calocaris), 67

barnardi (Ebalia), 62

barnardi (Munidopsis), 66

barnardi (Paguristes), 65

bellmarleyi (Metacrangon jacqueti), 76

bellmarleyi (Sclerocrangon), 76

Betaeus, 74

bituberculata (Cryptodromiopsis), 61

bituberculata (Eudromia), 61

bonaspei (Willemoesia), 69

bouvieri (Thalamita), 59

bouvieri (Parapagurus), 65

bovis (Rhynchoplax), 57

brachyphallus (Xenophthalmodes), 60

bullatus (Alpheus), 73

caeruleus (Penaeus), 69

Callianassa, 67, 68

Calocaris, 67

Cancellus, 65

capensis (Neolithodes), 64

chacei (Munidopsis), 67

Cleistostoma, 58

comatum (Solenocera), 70

cornuta (Dromidiopsis), 61

Corycodus, 63

Cryptodromiopsis, 61

ctenifera (Eualus), 71

ctenifera (Spirontocaris), 71

Cymonomus, 63

Dairoides, 59

Dehaanius, 57

delagoae (Menaethiops), 57

delagoae (Palaemon), 75

delagoae (Palinurus gilchristi), 69

delagoae (Periclimenes), 75

delagoae (Pisidia), 66

delagoae (Porcellana), 66

delagoae (Thalamita), 59

demani (Polycheles), 69

dentatus (Glyphocrangon), 73

difficilis (Hapaloptyx), 66

Diogenes, 65

disjunctipes (Corycodus), 63

disjunctipes (Nasinatalis), 63

dissodontonotus (Alpheus), 73

dissothrix (Dromidia), 61

Dromidia, 61

Dromidiopsis, 61

Dynomene, 62

Ebalia, 62, 63

elegans (Eurynome), 57

engyops (Paguristes), 65

equidens (Macrobrachium), 75

Ethusa, 63

Eualus, 71, 72

eucheir (Xeinostoma), 64

Eudromia, 61

Eudromidia, 61

Eurynome, 57

Exhippolysmata, 72

extricatus (Diogenes), 65

fossula (Paracleistostoma), 58

gamianus (Paguristes), 65

gilchristi (Callianassa), 67

gilchristi (Leander), 74

gilchristi (Palinurus), 68

glomus (Ebalia), 63

gloriosus (Sergestes), 71

Glyphocrangon, 73

Haliporoides, 70

Hapaloptyx, 66

hendersoni (Anapagurus), 64

TYPE SPECIMENS OF DECAPODA (CRUSTACEA)

hendersoni (Eudromia), 61

hendersoni (Eudromidia), 61

Heteronucia, 63

Hexapus, 60

Hippolysmata, 72

Hippolyte, 72

Homola, 62

Hymenopenaeus, 70

Ilyograpsus, 59

inhacae (Thalamita), 59

Ischnopontonia, 75

jacqueti (Metacrangon), 76

Jasus, 68

jucundus (Betaeus), 74

kilburni (Parapagurus), 66

kraussi (Callianassa), 67

laevis (Petalomera), 62

Latreillopsis, 62

Leander, 74, 75

Lebbeus, 72

Leontocaris, 72

lepidota (Cryptodromiopsis), 61

liochele (Pylopagurus), 66

Lithadia, 62

longispina (Axius), 67

longispina (Metixonaxius), 67

lophos (Ischnopontonia), 75

lophos (Philarius), 75

luciae (Alpheus), 74

Lupocyclus, 59

Lybia, 60

Macrobrachium, 75

macrotrichus (Paguristes), 65

makrothrix (Cancellus), 65

margaritatus (Dairoides), 59

Maxillothrix, 62

mcleayi (Portumnus), 59

mcleayi (Xaiva), 59

megalocheir (Philocheras), 76

megalocheir (Pontophilus), 76

Menaethiops, 57

Metacrangon, 76

Metapenaeus, 70

Meticonaxius, 67

monoceros (Metapenaeus), 70

monodon (Penaeus), 69

mossambicus (Palinustus), 69

Munidopsis, 66, 67

Nasinatalis, 63

natalensis (Acetes), 71

natalensis (Callianassa), 68

natalensis (Menaethiops), 57

Neolithodes, 64

nitidus (Plesiopenaeus), 70

nonalter (Alpheus), 74

Notostomus, 71

Nursia, 63

Ogyrides, 73

Ommatocarcinus, 60

pacificus (Palaemon), 74, 75

Paguristes, 65

Palaemon, 74, 75

Palicus, 58

Palinurus, 68, 69

Palinustus, 69

paludicola (Ilyograpsus), 59

palliola (Hippolyte), 72

Paracleistostoma, 58

Paramola, 62

Parapagurus, 65

parkeri (Jasus), 68

parkeri (Projasus), 68

paulsoni (Leontocaris), 72

pax (Eualus), 72

pax (Spirontocaris), 72

Penaeopsis, 70

Penaeus, 69

Periclimenaeus, 75

Periclimenes, 75

peringueyi (Leander), 75

Petalomera, 62

Philarius, 75

Philocheras, 76

pilosimanus (Parapagurus), 65

pilumnoides (Dynomene), 62

Pisidia, 66

planiforma (Retropluma), 58

Platymaia, 57

Plesiopenaeus, 70

plumosa (Lybia), 60

Polycheles, 69

Pomatocheles, 64

Pontophilus, 76

Porcellana, 66

Portumnus, 59

postulans (Nursia), 63

prehensilis (Sergestes), 71

Processa, 73

Projasus, 68

pulcher (Ommatocarcinus), 60

Pylopagurus, 66

te)

80 ANNALS OF THE SOUTH AFRICAN MUSEUM

Retropluma, 58

rhizophorae ({lyograpsus), 59

Rhynchoplax, 57

rosaceus (Paguristes), 65

rostratus (Salmoneus), 74

saldanha (Spirontocaris), 72

saldanhae (Lebbeus), 72

saldanhae (Ogyrides), 73

Salmoneus, 74

Sclerecrangon, 76

Sergestes, 71

sexlobata (Palicus), 58

sinespina (Ethusa), 63

Solenocera, 70

spinosissimus (Achaeus), 56

spinulicauda (Penaeopsis), 70

spiralis (Thaumastoplax), 60

Spirontocaris, 71, 72

stebbingi (Hexapus), 60

stebbingi (Stylodactylus), 71

Stylodactylus, 71

Synalpheus, 74

sil

Thalamita, 59

Thaumastoplax, 60

triarthrus (Haliporoides), 70

triarthrus (Hymenopenaeus), 70

trifurcus (Cymonomus), 63

tuberculosa (Ebalia), 63

tugelae (Hippolysmata), 72

tugelae (Lupocyclus), 59

turbynei (Platymaia), 57

undulatus (Dehaanius), 57

Upogebia, 68

uropodialis (Periclimenaeus), 75

waltervadi (Alpheus), 74

Willemoesia, 69

Xaiva, 59

Xeinostoma, 64

Xenophthalmodes, 60

INSTRUCTIONS TO AUTHORS

Based on

CONFERENCE OF BIOLOGICAL EDITORS, COMMITTEE ON FORM AND STYLE. 1960.

Style manual for biological journals. Washington: American Institute of Biological Sciences.

MANUSCRIPT

To be typewritten, double spaced, with good margins arranged in the following order:

(1) Heading, consisting of informative but brief title, name(s) of author(s), address(es) of

author(s), number of illustrations (figures, enumerated maps and tables) in the article.

(2) Contents. (3) The main text, divided into principal divisions with major headings; sub-

headings to be used sparingly and enumeration of headings to be avoided. (4) Summary.

(5) Acknowledgements. (6) References, as below.

Figure captions and tables to be on separate sheets.

ILLUSTRATIONS

To be reducible to 12 cm x 18 cm (19 cm including caption). A metric scale to appear

with all photographs.

All illustrations to be termed figures (plates are not printed; half-tones will appear in their

proper place in the text), with arabic numbering; items of composite figures to be designated

by capital letters (A, B, C etc.).

REFERENCES

Harvard system (name and year) to be used: author’s name and year of publication given

in text; full references at the end of the article, arranged alphabetically by names, chronologi-

cally within each name, with suffixes a, b, etc. to the year for more than one paper by the same

author in that year.

For books give title in italics, edition, volume number, place of publication, publisher.

For journal articles give title of article, title of journal in italics (abbreviated according to the

World list of scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses,

volume number, part number (only if independently paged) in parentheses, pagination.

Examples (note capitalization and punctuation)

BuULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FiscHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris

88: 100-140.

FIsCcHER, P.-H., DUvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des

littorines. Archs Zool. exp. gén. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee

region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19605. Spawning behaviour, egg masses and larval development in Conus from the

Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51. -

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn: SCHULTZE, L.

Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und

zentralen Siid-Afrika. 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16:

269-270.

ZOOLOGICAL NOMENCLATURE

To be governed by the rulings of the latest International code of zoological nomenclature

issued by the International Trust for Zoological Nomenclature (particularly articles 22 and

51). The Harvard system of reference to be used in the synonymy lists, with the full

references incorporated in the list at the end of the article, and not given in contracted form

in the synonymy list.

Example

Scalaria coronata Lamarck, 1816: pl. 451, figs 5 a, b; Liste: 11. Turton, 1932: 80.

Brian Kensley

TYPE SPECIMENS OF DECAPODA (CRUSTACEA)

IN THE COLLECTIONS OF tee

SOUTH AFRICAN MUSEUM

“OF THE SOUTH AFRICAN

"MUSEUM

CAPE TOWN

TT Stitt

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 66 + Band

November 1974 November

Part 5 Deel

4 / BS

SEMPER <4 ES AFFERS

LIQUID NOVI AFRICH

ik CREAACEOUS STRAMIGRAPE Y

OF SOUTH-CENTRAL AFRICA

MICHAEL VR COOPER

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

Obtainable from the South African Museum, P.O. Box 61, Cape Town

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

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OUT OF PRINT/UIT DRUK

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Price of this part/Prys van hierdie deel

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1974

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Printed in South Africa by In Suid-Afrika gedruk deur

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

THE CRETACEOUS STRATIGRAPHY OF SOUTH-CENTRAL AFRICA

MICHAEL R. COOPER

South African Museum, Cape Town

(With 10 figures)

[Ms. accepted 15 October 1973]

CONTENTS

PAGE

Introduction : : ; : ; 81

Stratigraphy of the west coast , : : : : 81

Stratigraphy of the south coast : : : : 88

Stratigraphy of the east coast : ; 89

The Cretaceous history of Africa south of the Sahara 4 99

Summary . ; ! : : : : i 104

Acknowledgements : 3 : ; ‘ : . 104

References . : : : : : ‘ ; . 104

INTRODUCTION

Recent studies by Reyment (1969, 1971) have led him to suggest that

‘Cretaceous (Albian to Turonian) transgressions around the coast of Africa

display a correlative pattern that suggests they may be the results of a see-saw

motion’ (1971: 1063). Field-work and biostratigraphical studies in Angola,

the south-east Cape, Zululand and southern Mocambique do not support this

conclusion. A study of the Cretaceous deposits of southern and central Africa

suggests the observed sequence of transgressions and regressions can best be

explained by eustatic sea-level changes, together with local coastal warping,

rather than oscillatory tiltings of the African continent.

STRATIGRAPHY OF THE WEST COAST

Along the west coast of Africa south of the Sahara, i.e. south of latitude

20°N, deposits of Cretaceous age are known from many localities. The occur-

rence of these sequences is important in the study of the break-up of Gondwana-

land and the early history of the South Atlantic Ocean.

According to Furon (1963: 232), the first genuine marine horizon in Ghana

is ‘that of the Anwiafutu Limestone with Plicatula auressensis and Exogyra

olisiponensis’, considered to be of Upper Cenomanian—Turonian age, whilst the

earliest marine beds in the Ivory Coast and Upper Volta are assigned to the

Upper Cenomanian on the basis of Plicatula fourneli Coquand.

The Cretaceous biostratigraphy of Nigeria (Fig. 1) is well known following

the studies of Reyment (1954a, 19546, 1955, 1965) and Barber (1957). The earliest

Ann. S. Afr. Mus. 66 (5), 1974: 81-107, 10 figs.

82 ANNALS OF THE SOUTH AFRICAN MUSEUM

recorded marine beds contain Oxytropidoceras of low Middle Albian age, this

genus dating the maximum transgression of the Albian in Nigeria (Reyment

1955). The remainder of the Albian appears to be present and is in places highly

fossiliferous. Undoubted Cenomanian is known only from Calabar Province

(Reyment & Tait 1972), where deposits with Euhystricoceras, Turrilites, Forbesi-

ceras, Acompsoceras and Acanthoceras quadratum Crick occur on the coast.

Euhystricoceras is confined to Lower Cenomanian strata while the remaining

forms are typical of the Middle Cenomanian. The only Upper Cenomanian

Pe 2S NN

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————

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ING =

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) Lower Cretaceous Series

7 Calabar /

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Fig. 1. The Cretaceous outcrop in Nigeria. After Reyment 1955.

species so far recorded (Reyment & Tait 1972) is Metoicoceras aff. ornatum

(Kennedy 1971).

The Lower Turonian of Nigeria is very rich in ammonites (Reyment

1954a, 19546, 1955; Barber 1957), although Upper Turonian strata appear to

be lacking in diagnostic fossils. The Coniacian is well established by the presence

of the ammonites So/gerites, Forresteria, Peroniceras (Peroniceras), P. (Reginai-

tes), Tissotia, and Gauthiericeras, Santonian (?Lower) is known from Nigeria

THE CRETACEOUS STRATIGRAPHY OF SOUTH—CENTRAL AFRICA 83

BAIE DE

BIAFRA

Recent volcanics

Tertiary - Quaternary

~ Senonian

ES Turonian Cretaceous

— Unfossiliferous strata

Basement

Fig. 2. The distribution of Cretaceous strata in Cameroun. After Belmonte 1966.

and Zaire by Texanites. According to Reyment (1966: 171) “The Santonian was,

in the Nigerian basin, a time of regression’. This possibly explains the absence

of the rich Pseudoschloenbachia-fauna which characterizes the Upper Santonian

(Collignon 1968) of the east coast. The Campanian and Maastrichtian of Nigeria

have yielded Libycoceras, Sphenodiscus, Didymoceras and Baculites. ;

Belmonte (1966) has summarized the latest stratigraphical data concerning

the Cretaceous deposits of Cameroun (Fig. 2). Albian is known only from near

the Nigerian frontier where cross-bedded sandstones and intercalated shales

have yielded plant fragments. Marine Cenomanian has not yet been recorded.

In the Mungo Valley the Cretaceous ‘forment une alternance d’épisodes

marins transgressifs’ (Belmonte 1966: 11), and Diebold (1962) has recognized

three sedimentary cycles.

The first cycle is a transgressive sequence dated at upper Lower Turonian

by the ammonites Kamerunoceras, Neoptychites, Choffaticeras, Hoplitoides, etc.

The second cycle is a regressive phase dated at Upper Turonian by Romaniceras

84 ANNALS OF THE SOUTH AFRICAN MUSEUM

aff. uchauxiense (Collignon) and R. aff. deverioides (de Grossouvre). A trans-

gressive sequence with Tissotia, Barroisiceras, and Peroniceras, dated at Conia-

cian, constitutes the third cycle.

The Santonian is well developed in the Douala basin, where a marine

intercalation has yielded Texanites quattuornodosus (Lasswitz) and T. aff. soutoni

(Baily). Reyment (#7 Muller 1966: 140) has recorded the Upper Santonian

Pseudoschloenbachia from the Mungo River Formation.

In the Douala basin the Campanian is represented by a transgressive

sequence yielding the ammonites Eupachydiscus, Karapadites, Menabites, and

Parapuzosia. These forms suggest a Middle to Upper Campanian age. The

presence of Parapachydiscus stallauense Imkeller is taken to provide evidence of

the Maastrichtian, since this species is known from the Lower Maastrichtian

of Nigeria.

The earliest marine horizon in Gabon has yielded an ammonite, the Lower

Aptian Deshayesites consorbinoides Sinzow according to De Klasz & Gageonnet

(1965: 285), although Reyment & Tait (1972) consider it better referred to the

Upper Aptian genus Neodeshayesites. These beds are succeeded higher up by

strata with Douvilleiceras, of upper Lower Albian age. Overlying beds have

yielded the ammonites Oxytropidoceras, Mortoniceras, and Elobiceras, and

probably span much of the remainder of the Albian. The Cap Lopez Formation

has been assigned a Cenomanian age on the basis of foraminifers. Turonian

strata with Wrightoceras, Bauchioceras, etc., is well known, although the presence

of the Upper Turonian is still in doubt (Reyment & Tait 1972). Coniacian strata

with Peroniceras, Gauthiericeras, and Barroisiceras discordantly overlie Turonian

rocks (Reyment & Tait 1972). Texanites provides evidence of the Santonian,

while Upper Campanian is known from the vicinity of Libreville by strata with

Hoplitoplacenticeras (Reyment 1966: 170).

Records of Aptian from the west coast are based largely on microfossil

evidence, although the bivalve Panis cabrai (Dartevelle & Freneix) has been

used to suggest an Aptian age for strata in Zaire and Angola. Presumably on

the basis of microfossils, Hoppener (1958: 75) recognized a ‘transgression

aptienne inférieure ou ante aptienne’, and a ‘transgression aptienne supérieur’

from Angola. No ammonites have yet been collected to support this contention.

In Angola (Fig. 3) the earliest undoubted marine beds occur at Dombe

Grande, to the south-east of Benguela, and have yielded the ammonites

Douvilleiceras mammillatum inaequinodum (Quenstedt) and D. orbignyi Hyatt

(Howarth 1965). The beds immediately above are rich in Oxytropidoceras

and Manuaniceras. These upper Lower Albian—lower Middle Albian beds

are characterized by a fauna rich in numbers but poor in species. Unfossiliferous

sediments overlying these beds would seem to span the remainder of the Middle

Albian and are succeeded by Upper Albian strata with Mortoniceras. No

ammonites have been recorded from the Middle Albian strata of Angola above

beds with Oxytropidoceras. Strata presumably of a similar age, at Hanha, the

supposed fossil locality of Haas (1942a), are also unfossiliferous. Near Hanha,

THE CRETACEOUS STRATIGRAPHY OF SOUTH—CENTRAL AFRICA

e@Quibala

™Gabela

4 Redondo

: Tertiary System

(TM) Upper Cretaceous Series

Lower Cretaceous Series

Salinas

w Sa da Bandeira

Fig. 3. Locality map showing the distribution of Cretaceous

strata in Angola. In part after Mouta & O’Donnell 1933.

86 ANNALS OF THE SOUTH AFRICAN MUSEUM

however, low Upper Albian strata are rich in Neokentroceras, Dipoloceras,

Mortoniceras, etc. The remainder of the Upper Albian in Angola is frequently

extremely rich in Mortoniceras, Elobiceras, Prohysteroceras, Anisoceras and

Stoliczkaia. Reyment & Tait (1972), following Howarth (1965), suggested that

Stoliczkaia was of Lower Cenomanian age in Angola, as did Kennedy (1971).

This is wrong. This misconception has arisen from Douvillé’s (1931) record of

Stoliczkaia dispar var. attenuata Douvillé from the uppermost Cenomanian of

Salinas (Cooper 1972). Further topotype material suggests that this specimen

belongs not to this Upper Albian—Lower Cenomanian genus, but should rather

be considered the homeomorphous development of some other acanthocerate

stock, possibly the early Turonian Benueites (W. J. Kennedy, pers. comm.).

The uppermost Albian of Angola, as exposed at Egito, is characterized by

Anisoceras perarmatum Pictet & Campiche, Stoliczkaia spp., and mortoniceratids.

At Egito the lowest fossiliferous horizons, resting upon crystalline base-

ment, have yielded Mortoniceras, suggesting the beginning of the Upper Albian

to have been a period of transgression. As noted by Reyment (1969), uppermost

Albian deposits occur closest to the coast, e.g. Egito, Catumbela, suggesting a

gradual regression following the early Upper Albian transgression.

Deposits at Salinas indicate the uppermost Cenomanian to have been a

period of transgression, yielding the ammonites Calycoceras naviculare (Mantell),

Austiniceras dibleyi Spath, Pseudocalycoceras angolaense (Spath), Sciponoceras

gracile (Shumard), Kanabiceras septemseriatum (Cragin) and Metoicoceras cf.

whitei Hyatt. The overlying beds comprise a condensed sequence spanning

Turonian and earliest Coniacian times (Cooper 1972), and contain the following

ammonite genera: Mammites, Subprionocyclus, Damesites, ‘Prionocyclus’,

Hauericeras, Baculites, Kossmaticeras, ?Subtissotia, Proplacenticeras, Gaudry-

ceras, Hypophylloceras, Anagaudryceras, and Mesopuzosia. The rich and varied

Turonian fauna of Nigeria is not known from Angola. These beds are dis-

conformably overlain by subaerially extruded alkaline volcanics of Upper

Coniacian age (Cooper 1972), in turn succeeded by early Santonian marine

beds with Protexanites, Texanites, Hauericeras and Damesites.

The stratigraphy of the Cuanza basin is at present poorly known. Hoppener

(1958) assigned an Upper Aptian age to oolitic limestones near the base of the

succession with Acteonella aff. fusiformis, Cerithium cf. albense, and Perna sp.

while a rich Upper Albian fauna is known from Cabo Ledo.

At the beginning of Cenomanian times the sea transgressed in Angola,

with Sharpeiceras laticlavum (Sharpe) occurring at Cabiri, while in the south

of the Cuanza basin, at Novo Redondo, the transgression extended about 20 km

inland, the lowest beds yielding the Lower Cenomanian Mantelliceras cf. saxbii

(Sharpe) (Cooper 19736). There was probably a gradual regression until Middle

Cenomanian times, since beds with Turrilites acutus Passy, Euomphaloceras

cunningtoni (Sharpe), Forbesiceras obtectum (Sharpe), etc., occur at the coast.

No higher beds are known from Novo Redondo. Haas (19425) recorded a

Romaniceras sp. from north of Cabiri, which suggests a Turonian age. Hoppener

THE CRETACEOUS STRATIGRAPHY OF SOUTH—CENTRAL AFRICA 87

(1958) also assigned a Turonian age to Coilopoceras cacobaensis sp. nov. (a

nomen nudum since it was neither figured nor described). Downfaulted strata at

Cabo Ledo have yielded an uppermost Turonian fauna with Prionocyclus and

Subprionocyclus (Basse 1963; Matsumoto 1965: 49; Howarth 1968: 225). Of

probable Coniacian age is Proplacenticeras reineckei (Haughton), while Hoppener

(1958) assigned ‘Placenticeras’ fritschi Hyatt and a Hemitissotia sp. to the

Coniacian. The occurrence of Texanites angolanus Haas and T. quinquenodosus

(Redtenbacher) to the north of Cabiri provides evidence of the Santonian. A

rich, predominantly heteromorph fauna from Barra do Dande suggests an

Upper Campanian—Maastrichtian age (Howarth 1965; Antunes & Sornay

1969). Amongst the genera recorded are Didymoceras, Nostoceras, Manam-

bolites, Solenoceras, Axonoceras, Polyptychoceras, and Sphenodiscus. Antunes

& Sornay (1969) consider these beds, the Barra do Dande Formation, to rest (?)

disconformably upon the Pambala Formation which they tentatively dated at

Upper Santonian—Lower Campanian on the basis of inoceramids assigned to

the bantu—balticus group. No ammonites have been collected to support this

evidence. The apparent absence of Hoplitoplacenticeras at Barra do Dande, a

common ammonite in the Upper Campanian fauna of the Egito outlier, suggests

that the transgression may have been slightly diachronous, occurring earlier

in the south than in the north. From far inland, at Carimba, Haughton (1925)

recorded Nostoceras, Didymoceras, Menuites, Libycoceras, etc., for which an

Upper Campanian—Lower Maastrichtian age appears the most precise dating.

A similar age seems the most likely for Haas’s (1943) fauna from the Maria

Theresa area. A major transgression undoubtedly took place in Angola during

Upper Campanian—Maastrichtian times. The base of the transgression is not

known from the Cuanza basin, but at Egito is dated at Upper Campanian

(vari Zone) probably reaching its maximum extent somewhat later. At San

Nicolau, a horizon rich in sharks’ teeth was assigned to the Maastrichtian

(Dartevelle 1942). Cooper (1972), on sedimentalogical grounds, doubted this

age determination. In retrospect, however, the occurrence of phosphatic

pellets associated with the sharks’ teeth suggests the possibility of a non-

sequence. Should such a break exist, then these beds possibly represent the

base of the late Campanian transgression.

From the Cavaco valley near Benguela, Spath (1951: 124) recorded -a

‘presumed Campanian’ fauna with Jnoceramus langi Choffat and ammonites

tentatively identified as Hauericeras gardeni (Baily), Damesites, and Neancy-

loceras from clays resting unconformably upon Upper Albian strata.

Haughton (1930) described a small inlier of marine Cretaceous from

Bogenfels in South West Africa, from which he recorded the ammonite

Proplacenticeras merenskyi (Haughton) in association with the oyster Exogyra

cf. columba (Lamarck) (= Rhynchostreon suborbiculatum (Lamarck)), these

beds being characterized by the abundance of the latter. At Salinas in southern

Angola, the uppermost Cenomanian is characterized by the abundance of

Rhynchostreon cf. suborbiculatum (Cooper 1972), which is absent from the

88 ANNALS OF THE SOUTH AFRICAN MUSEUM

overlying beds, nor has it been recorded from the Middle Cenomanian of Novo

Redondo. An uppermost Cenomanian age would appear the most likely for the

Bogenfels outlier. H. C. Klinger (pers. comm.), who has personally visited the

Wanderfeld IV locality of Haughton, considers the Cretaceous fauna to, in all

probability, represent remanié, although in this respect the apparent abundance

of the diagnostic Cenomanian oyster R. suborbiculatum is critical. Haughton

(1930) believed these ‘Exogyra beds’ to be unconformably overlain by the

Senonian with Protocardia hillana (Sowerby), Turritella (Zaria) bonei Baily and

T. (Haustator) meadi Baily. Together these forms suggest a possible Santonian

age.

STRATIGRAPHY OF THE SOUTH COAST

Along the south coast of South Africa (Fig. 4), marine strata at Knysna,

long correlated with the Uitenhage Group, have been assigned ‘A high

Mid-Kimmeridgian age’ (Klinger et a/. 1972) on the basis of the ammonite

242

KARROO BASIN

LTS

* Grahamstown

Cape Town

2 SPlettenbe =e Port Elizabeth

Knysna

Late Mesozoic deposits

26°

Fig. 4. The distribution of late-Mesozoic strata in the southern Cape. After Geol. Surv. S. Afr.

1970.

Hybonoticeras aff. hildebrandti (Beyrich), while Dingle & Klinger (1972) recorded

an ostracode assemblage with strong Callovian affinities. These are the earliest

Mesozoic marine sediments to be recorded from South Africa. At Robberg,

near Plettenberg Bay, casts of Megatrigonia conocardiiformis (Krauss) occur in

sandstones of the Enon Formation of the Uitenhage Group, and are probably

of Upper Jurassic age, since this species is also known from the Upper Kim-

meridgian—Tithonian of Mocgambique (Da Silva 1966).

To the north of Port Elizabeth, sediments assigned to the Uitenhage

Group (Cooper 1973a) crop out over wide areas, although it is only the upper

portion, the Sundays River Formation, that has yielded ammonites. The

cephalopod fauna is characterized by the abundance of species of Olcostephanus

s.s., together with Distoloceras, Bochianites, Eodesmoceras, Partschiceras,

?Leopoldia, and Belemnopsis. The absence of finely-ribbed forms of Olcoste-

THE CRETACEOUS STRATIGRAPHY OF SOUTH—CENTRAL AFRICA 89

phanus, e.g. O. filosus (Baumberger), which characterize the Lower Hauterivian

of France (Busnardo & Cotillon 1964), and of the subgenus O. (Jeannoticeras)

favours an uppermost Valanginian age for the Uitenhage fauna.

STRATIGRAPHY OF THE EAST COAST

To the south of Port St. Johns, at the Umgazana River mouth (Fig. 5),

Cretaceous conglomerates have yielded belemnites, the ammonite Bochianites

(Du Toit (1954) records the Upper Cretaceous Baculites, but this is a mis-

identification), and the bivalve Steinmanella cf. holubi (Kitchin). Together these

forms favour a correlation with the Uitenhage Group, and an uppermost

Valanginian age.

A short distance to the north of Port St. Johns are outcrops of unfossili-

ferous strata assigned to the Emboyti Formation. Although no marine fossils

have yet been recorded, the close lithological similarity with the Umgazana

Formation led Du Toit (1954) to suggest a correlation.

The age of the Pondoland Umzamba Formation has long been a source of

contention, but Spath (1953) was led to conclude that it represented a con-

densed sequence. The occurrence of Pseudoschloenbachia, Lewesiceras, Texanites,

Hauericeras, Barroisiceras, Eulophoceras, Forresteria, Pseudophyllites, Koss-

maticeras (Natalites), Pseudoxybeloceras, etc., suggests that the condensation

may span Upper Coniacian to Lower Campanian times, with the majority of

forms Santonian. Gevers & Little (1946) recorded an isolated outlier of Creta-

ceous, the Itongazi Formation, from between the Umkandandhlovu and Iton-

gazi Rivers in southern Natal. Although no ammonites have yet been recorded,

the majority of bivalves and gastropods are also common to the Umzamba

Formation, and leave little doubt as to their contemporaneity.

Two quarries at Need’s Camp, near East London, have yielded Upper

Senonian faunas although no diagnostic forms have been reported. On fora-

miniferal evidence, McGowran & Moore (1971) have assigned the Lower

Quarry an Upper Campanian—Maastrichtian age. The commonest macrofossils

are well-preserved sharks’ teeth, assigned to Jsurus and Carcharias, the coral

Caryophyllia, and bryozoans, together with the echinoid Coptosoma, the

brachiopod Lacazella, and the bivalves Exogyra, Neithea(?) and Inoceramus.

A similar age has been assigned to the Upper Quarry (King 1972), long con-

sidered of Eocene age, also on foraminiferal evidence. Macrofossils from this

quarry include teeth of the sharks Oxyrhina, Odontaspis and Carcharodon, the

gastropod Pyropsis, and the bivalves Plicatula, Panopea and Lima. No ammo-

nites have been recorded from either quarry. Lock (1973), however, considers

the Upper Quarry to be Miocene in age, with incorporated Cretaceous and

Eocene remanié.

Kennedy et al. (1973) have recently described an Upper Santonian—

Campanian borehole fauna from Durban, including the following species:

Anagaudryceras subsacya (Marshall), Texanites (Plesiotexanites) stangeri (Baily),

ANNALS OF THE SOUTH AFRICAN MUSEUM

ka Umkwelane Hill

Port Shepstone

Itongazi

Umzamba

Emboyti

ie)

Port St Johns

Umgazana

Need's Camp

East London

Fig. 5. Locality map showing the Cretaceous expo-

sures along the east coast of South Africa; the Zulu-

land outcrop is stippled. After Geol. Surv. S. Afr.

1970.

THE CRETACEOUS STRATIGRAPHY OF SOUTH—CENTRAL AFRICA 9]

Submortoniceras woodsi (Spath), Hoplitoplacenticeras plasticum Paulcke, Haueri-

ceras cf. gardeni (Baily), Kossmaticeras (Natalites) cf. acuticostatus (Spath), etc.

These authors considered this fauna to generally resemble that of the Umzamba

Formation.

In Zululand the earliest marine outcrops are of Barremian age (W. J.

Kennedy and H. C. Klinger, pers. comm.), with the ammonites Heteroceras,

Colchidites, Phylloceras serum Oppel, Eulytoceras phestum (Matheron), etc. The

occurrence of Tropaeum, Australiceras, Cheloniceras, Acanthohoplites, and

Diadochoceras (Haughton in Rennie 1936) indicates most of the Aptian to be

present. Beds containing Diadochoceras are overstepped by strata with

Douvilleiceras mammillatum (Schlotheim) (Kennedy & Klinger 1971); thereafter

deposition appears to have been continuous until the low Upper Cenomanian

with Calycoceras gr. naviculare (Mantell), although after deposition of beds

with Acanthoceras quadratum Crick, A. spp., Turrilites acutus Passy, Calycoceras

spp., Forbesiceras largilliertianum (dOrbigny), etc., the sequence is poorly

fossiliferous. These beds are unconformably overlain by Lower Coniacian strata

(Kennedy & Klinger 1971) with Proplacenticeras subkaffrarium (Spath), P.

umkwelanense (Etheridge), P. kaffrarium (Etheridge), Kossmaticeras theobal-

dianum (Stoliczka), Bostrychoceras indicum (Forbes), etc., followed by con-

tinuous deposition until the Lower Maastrichtian with Eubaculites, Pachydiscus

(Neodesmoceras), and Saghalinites. The uppermost Cenomanian and Turonian

was a period of non-deposition, the hiatus being represented by a Lithophaga-

bored hardground.

The earliest recorded marine horizon in Mocgambique, at Nacala (Fig. 6),

has been dated at Upper Kimmeridgian-Tithonian (Da Silva 1966) on the

basis of the bivalves Trigonia (Indotrigonia) smeei auct. and Astarte krenkeli

Dietrich. A cephalopod from these beds has not yet been described.

Slightly further to the north, at Fernao Veloso, strata with Olcostephanus

schenki (Oppel) (=O. baini (Sharpe)) and Haploceras (Neolissoceras) cf. grasia-

num (d’Orbigny) (Wray 1915; Spath 1930) provide evidence of the uppermost

Valanginian. Strata of a similar age also crop out at Mahiba Hill, west of Port

Amélia, from where Newton (1924) recorded fragments of Lytoceras, together

with the belemnite Duvalia. From this same locality, Spath (1930: 134) also

records a ‘portion of the periphery of a Neocomitid (Lyticoceras of the type of

L. regalis (Bean) or Neocomites neocomiensis (d’Orbigny) as figured by Sayn),

and the impression of a fragment of the Uitenhage Bochianites africanus are

decisive and unmistakable’. Neumayr (1885) recorded the Neocomian

Phylloceras semistriatum (d’Orbigny) from the Mocgambique region, an identifi-

cation subsequently confirmed by Zwierzycki (1913).

Aptian and Albian strata are best known from southern Mogambique

(Fig. 7), although exposures are poor and much of the area is capped by younger

Neogene deposits.

The best exposures are at Catuane, the so-called Maputoland locality of

Spath (1925), where the earliest beds have been assigned to the Upper Aptian

ANNALS OF THE SOUTH AFRICAN MUSEUM

Porto Amélia

Fernao Veloso

Sheringoma’ Plateau

Inhambane

|: INComaning

Fig. 6. The distribution of late-Mesozoic strata in Mocgambique.

De Freitas 1957.

Nacala:

Conducia e

Largely after

THE CRETACEOUS STRATIGRAPHY OF SOUTH—CENTRAL AFRICA 93

hi)

BAIA DE

LOURENGO MARQUES

Mahube R.

) ema

3 Powell's Camp

25] Senonian

re Aptian — M.Cenomanian

SWAZILAND

RIO MAPUTO

Lubemba

Fig. 7. The Cretaceous exposures of southern Mocgambique. After Servigos de Geologia

e Minas da Provincia de Mocambique, 1969.

(Da Silva 1962; Wachendorf 1968). On the farm Lubemba, from where both

Da Silva’s and Wachendorf’s collections came, recent collecting has yielded a

rich Lower Aptian (deshayesi-bowerbanki Zones) fauna dominated by the

abundance of Cheloniceras (Cheloniceras), together with the genera 7ropaeum,

Ancyloceras, ‘Neosilesites’, ‘Valdedorsella’, Phylloceras, Toxoceratoides, Pseudo-

saynella, etc. Wachendorf (1968) records Parahoplites campichei Pictet &

Renevier and P. melchioris Anthula from the Maputo River which provide

evidence of the Upper Aptian at this locality as well. The occurrence of

Phylloceras at Lubemba, which Da Silva (1962) thought to resemble P. velledae

(Michelin), led him to suggest the presence of Albian at this locality. This is

incorrect. Recent collecting has shown the Phylloceras to be closely associated

with Cheloniceras (Cheloniceras), and thus of Lower Aptian age. No Lower

94 ANNALS OF THE SOUTH AFRICAN MUSEUM

Albian fossils have yet been recorded from Catuane. The presence of the Middle

Albian is indicated by the occurrence of Oxytropidoceras, while Spath (1925)

has recorded a rich low Upper Albian fauna characterized by the heteromorphs

Anisoceras, Labeceras, and Myloceras, together with Mortoniceras, Elobiceras

and Hysteroceras. The uppermost Albian (dispar Zone) has not been recorded,

but its presence is to be suspected. Spath (1925) described Sharpeiceras florencae

Spath from Maputoland, and thus presumably Catuane. This genus is confined

to the Lower Cenomanian. The occurrence of Trigonarca cf. ligeriensis

(dOrbigny), Pterotrigonia (Scabrotrigonia) shepstonei (Griesbach), and

Amphidonte columba (Lamarck) (= Rhynchostreon suborbiculatum (Lamarck))

from 3 km to the east of Catuane, led Soares & Da Silva (1970) to suggest

the presence of the Cenomanian. This is confirmed by the recent collection

of a worn fragment of Calycoceras from this area. These beds are uncon-

formably overlain by the Mandajene Formation. Amongst 52 species of bivalves

and gastropods recorded from this formation (Soares & Da Silva 1970), 31 are

common to the Umzamba Formation of Pondoland. Soares & Da Silva (1970: 2)

considered the Mandajene Formation to be ‘of Santonian—Campanian age, or

even Maestrichtian towards Santana de Tinonganine’. Unfortunately, these

beds are poor in ammonites. Internal moulds of baculitids were compared with

Baculites capensis Woods (Soares & Da Silva 1970), while Spath (1925) recorded

Texanites aff. soutoni (Baily) from Maputoland, and presumably, therefore,

from this region. The dating of the base of the Senonian transgression in Zulu-

land, of which succession the Catuane deposits are merely a continuation, at

Lower Coniacian (Kennedy & Klinger 1971) suggests a similar age for the base

of the Mandajene Formation.

Cretaceous strata are again exposed to the west of Porto Henrique, at the

renowned Chalala locality (Haughton & Boshoff 1956; Wachendorf 1968),

where the presence of Cheloniceras (Cheloniceras), Toxoceratoides, Aconeceras,

Acrioceras, and Phylloceras provide evidence of the upper Lower Aptian

(deshayesi-bowerbanki Zones). Lacey (1961) recorded the presence of Bostrycho-

ceras and Calycoceras at Chalala, to which he assigned a Cenomanian or Lower

Turonian age. The heteromorph fragment was assigned to Bostrychoceras on

the basis of its rounded whorls and non-tuberculate ribs, while ‘The smaller

fragment . . . has tubercles and appears to be an indeterminate species of

Calycoceras which has an age span of Cenomanian to Lower Turonian. Very

similar forms such as the Aptian Cheloniceras or Roloboceras would appear to

be excluded by the presence of Bostrychoceras, since forms with round whorls

and non-tuberculate ribs do not occur before the Cenomanian’ (p. 9). These

are almost certainly misidentifications since a recent visit to Chalala revealed

only friable, unfossiliferous Neogene (?) deposits resting disconformably, with

a basal small-pebble conglomerate, upon Lower Aptian strata. Further to the

north-east, however, Upper Albian beds with Mortoniceras, Hysteroceras and

Labeceras are exposed and are overlapped by Middle Cenomanian strata with

Acanthoceras.

THE CRETACEOUS STRATIGRAPHY OF SOUTH—CENTRAL AFRICA 95

To the south-east of Porto Henrique, rocks of Cretaceous age are exposed

on the Catembe River. Presumably this is the Powell’s Camp locality of Spath

(1921), from where he described Aconeceras nisoides (Sarasin), Cheloniceras

gottschei (Kilian) and C. delagoense (Krenkel), considered to be of Upper

Aptian age. There is little doubt that these beds should be correlated with the

other similar deposits of southern Mocgambique, and be considered of upper

Lower Aptian age.

Lower Aptian strata are again exposed along the Mahube River, where

the genera Aconeceras, Toxoceratoides, and Cheloniceras were collected. These

beds are overlain by the Cenomanian (Soares & Da Silva 1970), characterized

by the abundance of Turrilites costatus Lamarck, virtually to the exclusion of

all else. Soares & Da Silva (1970) have also recorded Acanthoceras from this

locality. It seems likely that these beds can be correlated with the ‘Tuwrrilites

costatus faunal assemblage’ of Kennedy (1971), of low Middle Cenomanian age.

Kilian (1902) and Krenkel (1910) both described Aptian fossils collected

from ‘Delagoa Bay’. No Cretaceous is exposed around Delagoa Bay, and this

locality is obviously the regional name applied by the early travellers.

The only other early Cretaceous deposit known from Mogambique is the

rich Upper Albian—Cenomanian locality of Conducia, in the north, from

where Choffat (1903) recorded Anagaudryceras sacya (=A. choffati Shimizu),

Mariella bergeri (=M. conduciensis Breistroffer), Phylloceras semistriatum

@Orbigny, ?Baculites sp., ?Hamites sp., Desmoceras latidorsatum (Michelin),

*Desmoceras beudanti’ var. petersoni Choffat, Sharpeiceras laticlavium mozam-

biquense Choftat, Calycoceras marquescostae (Choffat), and Mortoniceras cf.

candollei (Pictet).

Cox (1925) described an entirely new gastropod fauna from Incomanini,

to which he assigned an Upper Maastrichtian age, a determination supported

by a study of the bivalves (Rennie 1935). The Maastrichtian is also present along

the Buzi River and at Sheringoma, from where Crick (1924) has recorded

Eubaculites cf. vagina (Forbes) and the nautiloids Hercoglossa mazambensis

Crick and H. sheringomensis Crick.

The late Mesozoic stratigraphy of Tanzania (Fig. 8) is not well known.

‘Nowhere is there good dating below the Callovian, but a considerable part of

the Ruvu Beds (q.v.) is apparently Bajocian/Bathonian in age’ (Quennell er al.

1956: 156). To the east of the Ngarama Plateau sediments assigned to the

Mandawa—Mahokondo ‘Series’ have yielded Upper Oxfordian (plicatilis-Zone)

ammonites of the genera Perisphinctes (Arisphinctes), P. (Dichotomosphinctes),

and Euaspidoceras. From the same locality the genera Ptychophylloceras,

Hecticoceras (Sublunuloceras), Sindeites, Indosphinctes, Choffatia, Grossouvria,

Kinkeliniceras, etc., provide evidence of the Middle and Upper Callovian. The

overlying ‘Septarian Marls’ have been dated at Lower Kimmeridgian (mutabilis—

pseudomutabilis Zones) by the presence of Streblites, Physodoceras, Idoceras,

Taramelliceras, Virgatites, Perisphinctes (Lithacoceras), P. (Pachysphinctes),

Simoceras, etc. (Quennell et al. 1956).

ANNALS OF THE SOUTH AFRICAN MUSEUM

Dar-es-salaam

Kilwa Kivinje

Mandawa - Mahokondo

anticline

Ngarama

Plateau

Tendaguru

Fig. 8. Distribution of Jurassic (black) and Cretaceous (stippled)

strata in Tanzania. After Quennell et a/. 1956.

THE CRETACEOUS STRATIGRAPHY OF SOUTH—CENTRAL AFRICA 97

From Tendaguru, in the Mbemkuru Valley of southern Tanzania a Lower

Tithonian age has been favoured (Quennell et a/. 1956) for the ‘Trigonia smeei

Beds’. The ammonites include Haploceras, Hildoglochiceras, Subdichotomoceras,

etc. Above beds with Trigonia Undotrigonia) smeei (J. Sowerby) occur the non-

marine ‘Upper Saurian Beds’, which are also included in the Jurassic (Quennell

et al. 1956).

There is an apparent unconformity separating the succeeding ‘Trigonia

schwarzi Beds’, which have yielded the ammonites O/costephanus (Olcostephanus),

O. (Jeannoticeras), Phylloceras serum Oppel, Lytoceras hennigi Zwierzycki (to

which species Zwierzycki (1914) assigned the Mocgambique fragments),

Bochianites, etc. To the north-east of the Mandawa—Mahokondo anticline,

Neocomian strata are anticlinally-folded, and unconformably overlain by

Aptian strata (Quennell et a/. 1956). The presence of Heteroceras, Ancyloceras,

Procheloniceras, Cheloniceras (Cheloniceras) rauffi (Zwierzycki), and Diado-

choceras nodosocostatum (d’Orbigny) suggests at least part of the Barremian

as well as most of the Aptian to be present, while the occurrence of Puzosia

paronae Kilian was taken by Zwierzycki (1914) to indicate the Albian. Hennig

(1937) recorded Hysteroceras varicosum (J. Sowerby), ‘Turrilites’ aff. bergeri

Brongniart, Phylloceras broilii Krenkel, Puzosia kitchini (Krenkel) (= Tetra-

gonites kitchini) and P. africana Krenkel non Kilian (= Parasilesites austro-

africanus (Krenkel)) from the Namazatu area, to which he assigned on Albian

age. According to Murphy (1967: 36), Tetragonites kitchini ‘is dated as upper

Albian’. Hennig (1937) considered the Cenomanian and Turonian to be

absent from Tanzania, although Quennell ef a/. (1956) report Cenomanian

strata, presumably based on microfaunal evidence. Furon (1963) considered

the Cenomanian to be represented by beds with Rhynchostreon suborbiculatum

(Lamarck). It seems likely, however, that this species is of an earlier age along

the east coast than it is along the west coast. If, as suggested by Reyment & Tait

(1972), Africa and Brazil were connected until the early Turonian, then migra-

tion of R. suborbiculatum during the Cenomanian must have occurred via the

eastern shoreline. This species has not yet been recorded from Zululand and

does not appear to have migrated this far south by the beginning of the Upper

Cenomanian. It is, however, known from the pre-uppermost Cenomanian

strata of Catuane (Soares & Da Silva 1970). This suggests that its age in East

Africa is either Lower or Middle Cenomanian.

Bate & Bayliss (1973) have recently described the stratigraphy of the Cre-

taceous succession in the Wami River area, northern Tanzania. The sequence

is considered to extend from the Aptian to the Turonian, inclusive. Age deter-

minations are based solely on microfossils, and no macrofossils are reported.

The foraminifers Rotalipora greenhornensis (Morrow) and R. cushmani

(Morrow), together with abundant Praeglobotruncana delrioensis (Plummer)

were considered diagnostic of the Upper Cenomanian. Middle and Upper

Turonian strata are recognized on the basis of the foraminifers G/obotruncana

helvetica Bolli, G. linneiana (d’Orbigny), Hedbergella delrioensis (Carsey),

98 ANNALS OF THE SOUTH AFRICAN MUSEUM

Praeglobotruncana stephani (Gandolfi) and Clavihedbergella simplex (Morrow),

together with the ostracodes Cytherelloidea turonica Bate, Cytheruna moorei

Bate, C. /unzangaziensis Bate, Curfsina turonica Bate, Cythereis luzangaziensis

Bate, Cytherella afroturonica Bate, Akrogmocythere wamiensis Bate, Paracypris

wamiensis Bate and Sphaeroleberis africana Bate.

It would be most unusual were this succession complete, in view of the

fact that breaks in sedimentation are known to exist both to the north and to

the south. Consequently, ammonites are needed to corroborate the microfaunal

evidence.

The Senonian in Tanzania is known mainly on the basis of microfossils,

although Spath (in Quennell ef al. 1956) assigned an ammonite to the genus

Pachydiscus s.|.

Tavani (1949) considered all horizons from Barremian to Turonian, inclu-

sive, to be present in the Cretaceous deposits of Somalia. No diagnostic Barre-

mian ammonites have yet been recorded, unless Pseudothurmannia incertus

Tavani is correctly identified, and the presence of this stage is based largely on

bivalves and gastropods. Much of the Aptian is present, a rich Cheloniceras

fauna recording the presence of upper Lower Aptian (deshayesi-bowerbanki

Zones) strata. The Upper Aptian is represented by beds with Epicheloniceras

subnodosocostatum (Sinzow), Parahoplites weissi Neumayr & Uhlig, P. rudis

Tavani, Acanthohoplites cf. aschiltaensis (Anthula), A. mustahilensis Tavani, etc.

Tavani’s (1949) record of Hypacanthoplites milletianus (d’Orbigny) suggests the

possibility of the presence of uppermost Aptian—lowest Albian (jacobi-tarde-

furcata Zones) strata, beds which appear to be absent from the other strati-

graphic sequences of the east coast. This identification should be regarded as

tentative, since as pointed out by Casey (1965: 435) ‘although it (Hypacanthop-

lites milletianus) is one of the most widely quoted of Albian ammonites, it has

been generally wrongly identified’.

Albian strata are known from Bugda Acable where a rich Douvilleiceras

fauna yielding D. mammillatum (Schlotheim), D. monile Tavani (non J. Sowerby),

D. spinosum Tavani, D. benonae Besairie, D. variabile Tavani, and D. cheloni-

ceratiforme Tavani indicates the upper Lower Albian (mammillatum Zone).

Middle Albian strata have yielded Brancoceras zrissense Pervinquiére and B.

senequieri (d’Orbigny), while Owen (1971: 134) considers the occurrence of the

endemic lyelliceratid Somalites vertebralis Tavani together with Brancoceras

(although Tavani (1949) records Somalites in association with Upper Aptian

forms) ‘may well indicate the equivalent of the /ye//i Subzone (Middle Albian)’.

The occurrence of a Hysteroceras sp. and Labeceras crassum Spath provides

evidence for the presence of low Upper Albian sediments, although mortoni-

ceratids are still unrecorded.

Tavani (1949: 71) considers the Ferfer Gypseous ‘Series’ to be of Ceno-

manian age, as also the base of the Belet Uen ‘Series’, noting ‘qui in Somalia

invece il Cenomaniano presenta una estrema scarsita di Ammoniti, rappresentati

soltanto da alcuni esemplari appartenenti ai generi Heterotissotia and Placen-

THE CRETACEOUS STRATIGRAPHY OF SOUTH—CENTRAL AFRICA 99

ticeras’. Heterotissotia figarii Greco, known from the Coniacian of Egypt,

H. complanata Yavani, and Proplacenticeras tumulicum (Blandford) are not of

Cenomanian age, but are indicative of the Coniacian. Thus, as in Zululand,

transgressive Coniacian deposits rest disconformably upon Albian and possibly

early Cenomanian strata. According to Furon (1963: 304) the Senonian is

known from Somalia by the bivalves Venericardia beaumonti (d’Archiac) and

‘Exogyra overwegi von Buch.

THE CRETACEOUS HISTORY OF AFRICA SOUTH OF THE SAHARA

It is now possible to recognize the following Cretaceous history for southern

and central Africa. The Africa—South America rift was almost certainly in

existence as a rift valley during the late Jurassic—early Cretaceous (Reyment &

Tait 1972), with the invading sea reaching Gabon prior to the ?Upper Aptian.

Along the east coast rifting began in the north and reached Knysna by at least

Upper Jurassic times. With regard to the Jurassic deposits, Klinger ef al.

(1972: 658) noted that ‘Occurrences of material comparable to our specimen

(H. aff. hildebrandti) in Kutch and Kenya are thought to be Middle Kimmerid-

gian (Spath 1927, 1930a; Arkell 1956). Hybonoticeras also occurs in the Middle

Kimmeridgian of Somalia and Ethiopia (Arkell 1956) whilst in Madagascar,

where the genus occurs in both Kimmeridgian and Tithonian, the closest

comparisons are with forms from Collignon’s (1959) Middle Kimmeridgian.’

The widespread occurrence of strata of this age along the east coast would seem

to date a transgressive episode.

At the end of the Valanginian another marine transgression resulted in the

deposits of the Sundays River Formation of the Uitenhage Group (S.E. Cape),

the Umgazana and Emboyti Formations of Pondoland, the deposits at Mahibi

Hill and Fernao Veloso in Mogambique, and the ‘Trigonia schwarzi Beds’ of

Tanzania. The Hauterivian appears to have been largely a period of marine

regression.

Along the east coast Barremian deposits are known from Somalia and

Ethiopia (Furon 1963), Tanzania, Zululand, and also Madagascar. The deposits

are generally of limited extent and poorly fossiliferous, although they would

seem to date a transgressive episode. Possibly the ‘Aptienne inférieure ou ante-

aptienne’ transgression of Angola (Hoppener 1958) was contemporaneous.

The widespread occurrence of upper Lower Aptian strata in southern

Mogambique suggests that this was a period of marine transgression, not

Upper Aptian as suggested by Furon (1963: 362), a fact supported by the trans-

gressive Lower Aptian deposits of the Mangoky sector of Madagascar (Haughton

1963: 266). Strata with rich Cheloniceras faunas are also known from Somalia

and Tanzania, and are undoubtedly related to this transgression. The Upper

Aptian was generally regressive, although the sea did not leave the continental

margin until the end of the Aptian. In Zululand the Aptian—Albian boundary

is a non-sequence (Kennedy & Klinger 1971), as also in Madagascar (Coltignon

1963): 2).

100 ANNALS OF THE SOUTH AFRICAN MUSEUM

By the beginning of the Albian the west coast rift was open at least as far

as Nigeria, and the first truly marine transgression, dated at upper Lower Albian

(mammillatum Zone), affected both east and west coasts, indicating it to have

been eustatic in origin. This was followed on both coasts by continuous deposi-

tion until the end of the Albian although in Angola, above beds with Oxytropi-

doceras, the Middle Albian is unfossiliferous, suggesting a slight regression. It

is thus interesting to note Owen’s (1971: 138) remark ‘Superficially, it seems

that there is in many parts of the Earth a major break in sedimentation, par-

ticularly at the top of the Middle Albian. General sedimentation occurred once

again in early Upper Albian times.’ The mammillatum Zone transgression appears

locally to have attained its maximum extent only at the beginning of the Middle

Albian, e.g. Nigeria, and in the Rodo Valley of Madagascar (Haughton 1963:

264).

At the beginning of the Upper Albian there was a widespread west coast

transgression, followed by a slow regression for the remainder of the Albian,

with faunal evidence (Reyment & Tait 1972) suggesting West Africa and Brazil

to still have been connected during the Middle Albian. Reyment (1969) con-

siders that for a short time during the Upper Albian the South Atlantic appears

to have been open along its entire extent. The opening can possibly be related

to flooding of the Nigeria~Pernambuco connection by this transgression. The

occurrence of Upper Albian strata, apparently overlapping Lower Aptian

sediments, near Porto Henrique in Mocambique suggests this transgression was

eustatic.

The final break between Africa and South America is believed by Reyment

(1969) to have occurred only at the end of the Lower Turonian. With regard to

the Cenomanian, Reyment & Tait (1972: 64) wrote “The role of the Cenomanian

in the Atlantic is important. In all cases known to us the deposits of Cenomanian

age in the South Atlantic coastal basins accumulated in a regressive sea.’

In Angola, however, Lower Cenomanian strata are known from far inland

both at Cabiri and Novo Redondo, while uppermost Albian is not known from

these localities. There seems little doubt that in Angola the Lower Cenomanian

was a period of transgression. Possibly the somewhat younger beds with

Turrilites costatus Lamarck along the Mahube River in southern Mocambique

are also related to this transgression, resting as they do upon Lower Aptian

strata. Following this transgression, there appears to have been a gradual

regression until the middle Middle Cenomanian. The highest Cenomanian beds

at Novo Redondo in Angola contain Turrilites acutus, Forbesiceras obtectum,

Euomphaloceras cunningtoni, etc. A similar regression would appear to have

occurred in Nigeria following deposition of beds with Acanthoceras quadratum

Crick, Turrilites sp., Forbesiceras sp., etc. In Zululand a rich Middle Cenomanian

fauna with Turrilites acutus, Acanthoceras quadratum, etc., is overlain by poorly

fossiliferous sediments of low Upper Cenomanian age, The paucity of fossils

in these Upper Cenomanian sediments is possibly related to the slight change in

environment caused by this Middle Cenomanian regression. This synchronous

THE CRETACEOUS STRATIGRAPHY OF SOUTH—CENTRAL AFRICA 10]

Fig. 9. Africa during uppermost Cenomanian times. T marks the onshore occur-

rences of uppermost Cenomanian transgressive deposits, with the stippled area

indicating the extent of the Neolobites transgression in North Africa. R marks

Cenomanian localities at which the uppermost Cenomanian is a non-sequence.

In part after Reyment 1969.

regression on both sides of the continent suggests the regression was eustatic

and not tectonic as suggested by Reyment (1969).

During the uppermost Cenomanian (Metoicoceras gourdoni Zone) the sea

again transgressed along the west coast, with deposition of the well-known

Salinas fauna of Angola, the ‘Exogyra Beds’ at Bogenfels in South West Africa,

and the beds containing Meroicoceras aff. ornatum Moreman in Nigeria. This

transgression is also indicated by the Neo/obites faunas of North Africa (Fig. 9).

Following this transgression, at San Nicolau in Angola, the sea appears to

have remained static, with minimal sediment input (represented by a condensed

sequence), until earliest Coniacian times. In Zululand, however, uppermost

Cenomanian and Turonian rocks are absent, and low Upper Cenomanian

strata are unconformably overlain by Lower Coniacian sediments. The hiatus

102 ANNALS OF THE SOUTH AFRICAN MUSEUM

in Zululand is represented by a hardground, indicating submarine erosion

(Rose 1970) and non-deposition. Thus, for almost the exact period represented

by the condensed sequence at San Nicolau on the west coast, there was sub-

marine erosion and non-deposition along the east coast. As pointed out by

Jenkyns (1971: 330) “There is an obvious genetic connection between condensed

sequences and hardgrounds; condensed sequences are produced by minimal net

sedimentation, hardgrounds by nil or negative sedimentation’. Thus, condensa-

tion to the west and non-deposition to the east represent slightly differing

manifestations of the same regional event.

The absence in Angola of such typical early Turonian forms as Wright-

oceras, Bauchioceras, Paravascoceras, Fagesia, Paramammites, Nigericeras, etc.,

indicates, as suggested by Reyment (1969), that these faunas in West Africa

were derived by a transgression from the north, i.e. across the Sahara, the sea

only reaching West Africa during the Lower Turonian (Reyment 1971).

At San Nicolau, in Angola, the Turonian-earliest Coniacian condensed

sequence is unconformably overlain by alkaline extrusives (Cooper 1972). The

absence of pillow-structures together with the unfossiliferous nature of the inter-

bedded sediments suggest subaerial eruption. These volcanics are overlain by a

Lower Santonian marine horizon with Texanites, Protexanites, Hauericeras,

and Damesites (Cooper 1972). Local marginal warping during the Coniacian

in Angola can possibly explain the subaerial deposition, the volcanics being

related to the tectonic instability of the time. This is supported by the fact that a

rich (?)Upper Coniacian fauna with Peroniceras dravidicum (Kossmat),

Gauthiericeras margae (Schliter), Barroisiceras haberfellneri (Hauer), etc., is

known from the Ogoové Basin of Gabon (Furon 1963). In South West Africa,

Martin (1953) has attributed the reversal of the drainage of the Tsondab Valley,

which now flows to the west, as having occurred at some time during the

Cretaceous. Possibly it occurred at the beginning of the Santonian.

Along the east coast of South Africa, Kennedy & Klinger (1971: 185) were

led to regard the base of the Senonian ‘as a diachronous transgressive horizon

sedimentation beginning earlier in the north than in the south’. The Mandajene

Formation of southern Mogambique can be related to this transgression as can

the deposits in Somalia with Heterotissotia and Proplacenticeras. A similar

transgression occurred along the west coast in Gabon, where Coniacian strata

rest discordantly upon Turonian rocks (Reyment & Tait 1972), and it would

seem to be eustatic in origin.

The Upper Santonian-Lower Campanian appears to have been, in

general, a period of regression. Lower Santonian texanitids are common along

the west coast, although the only record of Upper Santonian appears to be of

a typical Pseudoschloenbachia from Cameroun (Reyment in Miiller 1966: 140).

According to Reyment (1966) the Santonian was regressive in the Nigerian

basin, while Haughton (1963: 283) noted that the Upper Santonian deposits of

the Bas-Zaire ‘are less widespread, a regression having commenced in this sub-

stage’. Similarly, Furon (1963: 362) noted that the Santonian was generally

THE CRETACEOUS STRATIGRAPHY OF SOUTH—CENTRAL AFRICA 103

West Coast | East Coast

| es

= z=

O a = j

5 z > Z | Die he: Ra 8 ry

Bo) panne hia gis a mT ee ee ee

) a @ o o > 9 o Py ey o s

Se 0 cee? Senta Ria

8 ia Bahk

T aaa Tacoma ai |

| fx oe

Maastrichtian |

=| en a

Campanian

=|. | Bae Ae

Santonian

Coniacian ; & Meera ie i iil s

ip)

‘ A a

Turonian

2 | R

Cenomanian

1? >)

[ Bocoog, oe Sco each toenoenossnaccuacd PaceErcoood leDoRRSeeoc ‘T

Albian 2

es es eee ee eee 52 Keeascoeood eeeacos - ooo

ll 4 i si

Aptian

Sune a f

ey) - ii

1 R

Neocomian | He. Mw T

Fig. 10. A comparison of the known Cretaceous stratigraphic sequences along the

east and west coasts of south-central Africa. T marks the start of a marine trans-

gression while R marks regressive phases.

104 ANNALS OF THE SOUTH AFRICAN MUSEUM

regressive in Madagascar. Thus, this regression appears to have been eustatic.

The transgressive Lower Santonian of Angola is related to local coastal tilting,

and is not thought to be eustatic in origin. The Senonian beds at Bogenfels in

South West Africa probably have a similar origin.

A major transgression occurred towards the end of the Campanian,

beginning during Hoplitoplacenticeras vari Zone times but only reaching its

maximum extent somewhat later. The occurrence of similar late Cretaceous

outliers at Need’s Camp along the east coast, and possibly in the Buzi and

Sheringoma regions of southern Mocgambique, suggests this transgression was

also eustatic. This is supported by the transgressive Upper Campanian deposits

of the east coast of Madagascar (Furon 1963).

SUMMARY

A comparison of the Cretaceous sequences along the east and west coasts

of southern and central Africa has allowed the recognition of at least nine

episodes of marine transgression during the Cretaceous. These are dated at

uppermost Valanginian, Barremian, upper Lower Aptian, upper Lower Albian,

basal Upper Albian, Lower Cenomanian, uppermost Cenomanian, Lower

Coniacian, and Upper Campanian, by the associated ammonite faunas. They

appear to best be explained by eustatic changes in sea-level rather than oscilla-

tory tiltings of the African continent.

ACKNOWLEDGEMENTS

I am extremely grateful to Professor R. A. Reyment, Uppsala, for his con-

structive criticism of the original draft, as well as for many helpful suggestions.

Dr W. J. Kennedy and Dr H. C. Klinger critically reviewed the manuscript,

whilst also providing me with the latest stratigraphical details of their work in

Zululand, for which I am most grateful. Once again Dr Kennedy, with his

usual perspicacity, has saved me from a number of errors.

REFERENCES

ANTUNES, M. T. & SORNAY, J. 1969. Contribution 4 la connaissance du Crétacé supérieur de

Barra do Dande, Angola. Revta Fac. Ciénc. Univ. Lish. (2, C) 16: 65-104.

BARBER, W. 1957. Lower Turonian ammonites from north-eastern Nigeria. Bull. geol. Sury.

Nigeria 26: 1-86.

Basse, E. 1963. Quelques ammonites nouvelles du Crétacé supérieur d’Angola. Bull. Soc.

géol. Fr. 4: 871-876.

BATE, R. H. & BAytiss, D. D. 1973. The Cretaceous of the Wami River area, Tanzania. News/.

Stratigr. 2: 155-162.

BELMONTE, Y. C. 1966. Stratigraphie du bassin sédimentaire du Cameroun. Proc. W. Afr.

micropaleont. Collog. 2, Ibadan 1965: 7-24. Leiden: Brill.

BusNARDO, R. & CoTILLON, P. 1964. Stratigraphie du Crétacé inférieur dans la région des

gorges du Verdon (Basses-Alpes et Var). C.r. somm. Séanc. Soc. géol. Fr. 8: 321.

CARVALHO, G. S. de. 1961. Geologia do deserto de Mocgamedes (Angola). Mems Jta Invest.

Ultramar (2) 26: 1-227,

THE CRETACEOUS STRATIGRAPHY OF SOUTH—CENTRAL AFRICA 105

Casey, R. 1965. A monograph of the Ammonoidea of the Lower Greensand. Palaeontogr.

Soc. (Monogr.) 6: 399-546.

CuorFAT, P. 1903. Contributions a la connaissance géologique des colonies portugaises d’Afri-

que. 1. Le Crétacique de Conducia. Comungées Servs geol. Port. 1903: 1-32.

COoLLIGNON, M. 1963. Atlas des fossiles caractéristiques de Madagascar (Ammonites). X, Albien.

Tananarive: Service Géologique.

COLLIGNON, M. 1968. La coupe du Santonien et du Campanien a Mitraiky dans l’ouest de

Madagascar. C.r. Sem. Géol. Tananarive 1967: 21-29.

Cooper, M. R. 1972. The Cretaceous stratigraphy of San Nicolau and Salinas, Angola. Ann.

S. Afr. Mus. 60: 245-251.

Cooper, M. R. 1973a. A revision of the Lower Cretaceous (Upper Valanginian) Cephalopoda

from the Uitenhage Formation of South Africa. Unpubl. M.Sc. thesis, Univ. Natal,

Durban.

Cooper, M. R. 19735. Cenomanian ammonites from Novo Redondo, Angola. Ann. S. Afr.

Mus. 62: 41-67.

Cox, L. R. 1925. Cretaceous Gastropoda from Portuguese East Africa. Ann. Transv. Mus.

11: 201-216.

Crick, 1924. On Upper Cretaceous Cephalopoda from Portuguese East Africa. Trans. geol.

Soc. S. Afr. 26: 130-140.

DARTEVELLE, E. 1942. Le Crétacé supérieur de Mossamedes (Contribution a la géologie de

Angola). Bull. Soc. belge Géol. Paléont. Hydrol. 50: 186.

DreBoLp, P. 1962. Notes on the geology of the southern Cameroun. Occ. Pap. geol. Sury.

Nigeria 10.

DINGLE, R. V. & KLINGER, H. C. 1972. The stratigraphy and ostracod fauna of the Upper

Jurassic sediments from Brenton, in the Knysna outlier, Cape Province. Trans. R. Soc.

S. Afr. 40: 279-298.

Du Tort, A. L. 1954. The geology of South Africa. Edinburgh: Oliver & Boyd.

DovuviLLE, H. 1931. Contribution a la géologie de Angola. Bolm Mus. Lab. miner. geol.

Univ. Lisb. 1: 17-46.

Furon, R. 1963. The geology of Africa. Edinburgh: Oliver & Boyd.

GEVERS, T. & LitTLe, J. de V. 1946. Upper Cretaceous beds between the Itongazi and Umkand-

andhlovu Rivers, Alfred County, Natal. Trans. geol. Soc. S. Afr. 48: 27-29.

Haas, O. 1942a. The Vernay Collection of Cretaceous (Albian) ammonites from Angola.

Bull. Am. Mus. nat. Hist. 81: 1-224.

Haas, O. 19426. Some Upper Cretaceous ammonites from Angola. Am. Mus. Novit. 1182: 1-24.

Haas, O. 1943. Some abnormally coiled ammonites from the Upper Cretaceous of Angola.

Am. Mus. Novit. 1222: 1-17.

HAuGutTon, S. H. 1925. Notes on some Cretaceous fossils from Angola (Cephalopoda and

Echinoidea). Ann. S. Afr. Mus. 22: 263-288.

HAUGHTON, S. H. 1930. On the occurrence of Upper Cretaceous marine fossils near Bogenfels,

S.W. Africa. Trans. R. Soc. S. Afr. 18: 361-365.

HAUGHTON, S. H. 1963. Stratigraphic history of Africa south of the Sahara. Edinburgh: Oliver

& Boyd.

HAuGuton, S. H. & Bosuorr, J. C. 1956. Algumas amonites Aptianas de Chalala Gane

Oriental Portuguesa). alkip Servs Ind. Geol. Lourenco Marques 17: 1-24.

HENNIG, E. 1937. Der Sedimentstreifen des Lindi-Kilwa-Hinterlandes (Deutsch- Ostafrika).

ie rieconieeraphica Suppl. 7 (Reihe 2, Teil 2, Lfg. 2): 99-186.

Hoppener, H. 1958. Brief report on the palaeontology of the Cuanza basin—Angola. Bolm

Soc. eee! Port. 12: 75-82.

Howarth, M. K. 1965. Cretaceous ammonites and nautiloids from Angola. Bull. Br. Mus.

nat. ise (Geol.) 10: 335-412.

Howarth, M. K. 1968. A mid-Turonian ammonite fauna from the Mocamedes desert, Angola.

Gargia de Orta 14: 217-228.

JENKYNS, H. C. 1971. The genesis of condensed sequences in the Tethyan Jurassic. Lethaia 4:

327-352.

KENNEDY, W. J. 1971. Cenomanian ammonites from southern England. Spec. Pap. Palaeont.

8: 1-133.

KENNEDY, W. J., & KiinGeR, H. C. 1971. A major intra-Cretaceous unconformity in eastern

South Africa. J] geol. Soc. Lond. 127: 183-186.

106 ANNALS OF THE SOUTH AFRICAN MUSEUM

KENNEDY, W. J. , KAUFFMAN, E. G. & Klinger, H. C. 1973. Upper Cretaceous invertebrate

faunas from Durban, South Africa. Trans. geol. Soc. S. Afr. 76: 95-111.

KILIAN, W. 1902. Uber Aptien in Siidafrika. Ch/. Miner. Geol. Paldont. 3: 465-468.

Kina, L. C. 1972. Geomorphic significance of the late Cretaceous limestones at Needs Camp,

near East London. Trans. geol. Soc. S. Afr. 75: 1-3.

KLAsz, I. DE & GAGEONNET, R. 1965. Biostratigraphie du bassin gabonais. (Ist West African

Micropaleontological Colloquium, Dakar 1963.) Mém. Bur. Rech. géol. miniér. 32:

277-303.

KLINGER, H. C., KENNEDY, W. J. & DINGLE, R. V. 1972. A Jurassic ammonite from South

Africa. Neues Jb. Geol. Paldont. Mh. 11: 653-659.

KRENKEL, E. 1910. Die Aptfossilien der Delagoa-Bai (Sudostafrika). Neues Jb. Miner. Geol.

Paldont. 1: 142-168.

Lacey, W.S. 1961. Report on fossils from Chalala and Mangulane in the province of Mocam-

bique. Bolm Servs Geol. Minas Lourenco Marques 27: 5-15.

Lock, B. E. 1973. Tertiary limestones at Needs Camp, near East London. Trans. geol. Soc. S.

Afr. 76: 1-5.

Matsumoto, T. 1965. A monograph of the Collignoniceratidae of Hokkaido. Part I. (Studies

of the Cretaceous ammonites from Hokkaido and Saghalien—XIV.) Mem. Fac. Sci.

Kyushu Univ. (D) 16: 1-80.

Martin, H. 1953. Notes on the Dwyka succession and on some pre-Dwyka valleys in South

West Africa. Trans. Proc. geol. Soc. S. Afr. 56: 37-41.

McGowran, B. & Moore, A. C. 1971. A reptilian tooth and Upper Cretaceous microfossils

from the Lower Quarry at Needs Camp, South Africa. Trans. geol. Soc. S. Afr. 74: 103-105.

MULLER, K. J. 1966. Some remarks on the youngest conodonts. Proc. W. Afr. micropaleont.

Colloq. 2, Ibadan 1965: 137-140. Leiden: E. J. Brill.

Murphy, M. A. 1967. The Aptian—Cenomanian members of the ammonite genus Tetragonites.

Univ. Calif. Publs geol. Sci. 69: 1-78

Neumayr, M. 1885. Phylloceras semistriatum Orb. von Mossambique. Denkschr. Akad. Wiss

Wien 50: 139.

NewTon, R. B. 1924. A contribution to the palaeontology of Portuguese East Africa. Trans.

geol. Soc. S. Afr. 26: 141-159.

OweEN, H. G. 1971. Middle Albian stratigraphy in the Anglo-Paris basin. Bull. Br. Mus. nat.

Hist. (Geol.) Suppl. 8: 1-164.

QUENNELL, A. M., MCKINLAY, A. C. M. & AITKEN, W. G. 1956. Summary of the geology of

Tanganyika. Part I. Introduction and stratigraphy. Mem. geol. Surv. Tanganyika. 1: 1-264.

RENNIE, J. V. L. 1935. Upper Cretaceous Lamellibranchia from Incomanini, Portuguese East

Africa. Ann. Transy. Mus. 18: 325-347.

RENNIE, J. V. L. 1936. Lower Cretaceous Lamellibranchia from northern Zululand. Ann. S.

Afr. Mus. 31: 277-391.

REYMENT, R. A. 1954a. New Turonian (Cretaceous) ammonite genera from Nigeria. Colon.

Geol. Miner. Resour. 4: 149-164.

REYMENT, R. A. 19545. Some new Upper Cretaceous ammonites from Nigeria. Colon. Geol.

Miner. Resour. 4: 248-270.

REYMENT, R. A. 1955. The Cretaceous Ammonoidea of southern Nigeria and the southern

Cameroons. Bull. geol. Surv. Nigeria 25: 1-112.

REYMENT, R. A. 1965. Aspects of the geology of Nigeria. Ibadan: Ibadan University Press.

REYMENT, R. A. 1966. Brief review of the stratigraphic sequences of West Africa (Angola to

Senegal). Proc. W. Afr. micropaleont. Colloq 2, Ibadan 1965: 162-175. Leiden: E. J. Brill.

REYMENT, R. A. 1969. Ammonite biostratigraphy, continental drift and oscillatory transgres-

sions. Nature, Lond. 224: 137-140.

REYMENT, R. A. 1971. Experimental studies of Cretaceous transgressions for Africa. Bull.

geol. Soc. Am. 82: 1063-1072.

REYMENT, R. A. & Tait, E. A. 1972. Biostratigraphical dating of the early history of the South

Atlantic Ocean. Phil. Trans. R. Soc. (B) 264: 55-95.

Rose, P. R. 1970. Stratigraphic interpretation of submarine versus subaerial discontinuity

surfaces: an example from the Cretaceous of Texas. Bull. geol. Soc. Am. 81: 267-279.

Soares, A. F. & SitvA, G. H. pA. 1970. Contribuigao para o estudo da geologia do Maputo.

Estratigrafia e paleontologia da regiaio de Madubula e suas relagdes com areas vizinhas.

Revta Ciénc. geol. Lourenco Marques (A) 3: 1-85.

THE CRETACEOUS STRATIGRAPHY OF SOUTH—CENTRAL AFRICA 107

SitvA, G. H. DA. 1962. Amonites do Cretacico inferior do rio Maputo (Catuane—Mocam-

bique). Bolm Servs Geol. Minas Lourenco Marques 29: 7-32.

SitvA, H. DA. 1966. Sobre a ocorréncia do Jurassico marinho no Norte de Mocgambique.

Reyvta Estud. ger. Univ. Mocambique (2) 3: 61-68.

SPATH, L. F. 1921. On Cretaceous Cephalopoda from Zululand. Ann. S. Afr. Mus. 12: 217-321.

SpATH, L. F. 1925. On Upper Albian Ammonoidea from Portuguese East Africa. With an

appendix on Upper Cretaceous ammonites from Maputoland. Ann. Transy. Mus. 11:

179-200.

SpATH, L. F. 1930. On the Cephalopoda of the Uitenhage Beds. Ann. S. Afr. Mus. 28: 131-157.

SpaTH, L. F. 1951. Preliminary notice on some Upper Cretaceous ammonite faunas from

Angola. Comungées Servs geol. Port. 32: 123-130.

SPATH, L. F. 1953. The Upper Cretaceous cephalopod fauna of Graham Land. Scient. Rep.

Falkld Isl. Depend. Surv. 3: 1-60.

TAVANI, G. 1949. Fauna Malacologica Cretacea della Somalia e dell’ Ogaden (2. Gastropoda —

Cephalopoda—conclusioni). Palaeontogr. ital. 45: 1-76.

WACHENDORF, H. 1968. Zur Unterkreide-Stratigraphie von Sud-Mocambique. Neues Jb. Geol.

Paldont. Abh. 129: 272-303.

Wray, D. A. 1915. Observations sur la géologie du district de Mocgambique. Comungdes

Servs geol. Port. 11: 69-84.

ZWIERZYCKI, J. 1913. Zur Frage der Unteren Kreide in Portugiesisch-Mozambique. Sber.

Ges. naturf. Freunde Berl. 7: 319-326.

ZWIERZYCKI, J. 1914. Die Cephalopodenfauna der Tendaguruschichten in Deutsch-Ostafrika.

Arch. Biontol. (3) 4: 7-96.

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THE ZOOGEOGRAPHY OF THE SOUTH AFRICAN AVIFAUNA

J. M. WINTERBOTTOM

South African Museum, Cape Town

(With 9 maps and 6 tables)

[MS accepted I November 1973]

CONTENTS

PAGE

Introduction . : : : a 09

Lowland and plfeane : : : eh dls

East African Coastal District . ‘ : 120

South Central Highlands District : ae pS

Western Districts : : j ; 124

Winter Rainfall Area . , ; tn LAS)

Karoo : 3 : : ; ; 130

Highveld . ; F : : a 50

Transitional Zone . A é , ; 131

Montane . : : : : : : 132

Freshwater : : f 3 ; ; 135

Coastal . 4 ‘ : ; ; ; 137

Oceanic . : : : ; ; y 137

Summary. ; : : 5 ; : 139

Acknowledgements . ; : ; : 140

References 5 : Q : ; i 140

Appendix. : : : A 5 : 144

INTRODUCTION

From the days of P. L. Sclater (1858) and A. R. Wallace (1876), Africa

south of the Sahara has been recognized as one of the major faunal regions of

the globe. Wallace, in subdividing this, the Ethiopian Region, constituted the

area south of the Zambezi and Kunene Rivers as the South African Sub-region

and this is the part of the continent dealt with in this paper.

The fauna of any area depends partly on geography and partly on climate,

the latter working on animals chiefly through the vegetation. South Africa is

the southern extremity of a continent separated on three sides from other land

masses by a wide expanse of ocean and on the fourth by deserts. It is generally

agreed that the African fauna has most in common with that of the Oriental

Region, especially in the case of the forest fauna (De Beaufort 1951); and that

the land fauna is sharply divided into forest and non-forest. It is also frequently

stated (e.g. by De Beaufort 1951 and Cloudesley-Thompson 1969) that the

forest fauna is the older of the two, but I have recently (Winterbottom 1973)

given reasons for doubting this. The Ethiopian Region was isolated from the

north by the Tethys Sea until the Miocene and by that time had developed its

Ann. S. Afr. Mus. 66 (6), 1974: 109-149, 9 maps, 6 tables.

109

110 ANNALS OF THE SOUTH AFRICAN MUSEUM

HOE ARC. tule

Map 1. Zoogeographical regions of the world.

own individuality. The great expansion of grasslands during this period must

have led to considerable speciation (Brodkorb 1960). Although there were times,

at least during the Pleistocene, when the present Sahara Desert was inhabitable

for a wide range of animals, it is important to note the very considerable climatic

differences between north and south. The present bird and butterfly fauna of

North America is almost entirely Palaearctic, showing that the Ethiopian

elements cannot compete with them in the temperate climate north of the

Sahara and the Palaearctic fauna is at a similar disadvantage in the tropical

Sudan. It is important to stress this becase the temperate areas south of the

Equator are inhabited by a fauna totally distinct from the Palaearctic and clearly

derived from the tropical fauna to the north. Much of South Africa has a

temperate climate and its fauna, though related to the tropical fauna to the

north, has obviously been in existence for a long time, since it has allowed the

evolution of at least one distinct family, the Promeropidae, and a number of

distinct genera, such as Geocolaptes and Chaetops. Hall (1970) finds that the

African passerine fauna is relatively rich, as compared with the Oriental and

Palaearctic, in fruit-eaters (8°) and seed-eaters (24%). The South African

avifauna, with only 2° fruit-eaters and 18° seed-eaters, is markedly less so

than the African fauna as a whole.

As with the temperate fauna in the extreme south-west, the desert fauna of

South Africa has long been isolated, or almost isolated, from the deserts to the

north, and genera such as Namibornis, Lanioturdus and Philetairus have evolved

THE ZOOGEOGRAPHY OF THE SOUTH AFRICAN AVIFAUNA ay

there. The genus Namibornis is a particularly enlightening one in this respect,

since it appears to be intermediate between the Turdine genus Cercome/a and

the Muscicapine genus Me/aenornis, so its ancestors must have arrived before

the groups diverged (Jensen & Jensen 1971). The question of links with the north-

eastern desert area of Somaliland is discussed below.

Of the endemic South African species, the majority, as might be expected,

have their nearest relatives either in South Africa itself or in adjoining areas to

the north. Apart from the southwest—northeast species pairs, which are discussed

below, there are five whose nearest relatives occur in the Palaearctic and about

14 whose affinities are obscure but which have obviously had a long history in

South Africa.

In summary, the terrestrial avifauna of South Africa contains 6° of

species which are found in other zoogeographical regions, 33° which range

widely over the whole Ethiopian Region, and 34% which occur also in East

as well as South Africa (Table 1). Among the non-passerines, the figures for the

first two of these categories are higher than in the passerines, being 12° for

cosmopolitan and 40° for Ethiopian species.

Table 1

Geographical affinities of breeding South African land birds (percentages)

South S.W.- West Total

Cosmo- Ethio- & East N.E. Central South No. of

politan pian Africa only Africa Africa Species

Non-passerine 12 40 32 4 0 iN) 225

Passerine Z 28 35 DD 2 31 241

All species 6 33 34 1 1 DS 466

If we consider the South African terrestrial avifauna at the generic level, we

need to use wider categories. These, as used in Table 2, are: Cosmopolitan,

being genera represented in both Old and New Worlds and in at least four

Table 2

Geographical affinities of genera of breeding South African land birds (percentages)

: coset dee: ee hae

= “3 3 3 at elas zz § S)

a = a) ~ as go q = eS fe)

Bb 8 8g 2) 33 2 2, < 2;

ane Seas Woy om We Mioe) “SS as | apg

ee se Se Soe Ee. ee

3S) re) aa ea) ea] aly OZ aa n< n KO

Non-passerine 22 22 3 1 4 1 44 0 1 96

Passerine vA 16 0 6 4 0 58 2 7 120

All species 13 19 1 4 4 1 53 1 5 216

il 2 ANNALS OF THE SOUTH AFRICAN MUSEUM

zoogeographical regions; Old World, being genera represented in at least three

of the four regions of the Old World; Pan-tropical Old World, genera represented

in the Ethiopian, Oriental and Australian Regions but not in the Palaearctic;

Ethiopian and Oriental; Ethiopian and Palaearctic; Ethiopian and Neotropical;

Ethiopian, being genera widespread in the Region; South and East African; and

South African. As at the species level, the percentage with Cosmopolitan

affinities is markedly higher among the non-passerines than among the pas-

serines (to obtain a comparable total to the Cosmopolitan species in Table 1,

the first six columns of Table 2 should be summed); but the percentage of

Ethiopian genera is much higher than the percentage of Ethiopian species (as

might be expected) and is higher in the passerines than in the non-passerines;

but the percentage of South African genera is small, confirming the view that

South Africa does not merit rank as a major division of the Ethiopian

Region.

Udvardy (1969) analyses the tendency for species to change their range by

the use of what he terms ‘dynamic potential’. Species with distribution limits in

the area being considered constitute the Intrinsic Dynamic Potential; species

which moved their limit during the time period considered represent the Realized

Dynamic Potential and those which did not move, the Unrealized Dynamic

Potential. He gives a table to show this in respect of the Carpathian Basin,

1860-1960. However, the application of this type of analysis to South Africa is

complicated by several factors:

(i) Lying as it does at the end of a continent, all the breeding species on the

South African list (665, excluding sea birds) find their limits in the area, so that

the Intrinsic Dynamic Potential is 665.

(ii) Since the area is bounded by the sea on the west, south and east, all

movement must be from the north.

(iii) Owing to the great size of the area, it is necessary to introduce an

additional category to those used by Udvardy, namely species advancing in the

area. There are 8 of these for the period 1900-1970; 9 are receding; 3 are

sporadic breeders; and 1 is newly arrived, giving a Realized Dynamic Potential

of 21 (Carpathian Basin, 37). This suggests that the South African avifauna as a

whole is rather static.

In discussing the zoogeography of southern Africa, it is essential to realize

that five different patterns are involved, since the barriers which are effective for

one set of species are the areas of distribution of others. This is obvious if we

consider the sea and the land, the former being a barrier to land birds and the

latter to sea birds. The five patterns here mentioned are:

(i) oceanic birds;

(11) coastal birds;

(ii1) freshwater birds;

(iv) montane birds;

(v) the rest.

THE ZOOGEOGRAPHY OF THE SOUTH AFRICAN AVIFAUNA 113

These patterns become more and more complicated as one proceeds from (i) to

(v) and, especially in (v), it is exceedingly difficult to disentangle geographical

from ecological factors. Since (v) is much the most widespread and has excited

greatest comment, I propose to deal with these patterns in the reverse order to

that listed above.

LOWLAND AND PLATEAU

The lowland and plateau areas, which occupy most of the land of South

Africa, vary in their climate and vegetation from the bare sand-dunes of the

Namib Desert* to the close woodland of the Baikiaea ‘mutemwa’ of western

Rhodesia and north-east Botswana and the sometimes impenetrable thickets of

the macchia vegetation of the southern Cape. The geographical distribution of

avian species within this complex is therefore naturally very varied. I made a

list of 51 species which are found throughout, or almost throughout, South

Africa; but it by no means follows that because a species has a wide geographical

range, it also has a wide ecological tolerance. Of the 51 species mentioned above,

three do not appear in any of the lists of dominant or abundant species in the

different habitats considered by Winterbottom (19726). Those with much the

widest ecological tolerance appear to be the two doves Streptopelia capicola and

S. senegalensis, followed by Lanius collaris, Dicrurus adsimilis, Ploceus velatus

and Sy/vietta rufescens. Generally speaking, therefore, even species very wide-

spread geographically are limited to one or a few habitats and occur only in

them, so that, in effect, their distribution is patchy rather than general.

There have been a number of attempts to deal with the zoogeography of

South Africa. Wallace (1876) made it one of the four sub-regions into which he

divided the Ethiopian Region; Chapin (1932) divided it into the South West

Arid and South East Veld Districts, the boundaries of each of which he extended

rather beyond Wallace’s (and our) limits of the Zambezi and Kunene. Bowen

(1933) produced an entirely different arrangement. Unfortunately, his maps are

on so small a scale that it is extremely difficult to make out the boundaries he

proposes; within our area, however, he recognizes two zones, four ‘climates’ and

five districts. Of these, his Rhodesian Savanna District is roughly equivalent to

our South Central Highlands; his Southeast Veldt to our Macchia and Highveld,

with the southern Karoo and Rhodesian plateau added; and his East African

Lowland to the two northern divisions of our East African Coastal. In the

south-west, he divides the area in two by a line running down the centre of

Damaraland—an impossible division: but if more realistically adjusted, this

may be taken as marking the boundary between our South Temperate and South

West Arid Districts. He leaves the coastal areas from Beira to Cape Town as a

transition zone, ‘in which both Tropical and Subtropical Zone species may

occur’.

Moreau (1952) divided South Africa in three, splitting off the south-west

* Places mentioned in the text are shown on Map 9.

114 ANNALS OF THE SOUTH AFRICAN MUSEUM

Map 2. Zoogeographical divisions of South Africa according to A. Chapin;

B. Bowen; C. Moreau; D. Poynton.

Map B

THE ZOOGEOGRAPHY OF THE SOUTH AFRICAN AVIFAUNA 115

10° [5° 20° age 30° 35°

Map D

116 ANNALS OF THE SOUTH AFRICAN MUSEUM

Cape from the rest and uniting Chapin’s South East Veld with vast areas the to

north as a Southern Savanna District. Winterbottom (1959) removed the High-

veld grasslands from Chapin’s South East Veld to his South West Arid and

divided the rest of that district between his Rhodesian Highlands and East

African Coastal Districts. Poynton (1964), like Moreau, divided off the south-

western Cape; he put most of the South West Arid into a Temperate Western

and the rest into a Subtropical Arid, except for the extreme north, where he pro-

posed a division running right across from Ovamboland to the eastern Highlands

of Rhodesia as a Subtropical Moist. He retained the highveld as a Temperate

Eastern. The coastal strip north to about St. Lucia, with the Transvaal and

Swaziland lowveld, made up a Subtropical Moist and the rest of the low country

on the east, including the Zambezi valley to the Victoria Falls, was his Tropical

division. Poynton’s divisions were based on the distribution of the Amphibia.

His distinction of the South-West Cape is entirely justifiable in respect of its

amphibian fauna but not for birds and mammals.

One further attempt needs to be discussed. Roberts (1940) proposed to

divide South Africa into 21 ‘small regions’, a concept which was taken over, with

minor modifications, by McLachlan & Liversidge (1957, 1970). These, although

not so called by the authors, are, in effect, ‘biotic provinces’ as developed by

Dice (1952). Of the divisions they propose, their Southern and Eastern Ever-

green Forest, Southern and South-western Cape and a few others have some

validity, though the detailed evidence on which they are based has never been

published. Many of the others, however, have no biological reality at all and the

birds which the authors cite, when they do list any, do not support their con-

clusions. Thus the only bird listed for the Little Karoo is Circus maurus, which I

personally have never seen there, though no doubt it does occur; but which is

found all over their Southern and South-western Cape, their Karoo, their

Eastern Grassveld and much of their Great Namaqualand and Highveld Grass-

veld. They cite four species for their Thornveld Kalahari, Turdoides bicolor,

Laniarius atrococcineus, Mirafra africanoides and Erythropygia paena, all of

which occur almost throughout the South West Arid District. Among their

Damaraland species, they list Tockus bradfieldi, whose range barely reaches the

extreme north-east of this division but extends across the northern parts of their

Sandveld Kalahari and Ngamiland to western Rhodesia and which they also

list as the sole species for their Rhodesian Woodlands.

As far as can be judged from their map, Durban and St. Lucia are in their

Eastern Littoral, Ndumu in their Eastern Lowveld and Lourenco Marques in

their Eastern Tropical Littoral. Analysis of the avifaunas of these places shows

that St. Lucia has much more in common with Ndumu than with Durban; that

Lourengo Marques has an avifauna as similar to that of Ndumu as St. Lucia’s

is; and that the avifauna of Durban has more in common with that of East

London than with that of St. Lucia. This is in accord with the suggestions made

below, which would group St. Lucia and Ndumu with Lourengo Marques in

one sub-district of the East African Coastal District and Durban and East

THE ZOOGEOGRAPHY OF THE SOUTH AFRICAN AVIFAUNA ey

London in another, with the area between St. Lucia and Durban as a transitional

zone.

In an endeavour to apply McLachlan & Liversidge’s concepts more closely,

to see what would come of a detailed analysis, I drew up a list of species not

distributed all over South Africa; listed those which occurred in each magisterial

district (and sub-district in the western Cape, as set out in Winterbottom, 19685)

in the Cape Province south of the Orange River; and compared the lists for each

district so obtained with those of all the adjoining districts, using the Co-efficient

of Community as defined by the formula:

ii =P fy = © cee

where c is the number of species in common and 7, n, are the numbers in the

smaller and larger faunas respectively. Lines were then drawn through district

)

/Eastern Karoo

rae [ite Karoo.

Map 3. Avifaunal areas of the Cape Province south of the Orange River.

boundaries where the co-efficients indicated faunal change. The results are shown

in Map 3 and differ considerably from McLachlan & Liversidge’s ‘small regions’

for the same area. The number of species used in the analysis was 329.

Apart from the unsatisfactory nature of the actual divisions proposed, the

whole concept of biotic provinces, like those of life zones and faunal groups (on

the second of which, see Winterbottom 1965), is an attempt to find a way out of

the difficulties—very real difficulties—of the classical method of regions and

districts. Unfortunately, however, these methods, at least to my mind, produce

results even less satisfactory than those they were designed to supersede.

It will be realized from what has been said above and what follows, that

modern zoogeographers reject Wallace’s division of South Africa as a distinct

sub-region of the Ethiopian Region and that the boundaries of the various zoo-

geographical areas which have been proposed do not coincide with the Zambezi

and Kunene Rivers at any point. For various reasons, the present paper is

118 ANNALS OF THE SOUTH AFRICAN MUSEUM

primarily concerned with Wallace’s sub-region, to which about 130 species are

restricted, but it is appreciated that all those subdivisions which reach the

Zambezi or Kunene continue north of the rivers, though the avifauna of these

northern areas is not considered in detail here.

My own views on the zoogeographical subdivisions in South Africa are

shown in Map 4. It will be seen that there are four districts (apart from montane

forest) partly or wholly within the area: the South Central Highlands, the East

African Coastal, the South West Arid and the Southern Temperate; plus an

extended Transitional Zone between the first and third. Comments on, and

justification of, the establishment of these districts follow.

The most fundamental pattern in bird distribution in South Africa is the

dichotomy between east and west. I have listed 106 species whose distribution

36 |

1. South Central Highlands District

2. South West Arid District

A. Damaraland Sub-district

B. Kalahari Sandveld Sub-district

C. Kalahari Thornveld Sub-district

3. East African Coastal District

G. Mocambique Sub-district

H. Southern Sub-district

. Transitional Zone

. South Temperate District

D. Karoo Sub-district

E. Highveld Sub-district

F. Macchia Sub-district

ai >

Map 4. Zoogeographical regions of South Africa.

does not extend west of the main eastern escarpment, though some of them

occur in the montane forest of that area; and 103 whose distribution, at least

north of Port Elizabeth, does not extend east of the same escarpment. There is

a third common pattern, that of species which occur in the north, often south to

the Limpopo or even the northern Transvaal, in the west but extend further

south, into Natal and sometimes the eastern Cape, in the east. These are species

whose main centre of distribution lies north of our limits.

Although the discussion in this paper is mainly of the geographical distribu-

tion patterns, these must obviously be influenced by ecological conditions. They

are not discussed in detail here since that has already been done elsewhere

(Winterbottom 1972a). Hall & Moreau (1970) say ‘It is the distribution of

THE ZOOGEOGRAPHY OF THE SOUTH AFRICAN AVIFAUNA 119

vegetation which governs the geographical lines along which speciation has

taken place’. But things are by no means as simple as this, as perusal of my

ecological paper will show. In fact, Hall & Moreau use as their base map (as does

Moreau 1966) a map produced by Cooke (1962) which is by no means always

suitable to show the relations between bird and plant distribution. This is

particularly the case with the ‘Dry Woodland, etc., mostly Acacia’, which is

extended east to include the highveld grasslands and thus obscures the ecological

preferences of over 40 species.

It is nevertheless important for zoogeographical purposes to know something

of the major habitats available to birds in each of the areas delimited in Map 4.

1. South Central Highlands

Most of the area is covered with Brachystegia woodland (miombo), fairly

close woodland of rather small, deciduous trees with grass ground cover. There

are also stretches of Baikiaea woodland of somewhat similar character and of

mopane woodland, more open and with more scattered grass. In Rhodesia, and

also in extra-limital Zambia, there are stretches of open grassland with scattered

Acacia trees.

2. East African Coast

Chiefly scrub Acacia woodland, with patches of temperate forest.

3. South West Arid

Chiefly Acacia woodland on sand, with more open grassland in the south-

east.

4. Southern Temperate

Macchia—bushes of Protea, Erica, etc.—in the extreme south; Karoo—

small, scattered bushes; with bare gravel and sand-dunes in the Namib; High-

veld—open grassland.

The climatic vicissitudes of the Pleistocene must have changed the distribu-

tion of these vegetation types; but it would not appear to have eliminated or

drastically reduced or seriously fragmented any of them. Clark (1967) produces

a map showing the vegetation at 50° of the present rainfall; and while it is

unlikely that the rainfall decreased simultaneously over the whole continent,

Clark’s map does support the conclusion that an arid corridor linked, or almost

linked, the South West Arid District with the Somali Arid, perhaps several

times. This explains the curious relationships between the avifaunas of the two

districts. The following species are found in both, but not in the intervening

areas:

Poliohierax semitorquatus, Eupodotis ruficrista, Mirafra africanoides,

Turdoides melanops, Nectarinia mariquensis, Ploceus rubigenosus, Passer motiten-

sis, Bubalornis albirostris, Estrilda erythronotus and Amadina fasciata.

A number of other species (e.g. Pterocles gutturalis, Francolinus sephaena)

have been excluded because they range rather far beyond South West Arid

limits. In addition, there are over 20 cases of closely-related pairs of species,

120 ANNALS OF THE SOUTH AFRICAN MUSEUM

some of which some systematists regard as conspecific. Yet the presence of

endemic genera in each area suggests that there was only a limited exchange of

fauna. The question has been more fully discussed elsewhere and an analysis of

the avifaunas of the South West Arid and Southern Temperate Districts (Winter-

bottom 1972c) shows that of the 90 characteristic species, 34 are specifically

identical or closely allied to species otherwise found in the Somali Arid District;

24 are nearly related to species which occur further east in South Africa; and the

32 remaining forms are peculiar to these districts and their origins are in many

cases obscure. Several of them, however, suggest more than one invasion from

the north and/or radiation in the south-west. The Pleistocene arid periods, when

the Kalahari sands were blown deep into the Congo basin, have left south-west

arid relicts further north—the populations of Certhilauda albofasciata in

Angola and Tanzania are examples.

While the temperate forest faunas have been repeatedly isolated and then

joined again, a process which facilitates and accelerates speciation, this has not

been the case with the non-forest habitats. Keast (1972) has pointed out that in

Australia the desert 1s central and as it expanded and contracted, the fauna of

the more mesic habitats were alternately driven into ‘refuges’ and then brought

together again, while the desert fauna itself remained undivided and was

presented with little opportunity for speciation. In Africa, however, it is the

deserts which are peripheral. Contact between the northern deserts and those of

the south-west has been minimal and the faunas are decidedly different (Winter-

bottom 1972c). The two existing deserts in the south-west are separated by the

somewhat less arid Karoo and are decidedly different in character. The Namib,

as shown by its remarkable flora and arthropod and herpetological fauna, is

very old; but it appears to have always been of very restricted area. The Kalahari,

by contrast, has undoubtedly fluctuated greatly in size. At the height of arid

periods, Kalahari-type vegetation apparently extended right across the continent

in the Limpopo basin to the Indian Ocean and northward deep into the Congo

basin. At such times, the south-western portions were under a much more severe

desert regime than they are today. On the other hand, during pluvial periods, the

northern and north-western parts of the present desert would have been bush-

veld. Nevertheless, at no time would either desert have been fragmented.

The avifauna of the Namib contains several endemic bird species; that of

the Kalahari does not—its avifauna is merely an impoverished version of that

of the Acacia steppe further west.

EAST AFRICAN COASTAL DISTRICT

The two major groups of east and west mentioned above characterize what

Winterbottom (1959) defined as the South West Arid and South East Coastal

Districts. The latter has two sub-districts within our limits, besides a third north

of the Rovuma River in East Africa. The division between the other sub-districts

may be drawn at St. Lucia, south-west of which there is a rapid fall-off in

tropical species; but the area from St. Lucia to Durban is regarded as a transi-

THE ZOOGEOGRAPHY OF THE SOUTH AFRICAN AVIFAUNA je |

tional zone between the two; and, as will be seen below, the fixing of the boun-

dary is by no means as simple as some authors have assumed, the whole of the

southern part being, as it were, a subtraction zone at the end of the tropical belt

further north.

Of the characteristic species of the northern of the two sub-districts, two

drop out between Ndumu and St. Lucia, nine between St. Lucia and Durban and

12 between Durban and Port St. Johns (these figures are based on Pooley &

Dixon (1966) for Ndumu; Natal Parks Board (undated) for St. Lucia; Lawson

(1971) for Durban; and McCulloch, Skead & Winterbottom (1970) for Port St.

Johns). The 23 tropical species which drop out along this stretch of coastal

plain are:

Guttera edouardi, Tauraco porphyriolophus, Ceuthmochares aereus, Eurysto-

mus glaucurus, Stactolaema leucotis, Pogoniulus bilineatus, Campethera abingoni,

Smithornis capensis, Andropadus flaviventris, Nicator chloris, Cossypha heuglini,

Erythropygia quadrivirgata, Apalis ruddi, Batis fratrum, Platysteira peltata, Mala-

conotus quadricolor, Prionops scopifrons, Nectarinia neergaardi, N. bifasciata,

N. senegalensis, Zosterops senegalensis, Hypargos margaritatus and Lagonosticta

senegala.

B ;

Ekman’s Index of faunal change* , Where A is the number of species

in the larger fauna, B in the smaller fauna and C of the species in common, is

largest (2,7) between Port Alfred and the Tsitsikama and lies between 2,2 and 2,4

between other localities from Ndumu to Swellendam, at the south-west boundary

of the district (Map 5).

* Ekman’s Index is actually much more complicated than this and I have not attempted to

assess his ‘positive zoogeographical values’—see Udvardy (1969) for a discussion.

Ndumu

C Bi wie

Apart st Johns

Tsitsikama

Sectors Gy oF iss London

oS 2

yy VI SSS SS Prt Alfred

Map 5. Ekman’s Index along the south-east coast.

[22 ANNALS OF THE SOUTH AFRICAN MUSEUM

Another method of measuring faunal change is by Schilder’s Index

(Udvardy 1969). The Percentage Occurrence Index derived from this, in respect

of the East African Coastal District between Ndumu and Swellendam would

consist, for each station, of the number of species present at Ndumu but not at

Swellendam in each plus the number of species present at Swellendam absent

from the other stations, expressed as a percentage of the Ndumu figure (which

will, obviously, be 100%). The full list is:

Netaimiaiee se 2 eke ee 0)

St; Oca 2s), oe, Ae ee

DUEbane we wars Bi" ar) Cane ae

Port: Seonns.e | 2.0 ke eee ee

Bast-vondon v5) re 2 ee eee

PortvAlized 0s (ee oe oS

Psitsikeammaes 3. Ga ee ee ee

Georgie av ee Se Ao Pe

MiosseltBaye hi .24 a woh ey eee

Swelllen@am) es. a: co a 0

It will be seen that the biggest faunal change is between Durban and Port

St. Johns (27%), not between Port Alfred and Tstisikama, and is high too

between St. Lucia and Durban. This agrees better with the suggestion that the area

between St. Lucia and Durban is a transition zone between the two sub-districts.

As mentioned above, there are many species which, although confined to

the east south of the Limpopo, extend westwards across the Rhodesian plateau

into Zambia and Angola and are, in fact, inhabitants of the South Central

Highlands District as well as of the Eastern Coastal. To what extent, therefore,

are we justified in keeping the two separate?

The avifauna of the Rhodesian plateau, omitting water birds, non-breeding

migrants, montane forest and montane grassland birds and species which just

impinge on it from the South West Arid District, I have calculated (from

Smithers, Irwin & Paterson 1957) to consist of 296 species. The avifauna of the

East African Coastal area from the Save River to Natal I count as 324 species

(based on Pinto 1953, and Clancey 1953). Of these, 236 species occur in both.

This gives a Coefficient of Community of 61,4. This is a little below the figure

usually taken to indicate identity of fauna (Hagmeier & Stults 1964); and in view

of the fact that both Pinto and Clancey cover areas some distance west of the

East African Coastal District, so that some South Central Highlands birds may

have crept into the Coastal list, the distinction between the two districts is

justified. The following key species characterize the East African Coastal District,

those in brackets being extralimital to South Africa:

Francolinus rovumae, Poicephalus cryptoxanthus, (Tauraco fischeri), Halcyon

senegaloides, Pogoniulus pusillus, P. simplex, Campethera notata, (Turdoides

squamulatus), Lioptilus nigricapillus, Phyllastrephus debilis, Andropadus impor-

tunus, Turdus fischeri, Cossypha dichroa, Erythropygia signata, E. quadrivirgata,

Sheppardia gunningi, Bradypterus barratti, B. sylvaticus, Apalis ruddi, (Erythro-

THE ZOOGEOGRAPHY OF THE SOUTH AFRICAN AVIFAUNA 123

cercus holochlorus), Batis fratrum, Erythrocercus livingstonei, (Anthus melindae),

(A. sokokoensis), (Macronyx aurantigula), Tchagra tchagra, Malaconotus quadri-

color, Prionops scopifrons, Lamprotornis corruscus, (Spreo fischeri), (Nectarinia

pembae), N. neergaardi, N. veroxii, Anthreptes reichenowi, (A. neglectus),

Ploceus subaureus, (P. golandi), (Euplectes nigroventris), Hypargos margaritatus

Serinus scotops and S. citrinipectus.

As I have shown elsewhere (Winterbottom 1965), there are two centres of

species concentration in the East African Coastal District, one in Mozambique

and the other opposite Zanzibar and Pemba; though further exploration of the

Be | Aliwal North |

| N j

! \ Ae

F a | | SS HIGHVELD | X

KAROO | SS Elliot

Middelburg ere Umrata

e \ 6

| |

Queenstown ; l; :

a | Cradock | aa 7”

Beaufort West . if

e Aberdeen : Gj

Somerset East) = Fort Beaufort ue

2 e | F

Jansenville | v La

5 s |’ King Williams Town@ 3

East London

ae | Grahamstown Zi

= Seu EAST AFRICAN COASTAL DISTRICT

= ! Le |

x | AIS NS

=! {Knysna = Elizabeth | 34

Pe Ne |

a '=— Mossel Bay’ SS

oF |

Sa lls

e 25° 24° 25° 26° 27° 28° 25° ie reas

Map 6. Lines of Maximal Faunal Change, south-east Cape.

coastal belt in northern Mozambique may show that this bipolarity is due to

imperfect knowledge.

The boundary of the South East Coastal District at its southern extremity

is based on a map (Map 6) showing the range-limits of 234 species by 1° squares.

This somewhat rough-and-ready method simplifies an exceedingly complicated

pattern. The northward bulge in the boundary between 24° and 27°E. is due to

the extension of the distribution of many coastal birds into the lush vegetation ©

of the inland mountains, while Karoo birds infiltrate south in the drier valleys

between the ranges. In the square 32-33°S., 26-27°E. this phenomenon is

particularly marked, for 125 species find their limits in this square.

SOUTH CENTRAL HIGHLANDS DISTRICT

Where my proposals differ most radically from those of previous workers is

in the extent of the South Central Highlands District. Benson & Irwin (1966), in

their fine paper on the Brachystegia avifauna, tentatively suggest that the whole

extent of this woodland might be regarded as forming a zoogeographical unit,

which would fuse Chapin’s Rhodesian Highlands and his East African High-

124 ANNALS OF THE SOUTH AFRICAN MUSEUM

lands Districts (except that part of the latter north of the Equator) and would

add the Rhodesian plateau to the district too. I am in complete agreement with

these suggestions, which, indeed, I had recommended in part over 30 years ago

(Winterbottom 1942), but it will be noted that I have (1) carried the zoogeo-

graphical districts across the Luangwa and Zambezi rifts, where the vegetation

is dominated by the mopane Colophospermum mopane; and (11) taken it further

south to include the middle Limpopo valley and the Transvaal and Swaziland

lowveld, some of which is also dominated by Colophospermum mopane and the

rest by Acacia and other trees.

The differences in the avifauna east and west of the Lebombo mountains

have been commented on by Vincent (1951); and my views on the northern and

eastern Transvaal have already been expressed (Winterbottom 1962). The

question of the mopane avifauna is not discussed in any detail by Benson &

Irwin, to whose paper it would have been irrelevant.

That the avifauna of mopane differs significantly from that of Brachystegia

is, I think, clear and the differences are discussed elsewhere (Winterbottom

1972a, 1972b). However, no such thorough investigation of the mopane avifauna

as that of Benson & Irwin for Brachystegia has ever been made. Benson & Irwin

list 96 Brachystegia species from the Rhodesian area of that woodland, of which

19 do not occur south of the Limpopo. A further 13 species are listed as of

limited distribution in Brachystegia and of these 12 occur south of the Limpopo

and the other is confined to the area north of the Zambezi. There are therefore

altogether 109 species typical of Rhodesian Brachystegia, of which 90 (83%

extend beyond that belt into South Africa. These include 17 of the 29 species

confined to Brachystegia where that occurs. Benson & Irwin exclude raptors

from their list; and these would swell the number of species common to Brachy-

stegia and the area south of the Limpopo and to the mopane woodland further

north to a significant extent.

I believe, therefore, that the inclusion of the mopane areas of the Zambezi

and Luangwa valleys and of the southern lowveld within the South Central

Highlands District is justified.

Excluding forms extra-limital to our area, such as Seicercus laurae, Cisticola

dambo and Myioparus boehmi, species confined to the South Central Highlands,

or virtually so, are as follows:

Centropus cupreicaudus, Coracias spatulata, Stactolaema whytii, Campethera

bennetti, Pinarornis plumosus, Monticola angolensis, Thamnolaea arnoti, Cama-

roptera Stierlingi, Cisticola pipiens, Lanius souzae, Parus rufiventris, Lamprotornis

acuticaudus, Ploceus olivaceiceps and Serinus mennelli.

It will be noted that by no means all of these are birds of the Brachystegia.

WESTERN DISTRICTS

As already indicated, Chapin (1923) placed all that part of South Africa

west of the highveld grasslands and the Rhodesian plateau in a single district,

the South West Arid; Moreau (1952) split off the macchia areas of the extreme

THE ZOOGEOGRAPHY OF THE SOUTH AFRICAN AVIFAUNA 125

south-west; and Winterbottom (1959) extended Chapin’s original district to

include the highveld as well as the macchia. There are some 37 species which are

widespread in this extended South West Arid District and virtually confined to it:

Melierax canorus, Francolinus levaillantoides, Otis kori, O. ludwigii, Eupo-

dotis afra, Cursorius rufus, Pterocles namaqua, Colius colius, Mirafra apiata, M.

sabota, Certhilauda albofasciata, Calandrella starki, C. conirostris, Eremopterix

verticalis, Hirundo spilodera, Anthoscopus minutus, Pycnonotus nigricans, Pari-

soma subcaeruleum, Melaenornis silens, M. infuscatus, Batis pririt, Oenanthe

Table 3

Coefficients of Community: western districts

Winter Damara- Sandveld Kalahari

Rainfall Karoo Highveld land Kalahari Woodland

Winter Rainfall .. = Sl 55 D5 24 34

RANG Obe hes sf co: Sik — 57 46 46 32

rehveld 2896) 5) By — 52 46 46

Damaraland.-.--. . 25 46 52 — 60 60

Sandveld Kalahari . 24 46 46 60 — 51

Kalahari Woodland . 30 32 A Gy a 60 Dil —

monticola, Cercomela schlegelii, Myrmecocichla formicivora, Prinia flavicans,

P. pectoralis, Laniarius atrococcineus, Malaconotus zeylonus, Onychognathus

nabouroup, Lamprotornis nitens, Nectarinia fusca, Sporopipes squamifrons,

Amadina erythrocephala, Uraeginthus granatinus, Serinus flaviventris, Emberiza

impetuani and E. capensis.

Nevertheless, there are differences between one part of this area and another

and Winterbottom (1969) accordingly divided it into six sub-districts: Winter

Rainfall; Karoo; Highveld; Damaraland; Sandveld Kalahari; and Kalahari

Woodland. Later, Winterbottom (1970) suggested that these sub-districts fell

into two groups and proposed to separate the first three as a Southern Temperate

District.

If we analyse the faunas of these various sub-districts, using the Coefficient

of Community and omitting aquatic forms, we find that the group consisting of

the Winter Rainfall, Karoo and Highveld have coefficients of 51-57 inter se; and -

the Damaraland and two Kalahari divisions have coefficients of 51-60 inter se.

Except in the case of Damaraland and the Highveld, between which the coeffi-

cient is 52, the coefficient between divisions in different groups in no case exceeds

46 (Table 3). Inspection of Table 3 also shows that the Winter Rainfall area is the

most distinct (average coefficient 37) and the Highveld least (average coefficient

52,8).

The anomalously high coefficient between Damaraland and the Highveld

is due to the fact that each contains a higher percentage of species of wide range

in the Ethiopian Region further north but which do not penetrate as far south

as the other subdivisions. McLachlan & Liversidge (1970) say of the Sandveld

126 ANNALS OF THE SOUTH AFRICAN MUSEUM

Kalahari ‘perhaps mainly characterised by its lack of birds’; and it is, in fact,

merely an impoverished version of the Damaraland fauna, 95% of its species

being recorded from Damaraland too.

Cooke (1962) produces a map showing the hypothetical vegetation at

50-60 °% of the present rainfall. This shows that the Karoo would be divided into

a northern and a southern section by a broad corridor of desert across Great

Namaqualand, linking the Namib with the Kalahari. I can find no trace of this

in the present bird distribution, and it is odd that, of the list of Namib birds

given by Willoughby & Cade (1967), not a single one is confined to the Namib

and Kalahari; all are either Namib endemics or widely distributed in other parts

of the south-west.

In commenting on his previous suggestions, Chapin (1932) remarked, with

his usual perspicacity, that his treatment of South Africa was the least satis-

factory part of his classification and that more districts might be needed. If we

split the enlarged South West Arid District on the lines proposed above and con-

firmed by the figures in Table 3, we find three genera confined to the restricted

South West Arid and four to the Southern Temperate; with one (Philetairus)

occurring in parts of each. The endemic genera and species of these two districts

ake):

South West Arid District

Genera: Namibornis

Achaetops

Lanioturdus

Doubtful genus: Aethocichla

Species: Francolinus hartlaubi T. gymnogenys

F. adspersus* Monticola brevipes

Pterocles burchelli Melaenornis mariquensis

Poicephalus rueppellii Erythropygia paena

Agapornis roseicollis Namibornis herero

Tockus monteiri Achaetops pycnopygius

T. bradfieldi* Lanioturdus torquatus

Mirafra chuana Laniarius atrococcineus

Turdoides bicolor Lamprotornis australis*

Vidua regia

In addition, the Southwest-—Northeast species Poliohierax semitorquatus,

Turdoides melanops, Bubalornis albirostris, Estrilda erythronotos and Ploceus

rubiginosus are confined to this district in South Africa.

Southern Temperate District

Genera: Geocolaptes Chaetops

Calendula Stenostira

Doubtful genera: Sige/us Euryptila

THE ZOOGEOGRAPHY OF THE SOUTH AFRICAN AVIFAUNA 7

Species: Circus maurus

Francolinus africanus

F. capensis

Bubo capensis*

Eupodotis vigorsii

Merops apiaster

Geocolaptes olivaceus

Certhilauda curvirostris

C. albescens

Calendula magnirostris

Eremopterix australis

Hirundo albigularis

Macronyx capensis

Chaetops frenatus

Parisoma lyardi

Stenostira scita

Cercomela sinuata

Cotracinac

Monticola rupestris

M. explorator

Erythropygia coryphaeus

Cisticola subruficapilla

Prinia maculosa

P. substriata

Tchagra tchagra*

Spreo bicolor

Ploceus capensis*

Serinus tottus

S. alario

S. albogularis

Melaenornis silens*

Species marked * extend somewhat beyond the boundaries of the district in

which they are listed. Not included in the Southern Temperate list are species

confined, or virtually confined, to one sub-district (18 in all). These will be

discussed later. Merops apiaster, of course, also breeds in the Palaearctic and is

widespread in Africa as a non-breeding migrant; and we might have added

Ciconia ciconia to the Southern Temperate list, since it does the same, though its

breeding population in South Africa is very small.

Table 4

Zoogeographical affinities: western species

% of Avifauna

District

Cosmo- S. & E. S.W-N.E. S.W. Sectional

politan Ethiopian African only Endemic Endemic

South West Arid .. 11 37 24 3 16 9

Southern Temperate . 11 33 23 3 16 14

In Table 4, the zoogeographical affinities of the avifauna are set out. Six

categories have been used: Cosmopolitan, being species found in the Ethiopian

and one or more other regions; Ethiopian, being species of wide distribution

within that region; South and East African, being species of wide distribution

in South and East Africa but not extending to West Africa; Southwest—North-

east only, being species found in south-western and north-eastern Africa but not

in the intervening area; South West endemics, which are found in both western

districts; and sectional endemics, found in only one or the other of these two

districts. It will be seen that the Southern Temperate District contains a higher

128 ANNALS OF THE SOUTH AFRICAN MUSEUM

percentage of endemics and a lower percentage of Ethiopian species than the

South West Arid.

Moreau (1966) analyses faunas in five categories: A, aquatic birds; B,

raptorial and scavenging birds; C, game and other ground birds; D, other non-

passerines; and E, passerines. Omitting category A, the results for the two

districts of the south-west area are set out in Table 5. It will be seen that the two

present rather different faunas, the first having more ground birds and passerines

and the second more raptors and ‘other non-passerines’. That the Southern

Temperate should have more ground birds is to be expected in view of the small

extent of woodland in its area, nearly all of which is more or less open country.

Moreau (1966) has drawn attention to the relative decline in group D species as

one proceeds further from the Equator; and the Southern Temperate District is

almost wholly extra-tropical, whereas much of the South West Arid is within the

tropics, so that the difference in Group D is to be expected. Moreau attributes

the decline to the relative paucity of large insects, which are a major food

requirement for many Group D species, in more temperate regions. Many of

these species are also dependent on tree-holes for nest-sites and these, too, are in

shorter supply further from the Equator.

Table 5

Numbers and percentages of species by categories

No. of Species Percentage

District

B C D E Total B C D E

Southern Temperate . . 29 41 44 165 279 10 5 16 59

NSOUGMNVESt Anil i eee mms) 31 58 154 288 16 11 20 53

SomalivAnid” a5 see ee SS 24 68 161 292 13 8 23 55

Sudanese Arid 55 pa 29 74 163 308 14 9 24 53

I have included in Table 5 an analysis on similar lines of the avifaunas of

Chapin’s Somali Arid and Sudanese Arid Districts. These figures show that the

South West Arid avifauna is intermediate between these wholly tropical faunas

and that of the Southern Temperate District in respect of group D. All three of

the more tropical faunas have fewer passerines and ground birds but more

raptors than the southern fauna. The relationships between these arid faunas are

more fully discussed elsewhere (Winterbottom 1972c).

Moreau (1966) uses the same classification as that discussed above to

analyse the faunas of ‘sample-areas’ in certain vegetation types. He gives one

example for the South West Arid District. This relates to Acacia woodland in

the central Kalahari and, as may be seen from Table 6, represents a decidedly

impoverished fauna when compared with Damaraland areas further west, even

with Valencia, on the borders of the Namib (Kemp & Kemp 1972); or with the

THE ZOOGEOGRAPHY OF THE SOUTH AFRICAN AVIFAUNA 129

Kalahari Gemsbok National Park further south (Prozesky & Haagner 1962), but

not with the miserable fauna recorded from Tsabong (Pianka & Huey 1971).

Moreau explains that the poor fauna of his Kalahari sample is partly because it

Table 6

Numbers of species in sample areas, South West Arid District

Locality Latitude Longitude CG D E Total

@entral Kalahari... . .. ©. 24°S. (Os Pa Be 2 16 43 71

Smut 2 2 , |. 21°38'S, 1835178, 14 28 58 100

SISSckaMemeees hs, se 19°22’S, gpa eh 16 38 65 119

Oiwasandue ss. . . . 19°12'S. 14°28’E. 14 22 61 97

GemsbokiPatk. .. . .- 26-27°S. 20-21°E. DS 20 66 11

Saienci@weeeus . ., » 23 LO’S. 162572: 9 25 46 78

Sapo een: « . = 6. 2608'S. PLL IRSA Ele 6 Z 19 32

contains only the birds present in the dry season and therefore omits all breeding

migrants. However, this cannot account for Pianka & Huey’s results.

Similar lists for 12 Karoo localities in the Southern Temperate District have

an average for groups C-E species of 45,6, with a range of 32-61. They are

markedly inferior to the South West areas in all groups. Six Winter Rainfall

localities have an average for the same groups of 48,3, with a range of 38-59,

suggesting a similar type of avifauna to that of the Karoo. A single sample from

the Highveld (based on Boddam-Whetham 1965) has 88 species in groups C—E,

including 22 in group D; and is thus intermediate between the Karoo areas and

those of the South West Arid. Thus the Southern Temperate District in general

contains rather fewer species as a whole than the South West Arid (Table 5) and

decidedly fewer in any one locality.

The inter-relations of the various sub-districts of the South West Arid and

Southern Temperate Districts have been discussed above and also elsewhere

(Winterbottom 19685, 1971a). However, some comments may be made on the

Southern Temperate sub-districts and particularly on the Highveld. x

Winter Rainfall Area

There are six indigenous species in the Winter Rainfall area: Anthus

crenatus, Pycnonotus capensis, Bradypterus victorini, Promerops cafer, Nectarinia~

violacea and Serinus leucopterus. It is, as remarked above, the most distinct of

the three Southern Temperate sub-districts and only 13 of the widespread south-

western birds (p. 125) but 25 of the Southern Temperate endemics occur as

breeding species. That this avifauna is primarily derived from areas further north

and east, especially the Karoo and the South Eastern Coastal District, is shown

by the fact that an analysis of the terrestrial birds in Moreau’s groups C, D and

E gives the proportions of one indigenous species to five found also in other

South Temperate divisions to 11 widespread in South Africa. In view of the

marked individuality of the lower vertebrates and of the plants of this area, some

explanation would seem to be needed; and I have suggested (Winterbottom

130 ANNALS OF THE SOUTH AFRICAN MUSEUM

1968c) that the most important factors may ‘have been the same ones, whatever

they are, which have been responsible for the reduction in size of the avifauna

as one proceeds from low latitudes to high; with the relatively lesser ecological

diversity of the vegetation type as a contributing factor’.

Karoo

Five species are endemic to the Karoo; Alaemon grayi, A. burra, Calandrella

sclateri, Camaroptera subcinnamomea and Eremomela gregalis. Of these, A. grayi

is confined to the Namib. No less than 35 of the 37 south-western endemics

occur widely on the Karoo; and 26 of the Southern Temperate endemics.

Highveld

The highveld grasslands of South Africa have their closest, and virtually

their only, counterpart in the rest of the Ethiopian Region in the high grasslands

of Ethiopia, from which they are cut off by some 2 500 miles (4000 km) of

intervening tropical and subtropical vegetation. Nevertheless, out of 143

terrestrial species listed by Urban & Brown (1971) for the highland grasslands of

Ethiopia, 77 (54%) are recorded for the Orange Free State (Van der Plaat 1961)

and/or Lesotho (Jacot-Guillarmod 1963).

In considering the avifauna of the highveld and its affinities, two recent

man-made changes must be taken into consideration. The first, in point of

beginning, is the progressive degradation of the grassveld and its replacement by

Karoo as a consequence of 300 years of over-grazing and other agricultural

malpractices. Acocks (1953), our chief authority on such matters, believes that

the Karoo has advanced at least 2° eastward in the last 450 years (Map 7). The

original westward boundary of the grassveld coincides with a climatic change

from temperate to tropical. It is necessary to bear this in mind since a number of

typical highveld birds, such as Eupodotis caerulescens, Anthropoides paradisea,

Hirundo spilodera and the nominate race of Certhilauda albofasciata, still

inhabit this former grassveld area. This is in significant contrast to the Karoo

enclave in the Breede River valley, which is natural Karoo and where, for

instance, the subspecies of Serinus flaviventris and Certhilauda curvirostris are

the Karoo forms quintoni and gilli respectively and not the subspecies of the

surrounding macchia, flaviventris and curvirostris.

The second change has been the considerable planting of trees, mostly

aliens, round houses and as wind-breaks, which has led to the invasion of, or at

least increase in numbers in, this originally treeless area by arboreal species

previously rare or absent (see, e.g., Freer 1965).

Both these changes affect attempts to assess the composition of the highveld

avifauna, the first by suggesting that some highveld endemics or near-endemics

range to a considerable distance beyond the grassveld; and the second by sug-

gesting that a number of woodland species inhabit the highveld too.

A full list of the characteristic birds of the grasslands will be found in

Winterbottom (1972a) but for zoogeographical purposes, there are six species

confined, or almost confined, to this division. They are:

THE ZOOGEOGRAPHY OF THE SOUTH AFRICAN AVIFAUNA 131

Lourenco

Marques

Johannesburg

)

RoI

~—.

Probable boundary in1400 A.D

aie -Western boundary in1950

Durban

Port Elizabeth — (After Acocks,1953) |

Map 7. Highveld Grassland.

Geronticus calyus, Eupodotis caerulescens, Calandrella fringillaris, Mirafra

ruddi, Anthus chloris and Promerops gurneyi.

The last of these is not an inhabitant of grassland but of Protea bush and it

reappears in the Protea bush of the eastern highlands of Rhodesia. It is obviously

closely related to P. cafer of the Winter Rainfall area, with which it occasionally

hybridises in the King Williams’ Town area, where the two species are sympatric

(Skead 1964).

TRANSITIONAL ZONE

This comprises a mosaic of Baikiaea, Acacia and mopane woodland, open

grassland, flood plains and swamps. Examination of the ranges of individual

132 ANNALS OF THE SOUTH AFRICAN MUSEUM

species show that many characteristic birds of the South Central Highlands

extend into this zone, where they meet other forms characteristic of the South

West Arid. This is, for instance, the area of overlap between Pycnonotus barbatus

and P. nigricans; Melierax metabates and M. canorus; Tockus alboterminatus and

T. bradfieldi; Mirafra rufocinnamomea and M. apiata; Turdoides jardeneii and

T. melanops; Prinia subflava and P. flavicans; Laniarius aethiopicus and L. bicolor:

though some of these species pairs are ecologically separated. It marks the

southern limit in the west of such species as Streptopelia semitorquata, Poicepha-

lus robustus, Macrodipteryx vexillarius, Merops pusillus, Andropadus flaviventris

and Monticola angolensis; and the northern limit of, for example, Laniarius

atrococcineus, Lamprotornis australis, Bubalornis albirostris, Pterocles namaqua

and P. burchelli, The mixed nature of its avifauna is apparent from these

examples. Smithers (1964) divides the Botswana part into several areas, but the

Coefficients of Community between these range from 60 to 75, indicating

identity of fauna.

MONTANE

Chapin (1932) pointed out that the montane areas of the Ethiopian Region,

scattered and fragmented as they are, formed a zoogeographical unit with a

distribution pattern quite distinct from that of the lower country. He accordingly

grouped them together as a Humid Montane Province, dividing it into two

districts, a Cameroon Montane District and an East African Montane District.

He did not, however, include any montane areas south of the Zambezi in this

province.

Moreau (1966) remarks of the montane forests south of the Zambezi that

they contain ‘less than half [the number of species] . . . occupying the montane

forests of Malawi’ and dismisses them from his discussion in two brief para-

graphs. Closer investigation, however, suggests that Moreau was exaggerating.

I compiled from Benson (1953) a list of 91 species inhabiting montane forests in

Malawi; of this list, 68 species (75°%) are recorded from forest in Rhodesia

and/or southern Mozambique and 58 (64°) from south of the Limpopo. One or

two species (e.g. Pogonocichla swynnertoni for Rhodesia and Cossypha dichroa

for South Africa) must be added to obtain the total avifaunas of these southern

forests but it is clear that at least the Rhodesian montane forests and that of

Gorongoza must be included in the same zoogeographical division as the

montane forests further north. Dowsett (1971) agrees.

As one proceeds south, the situation is complicated by the fact that the

forests descend to lower and lower altitudes and finally merge with the lowland

forest of the East African Coastal District to form a Southern Temperate

Forest (Liversidge 19595; Winterbottom 1968). It is still further complicated by

differences in the ecological preferences of species in tropical and temperate

areas. The Limpopo and Zambezi valleys must always have isolated these

southern montane forests into two blocks, cut off by the Zambezi from the

montane areas to the north. Whether one is justified in splitting off the montane

THE ZOOGEOGRAPHY OF THE SOUTH AFRICAN AVIFAUNA 133

and temperate forests south of the Limpopo as a distinct South Temperate

Forest District or whether it is better to regard them as a sub-district of the East

African Montane District is a moot point, but as they include a number of

endemic species, such as Cossypha dichroa, Bradypterus sylvaticus and Erythro-

pygia signata, besides lacking many forms widespread further north (e.g. Alethe

spp., Arizelocichla spp., Cryptospiza spp.), they cannot be ‘lumped’ with the

East African Montane forests without some distinction being drawn.

In an endeavour to find out what might have become of our temperate and

montane forests in South Africa during the Pleistocene, I first tried to ascertain

the factors controlling their natural distribution at present (i.e. ignoring the

effects of human interference). Acocks (1953) suggests that the minimum rainfall

for forest is 30-35 inches (c. 762-889 mm) per annum. But it is clear, from the

absence of forest at high altitudes on the Drakensberg and the mountains of the

south-west Cape, that some other factor must be involved as well. The most

probable of these is temperature.

Zeuner (1959) states that the minimum for tree growth is a temperature in

excess of 10°C. in summer; but this relates to the northern taiga and not to the

temperate broad-leafed forest. In fact, the only place in South Africa where this

condition might apply if the temperature fell by 5°C. is Port Nolloth and the

rainfall there is inadequate for tree growth anyway.

I therefore plotted the rainfall against the average minimum monthly

temperature of the coldest month for those meteorological stations in the vicinity

of which forest occurs or was known to occur in the recent past. The bulk of these

fell into a zone demarcated by the 850 mm isohyet and the 8°C. isotherm; but

in the extreme south, forest could occur with 700 mm of rain and a minimum

average temperature of 7°C. in the coldest month. In South America, Nothofagus

forest occurs at much lower temperatures than this but Nothofagus apparently

never occurred in Africa.

I then made the assumptions that during a hypothermal period, the

temperature would drop 5°C. and the effectiveness of the rainfall would increase

by 25%. Cooke (1962) has produced a map showing the hypothetical vegetation

with a rainfall of 150% of that at present, but this seemed to me excessive.

The next step was to plot those meteorological stations which met these two

requirements —i.e. those with a minimum monthly average temperature of 13°C. —

or more in the coldest month and a rainfall of at least 640 mm.

The map so obtained (Map 8) suggests that, except close to sea-level, forest

could not grow along the south coast except between Knysna and Humansdorp;

that there would be a block of forest along the coastal lowland from Alexandria

to St. Lucia and thence west of the Lebombo Mountains to about 24°30S.,

though the forest may not have been completely continuous. There would be

further montane forest blocks in the north-eastern Transvaal, major enlarge-

ments of the existing Woodbush and Soutpansberg forests but lower down the

mountains, though not descending into the Limpopo valley. It also appears that

the Kasane area, at the junction of the Zambezi and Chobe, would have been

134 ANNALS OF THE SOUTH AFRICAN MUSEUM

Way

Kyi

“‘\

19 21 23 25 27 29 31 33

Map 8. Suggested forest country during a pluvial period.

able to support forest—there is, of course, riverine (‘gallery’) forest there today.

This forest would not reach as far east as the Victoria Falls; and as its postulated

existence depends on the assumption that the increased effectiveness of the

rainfall would be the same there as in the south, the suggestion is decidedly

speculative.

The big coastal block referred to above is mapped by Cooke (1962) as

‘Tropical forest-savanna mosaic’ and he extends it up into coastal Mozambique

as far as Beira. I did not carry my own investigations into Mozambique but the

rainfall map of the South African Weather Bureau supports Cooke’s idea that

the conditions along the coast there would be much the same as in Natal and

Zululand and this has been adopted in my map.

The southern (Knysna) forest block of my map is shown by Cooke as

‘Subtropical mixed woodland’, which seems to me unlikely.

My map suggests that the temperate (montane) forest and the tropical low-

land forest would merge along the Pongola River about latitude 27°S., which is

some 4—5° further north than they do today. In East Africa, including Rhodesia,

a faunal change in continuous forest cover occurs rather abruptly at an altitude

which naturally varies according to the latitude; but this change does not

coincide with any change in the botanical nature of the forest. Further south,

where lowland and montane forest are separated by drier areas, the avifaunas of

the two differ somewhat but chiefly at the subspecific level. Further south still,

the two forest types merge into a temperate lowland forest and the avifauna

THE ZOOGEOGRAPHY OF THE SOUTH AFRICAN AVIFAUNA 15

contains elements of both, plus some endemics. It would presumably be forest of

this type which extended north to Zululand in the Pleistocene hypothermals and

its greater extent and fragmentation during arid periods would give more scope

for the development of these endemics.

I have not considered here whether the lowering of the sea level, and the

consequent exposure of much of the present Agulhas Bank, would affect the

issue. This has been discussed briefly elsewhere (Winterbottom 1968) and my

conclusion was that it would be unlikely to be important.

The avifauna of the montane grasslands and heath-Protea complex in

Rhodesia includes one endemic species, Prinia robertsi; a number of species of

wider distribution further south but confined to this area in Rhodesia, such as

Sarothrura affinis, Sphenoeacus afer, Malaconotus zeylonus, Promerops gurneyi,

Nectarinia famosa, and Serinus canicollis; and montane species of East Africa,

here at the southern end of their range, like Apus myoptilus and Nectarinia

kilimensis. Of a list of 58 non-forest montane species in Malawi (based on

Benson 1953), ten (17%) do not occur south of the Zambezi.

In the montane non-forest areas south of the Limpopo, there are a number

of other species restricted to this type of habitat there but occurring down to

sea-level.further south. Examples are Geocolaptes olivaceus and Serinus tottus.

Although the montane forests north of the Limpopo are regarded as a

distinct district, this is not the case with the lowland forests of East Africa. These

differ from the West African lowland forests in their broken and slighter extent,

so that many of the species extend out of the forest into dense scrub. I have

listed only six species (Neocossyphus poensis, Bias musicus, Malacocincla rufi-

pennis, Dicrurus ludwigii, Nectarinia olivacea and Mandingoa nitidula) which

occur in the West African forests and again in the coastal forests of the east but

not in between, and two cases in which one member of a superspecies inhabits

the West African forests and another the East African (Macrosphenus concolor

and M. kretschmeri; and Erythrocercus mccalli and E. holochlorus), so that the

avifaunas of the two are quite distinct and it is simplest to regard the East

African coastal forests as a habitat of the East African Coastal District. This

difference between montane and lowland forest becomes intelligible if we

realize that when forest was at its maximum, during a pluvial, the lower tempera-

ture would favour montane, rather than lowland, forest; that the avifaunas of”

the two forest types are largely distinct (Moreau (1952) estimates that 90 % of the

montane forest avifauna is confined to that habitat and does not occur in low-

land forest); and that therefore there is unlikely to have been any direct con-

nection between the lowland forest of east and west.

FRESHWATER

As I have shown elsewhere (Winterbottom 1967), the freshwater avifauna

of eastern and northern South Africa forms part of the East African tropical

aquatic avifauna; but that of the west and south-west is rather different.

The tropical avifauna, which extends south to St. Lucia, has been fully

136 ANNALS OF THE SOUTH AFRICAN MUSEUM

discussed and described elsewhere (Winterbottom 1967, 1972a). It comprises 52

species, excluding Palaearctic migrants and species characteristic of reed-beds

(e.g. Acrocephalus spp.). A further ten species are sufficiently widespread to

suggest that they, too, should be included. The fauna, which has already been

set out in papers cited above and also in Ruwet (1965), is:

Phalacrocorax lucidus, P. africanus, Anhinga rufa, Pelecanus rufescens,

Nycticorax nycticorax, Butorides striatus, Ardeola rufiventris, A. ralloides, A. ibis,

Egretta ardesiaca, E. garzetta, E. intermedia, E. alba, Ardea cinerea, A. melano-

cephala, A. goliath, A. purpurea, Ixobrychus minutus, Scopus umbretta, Ibis ibis,

Leptoptilos crumeniferus, Ephippiorhynchus senegalensis, Anastomus lamelligerus,

Threskiornis aethiopica, Plegadis falcinellus, Bostrychia hagedash, Platalea alba,

Thalassornis leuconotus, Sarkidiornis melanotos, Dendrocygna viduata, D. bicolor,

Nettapus auritus, Plectropterus gambensis, Anas erythrorhyncha, Rallus caeru-

lescens, Limnocorax flavirostra, Gallinula chloropus, Porphyrio porphyrio, Balae-

arica pavonina, Actophilornis africanus, Vanellus senegallus, V. armatus, V. crassi-

rostris, Himantopus himantopus, Gallinago nigripennis, Larus cirrocephalus,

Glareola pratincola, Chlidonias leucoptera, Alcedo cristata, Ceryle rudis,

C. maxima.

For southern African areas, we should add Pelecanus onocrotalus.

The temperate aquatic avifauna of the south-west, based on species occur-

ring in at least 5% of 1 210 lists, consists of 49 species, as follows:

Podiceps cristatus, P. nigricollis, P. ruficollis, Pelecanus onocrotalus, Phala-

crocorax lucidus, P. africanus, Anhinga rufa, Ardea cinerea, A. melanocephaal,

A. purpurea, Egretta garzetta, E. intermedia, Ardeola ibis, Nycticorax nycticorax,

Scopus umbretta, Threskiornis aethiopica, Platalea alba, Phoenicopterus ruber,

P. minor, Plecotropterus gambensis, Alopochen aegyptiacus, Tadorna cana, Anas

smithii, A. undulata. A. erythrorhyncha, A. capensis, Netta erythrophthalma,

Oxyura punctata, Haliaeetus vocifer, Circus ranivorus, Limnocorax flavirostra,

Porphyrio porhpyrio, Gallinula chloropus, Fulica cristata, Charadrius marginatus,

C. pecuarius, C. tricollaris, Vanellus armatus, Gallinago nigripennis, Recurvi-

rostra avosetta, Himantopus himantopus, Burhinus vermiculatus, Ceryle rudis,

Alcedo cristata, Riparia paludicola, Acrocephalus gracilirostris, Bradypterus

babaecalus, Cisticola tinniens and Motacilla capensis.

Non-breeding migrants from the Palaearctic, visitors from the sea-coast

(e.g. Larus novaehollandiae) and species which forage on the shores but breed

elsewhere (e.g. Macronyx capensis) are excluded.

Of the above 49 species, only 22 (45°) occur also in the tropical list and the

Coefficient of Community is 25; so that the two are very distinct.

I have also suggested that the Karoo-Damaraland water birds may form a

distinct assembly. The fauna, based on South West African and Karoo lists,

numbers 28 species, namely:

Podiceps ruficollis, Phalacrocorax lucidus, P. africanus, Ardea cinerea,

A. melanocephala, Scopus umbretta, Ciconia nigra, Ibis ibis, Platalea alba,

Alopochen aegyptiacus, Tadorna cana, Anas smithii, A. undulata, A. sparsa,

THE ZOOGEOGRAPHY OF THE SOUTH AFRICAN AVIFAUNA 7

A. capensis, A. erythrorhyncha, Oxyura punctata, Gallinula chloropus, Fulica

cristata, Charadrius pecuarius, C. tricollaris, Vanellus armatus, Recurvirostra

avosetta, Himantopus himantopus, Larus cirrocephalus, Acrocephalus baeticatus,

Cisticola tinniens and Motacilla capensis.

The Coefficient of Community between this fauna and that of the south-west

is only 43; but Simpson’s Coefficient:

= <P 100

where c is the number of species in common and n, is the number in the smaller

fauna, is 82, suggesting that this fauna may be little more than an impoverished

version of that of the south-west. It should be noted, however, that Anas

undulata, despite the map in McLachlan & Liversidge (1970), and Cisticola

tinniens occur only in the south and not in South West Africa and the validity of

their inclusion is thus rather doubtful. Their exclusion would reduce Simpson’s

Coefficient to 75. With the tropical fauna, Simpson’s Coefficient is only 43. The

distribution pattern of most aquatic birds in South West Africa is a narrow strip

down the centre, between the Namib and the Kalahari, and only extending as

far south as about 25°S., though some of this may be due to lack of observations

at the time water is present in the south. On the whole, I think it would be better

to keep this fauna distinct as a South West Arid aquatic avifauna.

COASTAL

The birds of the South African coasts show an east-west difference in their

distribution, very largely correlated with the cold current on the west coast and

the warm one on the east. The rich avifauna of the west coast is characterized by

the presence as breeding species of Spheniscus demersus, Sula bassana, Phalacro-

corax lucidus, P. capensis, P. neglectus, P. africanus coronatus, Haematopus

moquini, Charadrius marginatus, Larus novaehollandiae, L. dominicanus, Sterna

bergii and S. balaenarum. The section from Cape L’Agulhas to Algoa Bay may

be regarded as a transition zone, east and north of which the coastal fauna is an

impoverished one, with, from Zululand north, a small tropical component

Dromas ardeola, Sterna fuscata and S. benghalensis being the most important;

but none of these breeds within our limits. The only coastal-breeding species

given by Clancey (1971) for southern Mozambique are Charadrius marginatus,

Sterna bergii and Hydroprogne caspia.

OCEANIC

As Liversidge (1959a) has pointed out, oceanic waters in South Africa

belong to the Southern Oceans of Alexander (1955). The Southern Oceans are

characterized by the wealth of Sphenisciformes and Procellariiformes, plus a

few species of other groups, such as Sterna vittata. The only uncertain point is

just where this Southern Oceans avifauna gives place to that of the Tropical

Oceans. Serventy, Serventy & Warham (1972) have placed the limit of the

138 ANNALS OF THE SOUTH AFRICAN MUSEUM

13° 17 ar 25° 2g" 33

[ I" i Luangwa River ‘ambezi River

/ ( ALA |

gene Py =.

oy a ae

OLAND Victoria Falls

<Otjivasandu @oissekab NGAMILAND

ar Zz eoturmfeld

a KALAHARI

Valencia

hee See i OES ho ae ee

io Kalahari

a mM em ok

7S) bon 4

©, GREAT ey v

NAMAQUALAND i

zal

4 Or Nnge RNY }

| Dates Port Nollot /

Fort StUohns

[ King Williams Town

1 ast London——

33 George Humanisdor, rt Alfred

Cape Peninsu a “e, ellendam e. e A\ Ba

os ne Mossel Bay cima

L _\ Cape Agulhas e

13 ii ae + 29 33 37

Map 9. Places mentioned in the text.

Sub-Antarctic avifauna at the Sub-tropical Convergence; but have, in effect,

admitted the invalidity of this by extending the boundary northwards to include

the southern coasts of Australia. The idea that the Sub-tropical Convergence

constitutes a faunal boundary for birds no doubt stems from Murphy (1936),

who considered that ‘the majority of oceanic birds are bound as peons to their

own specific types of surface water’. This is an exaggeration (Winterbottom

19715). In the Atlantic, the boundary between the Southern Oceans and the

Tropical Seas appears to be about the Tropic of Capricorn; on the east coast, it

may be about the same latitude—of the 13 species recorded by Rand (1962) ina

transect along 28°S., all but Sterna fuscata were birds of the Southern Oceans

fauna; but the breeding sea birds of Europa Island, in about 22°S., are tropical—

Sula sula, Fregata minor, Phaethon spp. and Sterna fuscata (Malzy 1966). The

distribution of marine birds in the Indian Ocean is poorly known (Watson,

Zusi & Storer 1963) but of the 18 tropical species, ten have been recorded within

South African limits, mostly as stragglers.

THE ZOOGEOGRAPHY OF THE SOUTH AFRICAN AVIFAUNA 139

Of the predominantly marine families, the Spheniscidae, Diomedeidae,

Procellariidae, Hydrobatidae, Sulidae, Phalacrocoracidae, Stercorariidae and

Laridae, South African seas are inhabited by 54 species, excluding such fresh-

water forms as Larus cirrocephalus and Chlidonias spp. in the Laridae and the

tropical terns, boobies, tropic-birds and frigate-birds which reach the area only

marginally. Of these, 11 species breed (and are more typical of the coasts than

the ocean) and another 11 are migrants from the north. The remaining 32 species

all breed in the south, ten of them (plus Larus dominicanus, which also breeds in

South Africa) south of 60°S. Attempts to divide the avifauna of the south into

Antarctic and Sub-Antarctic have been made by Murphy (1964) and Voous

(1965) but there are only three marine birds whose breeding is wholly confined to

the Antarctic (Thalassoica antarctica, Fulmarus glacialoides and Pagodroma

nivea) and the first two of these range far to the north out of the breeding season.

It may therefore be doubted if it is really profitable to make such a distinction.

As I have pointed out elsewhere (Winterbottom 19715), the influence of surface

waters has been over-emphasized (except possibly in the case of penguins, whose

distribution when not breeding 1s very poorly known) and I agree with Alexander

in including all waters south of the Tropical Convergence in a single zoo-

geographical district, the Southern Oceans. To this district, South African off-

shore waters belong. It should be noted, however, that not a single species of the

Procellariiformes breeds in South African waters.

SUMMARY

South Africa forms part of the Ethiopian Zoogeographical Region. Its

birds exhibit five distinct patterns of geographical distribution: oceanic birds,

coastal birds, freshwater birds, forest birds and the rest.

Almost the whole of oceanic waters off the South African coast are inhabited

by birds of the Southern Oceans Zoogeographical Region.

Coastal birds show a difference between east and west correlated with the

warm and cold currents; the eastern avifauna is impoverished and, in the north,

has a small tropical element.

Freshwater birds fall into three groups, an eastern tropical, a southern

temperate and a western.

The avifauna of the mountain forests of Rhodesia and Mozambique forms

a sub-district of the East African Montane District of Chapin (1932). The avi-

fauna of the temperate forests south of the Limpopo is rather more distinct; but

the tropical forests of the lowlands of Mozambique and Zululand are best

regarded as a habitat of the East African Coastal District.

For the remaining land birds, the pattern suggested by their distribution

comprises four districts, three of which extend beyond the borders of South

Africa, and a transitional zone. Sub-districts are recognizable for three of these

districts.

The districts and sub-districts are:

140 THE ZOOGEOGRAPHY OF THE SOUTH AFRICAN AVIFAUNA

South Central Highlands District

South West Arid District

Damaraland Sub-district

Kalahari Sandveld Sub-district

Kalahari Thornveld Sub-district

East African Coastal District

Mozambique Sub-district

Southern Sub-district

South Temperate District

Karoo Sub-district

Highveld Sub-district

Winter Rainfall Sub-district

The transition zone lies between the South Central Highlands and the

South West Arid Districts.

ACKNOWLEDGEMENTS

In matters of past plant distribution, I have had the advantage of conversa-

tions with Professor E. M. van Zinderen Bakker and Professor E. Schelpe, who

must not, however, be held responsible for the views expressed here. I have also

had discussions and correspondence on zoogeography with Dr J. C. Poynton,

whose views I have found stimulating—especially when I disagreed with them!

My colleagues, Professor W. R. Siegfried and Mrs M. K. Rowan, read the whole

MS and made helpful suggestions.

For anything concerning the distribution of passerine birds I have found

Hall & Moreau (1970) invaluable— without it, this paper would have contained

many more inaccuracies than no doubt it does. For non-passerines, I have used

White (1965), also a useful work of reference. For the detailed distribution of

birds within South Africa, I have used the various numbers of the South African

Avifauna Series, published by the Percy FitzPatrick Institute of African Ornitho-

logy, and the numerous MS lists filed in the Institute, especially my own and

those of C. J. Skead, whose meticulous work on the birds of the eastern Cape

Province has been invaluable.

The section on the western districts incorporates most of a paper presented

at the XV Congressus Internationalis Ornithologicus, where my views on the

desirability of two districts in this area were first put forward (Winterbottom

1970).

REFERENCES

Acocks, J. P. H. 1953. Veld types of South Africa. Mem. bot. Surv. S. Afr. 28: i-iv, 1-192.

ALEXANDER, W. B. 1955. Birds of the ocean. 2nd ed. London: Putnam.

BENSON, C. W. 1953. A checklist of the birds of Nyasaland. Blantyre, Lusaka: Nyasaland

Society.

BENSON, C. W. & Irwin, M. P. S. 1966. The Brachystegia avifauna. Ostrich, Suppl. 6: 297-321.

THE ZOOGEOGRAPHY OF THE SOUTH AFRICAN AVIFAUNA 14]

BoDDAM-WHETHAM, A. D. 1965. The birds of Kirklington, O.F.S. S. Afr. Avifauna Ser.

P. Fitzpatrick Inst. Afr. Orn. 32: 1-35.

Bowen, W. W. 1933. African bird distribution in relation to temperature and rainfall. Ecology

14: 247-271.

BRoDKOoRB, P. 1960. How many birds have existed? Bull. Fla St. Mus. biol. Sci. 5: 41-53.

CHAPIN, J. P. 1923. Ecological Aspects of bird distribution in tropical Africa. Am. Nat. 59:

106-125.

CHAPIN, J. P. 1932. Birds of the Belgian Congo. Part 1. Bull. Am. Mus. nat. Hist. 65: i-x, 1-756.

CLANCEY, P. A. 1953. A preliminary list of the birds of Natal and Zululand. Durban: Durban

Museum.

CLANCEY, P. A. 1971. A handlist of the birds of Mocambique. Part 1. Mems Inst. Invest. cient.

Mocamb. 10(A): 145-302.

CLARK, J. D. 1967. Atlas of African prehistory. Chicago, London: University of Chicago Press.

CLOUDESLEY-THOMPSON, J. L. 1969. The zoology of tropical Africa. London: Weidenfeld &

Nicolson.

Cooke, H. B. S. 1962. The Pleistocene environment in southern Africa: hypothetical vegetation

in southern Africa during the Pleistocene. Ann. Cape prov. Mus. 2: 11-15.

De BeEAuForT, L. F. 1951. Zoogeography of the land and inland waters. London: Sidgwick

Jackson.

DowseTT, R. J. 1971. The avifauna of the Makutu Plateau, Zambia. Revue Zool. Bot. afr. 84:

312-333.

EKMAN, S. 1952. Zoogeography of the sea. London: Sidgwick.

FREER, J. S. 1955. Reaction of birds to habitation. Bokmakierie 7: 48-49.

Hacmeier, E. M. & Stutts, C. D. 1964. A numerical analysis of the distribution patterns of

North American mammals. Syst. Zool. 13: 125-155.

HALL, B. P. 1970. Causal ornithogeography of Africa, Int. orn. Congr. 15 (Abstr.): 33-36.

HALL, B. P. & Moreau, R. E. 1970. An atlas of speciation in African passerine birds. London:

British Museum (Natural History).

JACOT-GUILLARMOD, C. 1963. Catalogue of the birds of Basutoland. S. Afr. Avifauna Ser.

P. Fitzpatrick Inst. Afr. Orn. 8: 1-111.

JENSEN, R. A. C. & JENSEN, M. K. 1971. First breeding records of the Herero Chat Namibornis

herero, and taxonomic implications. Ostrich, Suppl. 8: 105-116.

KEAST, A. 1972. Faunal elements and evolutionary patterns: some comparisons between the

continental avifaunas of Africa, South America and Australia. Int. orn. Congr. 15 (Proc.):

594-622.

Kemp, A. C. & Kemp, M. I. 1972. The birds of Valencia Ranch, South West Africa. S. Afr.

Avifauna Ser. P. Fitzpatrick Inst. Afr. Orn. 82: 1-15.

Lawson, W. J. 1971. Check list of the birds of Durban. S. Afr. Avifauna Ser. P. Fitzpatrick

inst. Aj. Orn. 73: 1-80.

LiversiIDGE, R. 1959a. The place of South Africa in the distribution and migration of ocean

birds. Ostrich, Suppl. 3: 47-57.

LiIveRSIDGE, R. 19596. Tropical mountain birds south of the Zambesi. Ostrich, Suppl. 3: 68-78.

McCuLLocnu, D., SKEAD, C. J. & WINTERBOTTOM, J. M. 1970. The birds of Port St. Johns.

S. Afr. Avifauna Ser. P. Fitzpatrick Inst. Afr. Orn. 71: 1-34.

MACKWORTH-PRAED, C. W. & GRANT, C. H. B. 1955. African handbook of birds. Ser. 1, 1.

Birds of eastern and north eastern Africa. London: Longmans, Green.

McLAcHLAN, G. R. & LiversipGe, R. 1947. Roberts Birds of South Africa. Johannesburg:

Central News Agency.

McLAcHLAN, G. R. & LiverRsIDGE, R. 1970. Roberts Birds of South Africa. 3rd ed. Johannes-

burg: Central News Agency.

Maizy, P. 1966. Oiseaux et mammiféres de I’Ile Europa. Mém. Mus. natn. Hist. nat., Paris

(nse A) 4s 23-27.

Moreau, R. E. 1952. Africa since the Mesozoic. Proc. zool. Soc. Lond. 121: 869-913.

Moreau, R. E. 1966. The bird faunas of Africa and its islands. New York, London: Academic

Press.

Murpny, R. C. 1936. Oceanic birds of South America. New York: American Museum of

Natural History.

Mourpnuy, R. C. 1964. Systematics and distribution of Antarctic petrels. In: CARRICK, R.,

HoLpGate, M. & Prevost, J., eds. Biologie Antarctique: 349-358. Paris: Hermann.

142 ANNALS OF THE SOUTH AFRICAN MUSEUM

PATERSON, J. M. 1958. Check list of birds of the eastern Cape Province. Port Elizabeth:

Eastern Cape Wild Bird Society.

PIANKA, E. R. & Huey, R. B. 1971. Bird species density in the Kalahari and the Australian

deserts. Koedoe 14: 123-129.

Pinto, A. A. DA R. 1953. Uma lista sistematica das aves da regiao do extremo sul da Provincia

de Mocambique. Bolm Soc. Est. Mogamb. 23 (77): 1-72.

Poo.ey, A. C. & Dixon, J. E. W. 1966. A check list of the birds occurring in the Ndumu Game

Reserve in Northern Zululand. S. Afr. Avifauna Ser. P. Fitzpatrick Inst. Afr. Orn. 39: 1-38.

PoynTOoNn, J. C. 1964. The biotic divisions of southern Africa as shown by the Amphibia. Jn:

Davis, D. H. S., ed. Ecological studies in southern Africa: 206-218. The Hague: Junk.

(Monographiae biol. 14.)

PROZESKY, O. P. M. & HAAGNER, C. H. 1962. A check list of the birds of the Kalahari Gemsbok

Park. Koedoe 5: 171-182.

RAND, R. W. 1962. Sea-birds south of Madagascar. Ostrich 33 (3): 48-51.

Roserts, A. 1940. The birds of South Africa. London: Witherby; Johannesburg: Central News

Agency.

Ruwet, J. C. 1965. Les oiseaux des plaines et du lac-barrage de la Lufira supérieure (Katanga

meridional). Liege: Editions F.U.L.R.E.A.C.

ScLATER, P. L. 1858. On the general geographical distribution of the members of the class

Aves. J. Linn. Soc. (Zool.) 2: 130-145.

SERVENTY, D. L., SERVENTY, V. & WARHAM, J. 1972. Handbook of Australian seabirds. Sydney:

Reed.

SHortT, L. L. 1970. The affinity of African with Neotropical woodpeckers. Ostrich, Suppl. 8:

35-40.

SKEAD, C. H. 1964. Sugarbirds in the Amatole Mountains, King William’s Town, Cape

Province. Ostrich 35: 236.

SMITHERS, R. H. N. 1964. A check list of the birds of Bechuanaland Protectorate and the Caprivi

Strip. Salisbury: National Museums of Southern Rhodesia.

SMITHERS, R. H. N., IRwin, M. P. S. & PATERSON, M. L. 1957. A check list of the birds of

Southern Rhodesia. Salisbury: Rhodesian Ornithological Society.

Upvarpy, M. D. F. 1969. Dynamic zoogeography. New York: Van Nostrand Reinhold.

UrsBan, E. K. & Brown, L. H. 1971. A check list of the birds of Ethiopia. Addis Ababa:

University Press.

VAN DER PiaaT, A. 1961. Lys van wilde voéls vir die Oranje-Vrystaat. S. Afr. Avifauna Ser.

P. Fitzpatrick Inst. Afr. Orn. 2: 1-69.

VINCENT, J. 1951. Note on some Swaziland birds. Ostrich 22: 123-125.

Voous, K. H. 1965. Antarctic birds. Jn: MIEGHEM, J. VAN & Oye, P. VAN, eds. Biogeography and

ecology in Antarctica: 649-689. The Hague: Junk. (Monographiae biol. 1£.)

WALLACE, A. R. 1876. The geographical distribution of animals. London: Macmillan.

Watson, G. E., Zusi, R. L. & Storer, R. 1963. Preliminary field guide to the birds of the Indian

Ocean. Washington: Smithsonian Institution.

Waite, C. M. N. 1965. A revised check list of African non-passerine birds. Lusaka: Government

Printer.

WILLouGusy, E. J. & Cave, T. J. 1967. Drinking habits of birds in the central Namib Desert

of South West Africa. Scient. Pap. Namib Des. Res. Stn 31: 1-35.

WINTERBOTTOM, J. M. 1942. A contribution to the ornithology of Barotseland. Jbis (14) 6:

18-27.

WINTERBOTTOM, J. M. 1959. Some zoo-geographical remarks on the South African avifauna.

Ostrich, Suppl. 3: 40-46.

WINTERBOTTOM, J. M. 1962. The zoo-geographical affinities of the avifauna of the northern and

eastern Transvaal. Ostrich 33 (2): 32-37.

WINTERBOTTOM, J. M. 1965. Faunal groups in the avifauna of southern Africa. Revue Zool. Bot.

afr. 71: 157-170ter.

WINTERBOTTOM, J. M. 1966. Avifaunal resemblances between the Neotropical and Ethiopian

Regions. Hornero 10: 209-214.

WINTERBOTTOM, J. M. 1967. The relationships of some African aquatic avifaunas. Revue Zool.

Bot. afr. 75: 148-155.

WINTERBOTTOM, J. M. 1968a. Remarks on some forest avifaunas of southern Africa. Ostrich

39: 146-149.

THE ZOOGEOGRAPHY OF THE SOUTH AFRICAN AVIFAUNA 143

WINTERBOTTOM, J. M. 1968. A check list of the land and fresh water birds of the western Cape

Province. Ann. S. Afr. Mus. 53: 1-276.

WINTERBOTTOM, J. M. 1968c. Remarks on the avifauna of the macchia of the southern Cape

Province. Revue Zool. Bot. afr. 77: 221-235.

WINTERBOTTOM, J. M. 1969. Check list of the birds of South Africa. Cape Town: South African

Ornithological Society.

WINTERBOTTIOM, J. M. 1970. The zoogeography of the south west arid area of Africa. Jnt. orn.

Congr. 15 (Abstr.): 238.

WINTERBOTTOM, J. M. 1971a. A preliminary check list of the birds of South West Africa. Wind-

hoek: S.W.A. Scientific Society.

WINTERBOTTOM, J. M. 19715. The position of Marion Island in the sub-Antarctic avifauna. Jn:

VAN ZINDEREN BAKKER, E. M., WINTERBOTTOM, J. M. & Dyer, R. A., eds. Marion and

Prince Edward Islands: 241-248. Cape Town: Balkema.

WINTERBOTTIOM, J. M. 1972a. Ecological distribution of birds in southern Africa. Monogr.

P. Fitzpatrick Inst. Afr. Orn. 1: i-vil, 1-82.

WINTERBOTTOM, J. M. 19726. The Mopane avifauna. S. Afr. Avifauna Ser. P. Fitzpatrick Inst.

Afr. Orn, 87: 1-19.

WINTERBOTTOM, J. M. 1972c. The avifaunas of the arid districts of Africa. Revue Zool. Bot.

afr. 85: 131-141.

WINTERBOTTOM, J. M. 1973. The antiquity of the African avifauna. Ostrich 44: 139.

ZEUNER, F. E. 1959. The Pleistocene period. London: Hutchinson.

144 ANNALS OF THE SOUTH AFRICAN MUSEUM

APPENDIX

English names of birds mentioned in the text

Achaetops pycnopygius —Damara Rockjumper

Acrocephalus baeticatus—South African Marsh-Warbler

gracilirostris—Cape Reed-Warbler

Actophilornis africanus— African Jagana

Agapornis roseicollis—Rosy-faced Lovebird

Agalaius phoeniceus —Red-winged Blackbird

Alaemon burra—Red Lark

grayi—Gray’s Lark

Alcedo cristata— Malachite Kingfisher

Alopochen aegyptiacus —Egyptian Goose

Amadina erythrocephala—Red-headed Finch

fasciata—Cut-throat Finch

Anas capensis —Cape Teal

erythrorhyncha—Red-billed Teal

smithii— Cape Shoveller

sparsa— Black Duck

undulata—Yellow-billed Duck

Anastomus lamelligerus —Openbill

Andropadus flaviventris— Y ellow-breasted Bulbul

importunus —Sombre Bulbul

virens — Little Greenbul

Anhinga rufa— African Darter

Anthoscopus minutus —Penduline Tit

Anthreptes neglecta—Ulunguru Violet-backed Sunbird

reichenowi—Blue-throated Sunbird

Anthropoides paradisea— Blue Crane

Anthus chloris—Yellow-breasted Pipit

crenatus — African Rock-Pipit

melindae — Malindi Pipit

sokokensis —Sokoke Pipit

Apalis ruddi—Rudd’s Apalis

Apus myoptilus—Scarce Swift

Ardea cinerea—Grey Heron

goliath—Goliath-Heron

melanocephala—Black-headed Heron

purpurea— Purple Heron

Ardeola ibis —Cattle-Egret

ralloides —Squacco Heron

rufiventris —Rufous-bellied Heron

Balaearica pavonina—Crowned Crane

Batis fratrum—Woodwards’ Batis

pririt—Pririt Batis

Bias musicus — Black-and-White Flycatcher

Bostrychia hagedash—Hadeda

Bradypterus babaecalus— African Sedge-Warbler

barratti—Scrub-Warbler

sylvaticus —Knysna Scrub-Warbler

victorini— Victorin’s Warbler

Bubalornis albirostris —Buffalo-Weaver

Bubo capensis —Cape Eagle-Owl

Burhinus vermiculatus —Water-Dikkop

Butorides striatus —Green-backed Heron

Calandrella conirostris —Pink-billed Lark

Sringillaris—Botha’s Lark

sclateri—Sclater’s Lark

THE ZOOGEOGRAPHY OF THE SOUTH AFRICAN AVIFAUNA

Calandrella starki—Stark’s Lark

Calendula magnirostris —Thick-billed Lark

Camaroptera stierlingi—Stierling’s Barred Warbler

subcinnamomea—Cinnamon-breasted Warbler

Campethera abingoni—Golden-tailed Woodpecker

bennetti— Bennett’s Woodpecker

notata— Knysna Woodpecker

Centropus cupreicaudus —Coppery-tailed Coucal

Cercomela schlegelii— Karoo Chat

sinuata—Sickle-winged Chat

tractrac—Tractrac Chat

Certhilauda albescens— Karoo Lark

albofasciata—Spike-heeled Lark

curvirostris —Long-billed Lark

Ceryle maxima—Giant-Kingfisher

rudis— Pied Kingfisher

Ceuthmochares aereus— Green Coucal

Charadrius marginatus —White-fronted Plover

pecuarius—Kittlitz’s Plover

tricollaris—Three-Banded Plover

Chlidonias leucoptera—White-winged Black Tern

Ciconia ciconia—White Stork

nigra—Black Stork

Circus maurus— Black Harrier

ranivorus— African Marsh-Harrier

Cisticola dambo—Black-tailed Cisticola

juncidis—Fan-tailed Cisticola

pipiens —Chirping Cisticola

subruficapilla—Grey-backed Cisticola

tinniens—Le Vaillant’s Cisticola

Colius colius—White-backed Mousebird

Coracias spatulata—Racket-tailed Roller

Cossypha dichroa—Chorister-Robin

heuglini— Heuglin’s Robin

Cursoirus rufus—Burchell’s Courser

Dendrocygna bicolor—Fulvous Tree-Duck

viduata—White-faced Duck

Dicrurus adsimilis— Fork-tailed Drongo

ludwigii— Square-tailed Drongo

Dromas ardeola—Crab-Plover

Egretta alba—Great White Egret

ardesiaca— Black Heron

garzetta—Little Egret

intermedia— Yellow-billed Egret

Emberiza capensis—Cape Bunting

impetuani—Lark-like Bunting

Ephippiorhynchus senegalensis —Saddle-billed Stork

Eremomela gregalis— Karoo Green Warbler

Eremopterix australis—Black-eared Finch-Lark

verticalis—Grey-backed Finch-Lark

Erythrocercus holochlorus—Little Yellow Flycatcher

livingstonei— Livingstone’s Flycatcher

mecalli—Chestnut-capped Flycatcher

Erythropygia coryphaeus—Karoo Robin

paena— Kalahari Robin

quadrivirgata— Bearded Robin

signata— Brown Robin

Estrilda erythronotos—Black-cheeked Waxbill

Euplectes axillaris—Red-shouldered Widowbird

145

146 ANNALS OF THE SOUTH AFRICAN MUSEUM

Euplectes nigroventris— Zanzibar Red Bishop

Eupodotis afer —Black Korhaan

caerulescens —Blue Korhaan

ruficrista —Red-crested Korhaan

vigorsii— Karoo Korhaan

Eurystomus glaucurus—Broad-billed Roller

Francolinus adspersus—Red-billed Francolin

africanus —Grey-winged Francolin

capensis —Cape Francolin

hartlaubi— Hartlaub’s Francolin

levaillantoides —Orange River Francolin

rovumae—Rovuma Francolin

sephaena—Crested Francolin

Fregata minor—Greater Frigatebird

Fulica cristata—Red-knobbed Coot

Fulmarus glacialoides— Antarctic Fulmar

Gallinago nigripennis— Ethiopian Snipe

Gallinula chloropus—Moorhen

Geocolaptes olivaceus—Ground-Woodpecker

Geronticus calvus— Bald Ibis

Glareola pratincola—Pratincole

Gutter edouardi—Crested Guineafowl

Haematopus moquini— Black Oystercatcher

Halcyon senegaloides— Mangrove-Kingfisher

Haliaeetus vocifer — Fish-Eagle

Himantopus himantopus — Black-winged Stilt

Hirundo albigularis—White-throated Swallow

spilodera—Cliff-Swallow

Hydroprogne caspia—Caspian Tern

Hypargos niveoguttatus —Red-throated Twinspot

Ibis ibis —Wood-Ibis

Ixobrychus minutus — Little Bittern

Lagonosticta senegala—Red-billed Firefinch

Lamprotornis acuticaudus—Sharp-tailed Starling

australis —Burchell’s Starling

corruscus — Black-bellied Starling

mevesii— Long-tailed Starling

nitens —Cape Starling

Laniarius aethiopicus—Tropical Boubou

atrococcineus —Crimson-breasted Shrike

bicolor —Western Boubou

Lanioturdus torquatus —White-tailed Shrike

Lanius collaris—Fiscal-Shrike

souzae —Souza’s Shrike

Larus cirrocephalus—Grey-headed Gull

dominicanus —Southern Black-backed Gull

novaehollandiae — Hartlaub’s Gull

Leptoptilus crumeniferus — Marabou

Limnocorax flavirostra— Black Crake

Lioptilus nigricapillus —Bush-Blackcap

Macrodiptervx vexillarius —Pennant-winged Nightjar

Macronyx aurantigula— Pangani Longclaw

capensis —Orange-throated Longclaw

croceus — Y ellow-throated longclaw

Macrosphenus concolor —Grey Longbill

kretschmeri—Kretschmer’s Longbill

Malacocincla rufipennis —Pale-breasted Iladopsis

Malaconotus quadricolor —Gorgeous Bush-Shrike

zeylonus — Bokmakierie

THE ZOOGEOGRAPHY OF THE SOUTH AFRICAN AVIFAUNA

Mandingoa nitidula— Green Twinspot

Melaenornis infuscatus —Chat-Flycatcher

mariquensis — Marico Flycatcher

silens — Fiscal-Flycatcher

Melierax canorus—Chanting Goshawk

metabates— Dark Chanting Goshawk

Merops apiaster—European Bee-eater

pusillus—Little Bee-eater

Mirafra africanoides—Fawn-coloured Lark

apiata—Clapper-Lark

chuana—Short-clawed Lark

ruddi—Rudd’s Lark

rufocinnamomea— Flappet-Lark

sabota—Sabota Lark

Monticola angolensis— Angola Thrush

brevipes—Short-toed Rock-Thrush

explorator —Sentinel Rock-Thrush

rupestris— Cape Rock-Thrush

Motacilla capensis—Cape Wagtail

Myioparus boehmi—Bohm’s Flycatcher

Myrmecocichla formicivora— Ant-eating Chat

Namibornis herero— Herero Chat

Nectarinia bifasciata—Purple-banded Sunbird

famosa— Malachite Sunbird

fusca—Dusky- Sunbird

kilimensis— Bronze Sunbird

mariquensis—Marico Sunbird

neergaardi—Neergaard’s Sunbird

olivacea— Olive Sunbird

pembae—Pemba Sunbird

senegalensis —Scarlet-chested Sunbird

veroxii— Grey Sunbird

violacea—Orange-breasted Sunbird

Neocossyphus poensis— White-tailed Ant-Thrush

Netta erythrophthalma—Southern Pochard

Nettapus auritus—Pygmy-Goose

Nicator gularis—Yellow-spotted Nicator

Nycticorax nycticorax —Night-Heron

Oenanthe monticola— Mountain-Chat

Onychognathus nabouroup —Pale-winged Starling

Ortygospiza locustella—Locust-Finch

Otis kori—Kori Bustard

ludwigii— Ludwig’s Bustard

Oxyura punctata—Maccoa Duck

Pagodroma nivea—Snow-Petrel

Parisoma layardi—Layard’s Tit-babbler

subcaeruleum—Tit-babbler

Parus rufiventris—Rufous Tit

Passer motitensis—Great Sparrow

Pelecanus onocrotalus—White Pelican

rufescens — Pink-backed Pelican

Phalacrocorax africanus —Reed-Cormorant

capensis —Cape Cormorant

lucidus — White-breasted Cormorant

neglectus— Bank-Cormorant

Phoenicopterus minor —Lesser Flamingo

ruber —Greater Flamingo

Phyllastrephus debilis—Slender Bulbul

Pinarornis plumosus—Boulder-Chat

147

148 ANNALS OF THE SOUTH AFRICAN MUSEUM

Platalea alba— African Spoonbill

Platysteira peltata— Black-throated Wattle-eye

Plectropterus gambensis —Spur-winged Goose

Plegadis falcinellus—Glossy Ibis

Ploceus capensis—Cape Weaver

golandi—Clarke’s Weaver

olivaceiceps —Olive-headed Weaver

rubiginosus —Chestnut-Weaver

subaureus— Yellow Weaver

velatus— Masked Weaver

Podiceps cristatus— Great Crested Grebe

nigricollis—Black-necked Grebe

ruficollis —Dabchick

Pogonocichla swynnertoni—Swynnerton’s Robin

Pogoniulus bilineatus—Golden-rumped Tinker-Barbet

pusillus—Red-fronted Tinker-Barbet

simplex —Green Tinker-Barbet

Poicephalus cryptoxanthus—Brown-headed Parrot

robustus —Cape Parrot

rueppellii—Riuppell’s Parrot

Poliohierax semitorquatus—Pygmy-Falcon

Porphyrio porphyrio—Purple Gallinule

Prinia flavicans—Black-chested Prinia

maculosa— Karoo Prinia

pectoralis—Rufous-eared Prinia

robertsi—Roberts’s Prinia

subflava—Tawny-flanked Prinia

substriata— Namaqua Prinia

Prionops scopifrons—Chestnut-fronted Helmet-Shrike

Promerops cafer —Cape Sugarbird

gurneyi—Gurney’s Sugarbird

Pterocles burchelli—Spotted Sandgrouse

gutturalis—Yellow-throated Sandgrouse

namaqua—Namaqua Sandgrouse

Pycnonotus barbatus—Black-eyed Bulbul

capensis —Cape Bulbul

nigricans —Red-eyed Bulbul

Rallus caerulescens—Cape Rail

Recurvirostra avosetta— Avocet

Riparia paludicola— African Sand-Martin

Sarkidiornis melanotus—Knob-billed Duck

Sarothrura affinis—Striped Flufftail

Scopus umbretta—Hamerkop

Seicercus laurae—Mtrs Boulton’s Warbler

Serinus alario— Black-headed Canary

albogularis —White-throated Seed-eater

citrinipectus — Lemon-breasted Canary

flaviventris— Yellow Canary

leucupterus — Protea Seed-eater

mennelli— Black-eared Seed-eater

scotops — Forest-Canary

tottus — Mountain-Siskin

Sheppardia gunningi—Gunning’s Robin

Smithornis capensis— African Broadbill

Spheniscus demersus —Jackass-Penguin

Sphenoeacus afer —Grassbird

Sporopipes squamifrons—Scaly Weaver

Spreo bicolor—Pied Starling

fischeri—Fischer’s Starling

THE ZOOGEOGRAPHY OF THE SOUTH AFRICAN AVIFAUNA

Stactolaema leucotis—White-eared Barbet

whytii— Whyte’s Barbet

Stenostira scita—Fairy-Flycatcher

Sterna balaenarum—Damara Tern

benghalensis— Lesser Crested Tern

bergii— Swift Tern

fuscata—Sooty Tern

vittata— Antarctic Tern

Streptopelia capicola—Cape Turtle-Dove

semitorquata—Red-eyed Dove

senegalensis— Laughing Dove

Sturnella magna— Meadowlark

Sula bassana—Cape Gannet

sula—Red-footed Booby

Sylvietta rufescens—Long-billed Crombec

Tadorna cana—South African Shelduck

Tauraco fischeri—Fischer’s Turaco

porphyriolophus—Purple-crested Lourie

Tchagra tchagra—Tchagra

Thalassoica antarctica— Antarctic Petrel

Thalassornis leuconotus—White-backed Duck

Thamnolaea arnoti—Arnot’s Chat

Threskiornis aethiopica—Sacred Ibis

Tockus alboterminatus—Crowned Hornbill

bradfieldi— Bradfield’s Hornbill

monteiri— Monteiro’s Hornbill

Turdoides bicolor—Pied Babbler

gymnogenys—Bare-cheeked Babbler

jardineii— Arrow-marked Babbler

melanops—Black-faced Babbler

squamulatus—Scaly Babbler

Turdus fischeri— Natal Thrush

Uraeginthus granatinus—Violet-eared Waxbill

Vanellus armatus—Blacksmith-Plover

crassirostris— White-winged Plover

senegallus—Wattled Plover

Vidua regia—Shaft-tailed Whydah

Zosterops senegalensis— Yellow White-eye

149

INSTRUCTIONS TO AUTHORS

Based on

CONFERENCE OF BIOLOGICAL EDITORS, COMMITTEE ON FORM AND STYLE. 1960.

Style manual for biological journals. Washington: American Institute of Biological Sciences.

MANUSCRIPT

To be typewritten, double spaced, with good margins arranged in the following order:

(1) Heading, consisting of informative but brief title, name(s) of author(s), address(es) of

author(s), number of illustrations (figures, enumerated maps and tables) in the article.

(2) Contents. (3) The main text, divided into principal divisions with major headings; sub-

headings to be used sparingly and enumeration of headings to be avoided. (4) Summary.

(5) Acknowledgements. (6) References, as below.

Figure captions and tables to be on separate sheets.

ILLUSTRATIONS

To be reducible to 12 cm x 18 cm (19 cm including caption). A metric scale to appear

with all photographs.

All illustrations to be termed figures (plates are not printed; half-tones will appear in their

proper place in the text), with arabic numbering; items of composite figures to be designated

by capital letters (A, B, C etc.).

REFERENCES

Harvard system (name and year) to be used: author’s name and year of publication given

in text; full references at the end of the article, arranged alphabetically by names, chronologi-

cally within each name, with suffixes a, b, etc. to the year for more than one paper by the same

author in that year.

For books give title in italics, edition, volume number, place of publication, publisher.

For journal articles give title of article, title of journal in italics (abbreviated according to the

World list of scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses,

volume number, part number (only if independently paged) in parentheses, pagination.

Examples (note capitalization and punctuation)

BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FIscHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris

88: 100-140.

FIsCcHER, P.-H., DUVAL, M. & RarFy, A. 1933. Etudes sur les échanges respiratoires des

littorines. Archs Zool. exp. gén. 74: 627-634.

Konun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee

region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 1960b. Spawning behaviour, egg masses and larval development in Conus from the

Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L.

Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und

zentralen Siid-Afrika 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16:

269-270.

ZOOLOGICAL NOMENCLATURE

To be governed by the rulings of the latest International code of zoological nomenclature

issued by the International Trust for Zoological Nomenclature (particularly articles 22 and

51). The Harvard system of reference to be used in the synonymy lists, with the full

references incorporated in the list at the end of the article, and not given in contracted form

in the synonymy list.

Example

Scalaria coronata Lamarck, 1816: pl. 451, figs 5 a, b; Liste: 11. Turton, 1932: 80.

J. M. WINTERBOTTOM

THE ZOOQOGEOGRAPHY OF

THE SOUTH AFRICAN AVIFAUNA

BOT + ek

NOVEMBER 1974

VOLUME 66 PART 7

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

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November 1974 November

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

PEDI SKIN DRESSING TECHNIQUE

E. M. SHAW & H. E. BOHME

Photographs by J. B. KRAMER

South African Museum, Cape Town

(With 28 figures)

[MS accepted 16 November 1973]

CONTENTS

PAGE

Introductions? Jo. 's. ee eh ears. CLS

IDSSCHDUONG el) ie ke os Me ee welioil

NcCknOWledeementsin le). Gees a eee Or,

INTRODUCTION

In May 1973 a visit was made to the BaPedi of the Nebo area of Sekhu-

khuniland in Lebowa, and at Chief Sekwati’s village a demonstration was seen

of the process of dressing an ox-skin. The following information about the

technique was obtained from Johannes Mogeng Kgolone, who had learnt the

craft from his father, and who practised as a specialist skinworker. He had

regular assistance from a brother, and called in friends to help with the more

strenuous processes involved. On the occasion that we were present at his home-

stead he rewarded them with beer, and when we gave a few individual presents

to those participating he firmly insisted on them all being handed over, which

they were without demur.

DESCRIPTION

After the skin is removed from the animal, it is pegged out to dry. It then

becomes hard and may be stored like this until needed. To begin the dressing

process the skin is soaked in clear water in a three-legged pot (Fig. 1). A stone is

placed on the skin to keep it submerged. The moisture softens the skin and the

hairs can be removed more easily. When the skin is very well soaked the process _

of scraping and further softening commences. This is a long procedure which,

according to the skinworker, can take from a day, when he has many helpers, to

four days. There is a more or less set sequence of events.

The damp skin is held outstretched by four or five men with the hairy side

facing upwards and one man proceeds to scrape the skin (removing the hair)

with an iron rod, bent in the shape of a U (Fig. 2). This is done till the surface

moisture and most of the loose hair is scraped off. The hairy side now has a

‘patchy’ look (Fig. 3).

Ann. S. Afr. Mus. 66 (7), 1974: 151-167, 28 figs.

151

152 ANNALS OF THE SOUTH AFRICAN MUSEUM

Figure 2

PEDI SKIN DRESSING TECHNIQUE 153

Figure 3

Then the scraped skin is turned over, so that the inner side faces upwards

and the hair side down, and is stretched and pegged with wooden pegs on to the

ground on a layer of dried grass. The grass is there to keep the wet skin clean. A

sack of dried grass is specially kept for this purpose. A stone is used to hit the

pegs into the ground (Fig. 4). Various tools are used for scraping and currying

but in no definite sequence.

=o SESS ee So ‘oy $ J - *¥

F .

154 ANNALS OF THE SOUTH AFRICAN MUSEUM

Figure 5

A thorny aloe leaf (/ilabo) (Fig. 5) and an adze (/epalo) (Fig. 6) are used at

this stage to scrape the skin. When using the adze each worker first wets his

working area and then proceeds to scrape in the following way: the haft of the

adze is held in the left hand, while the right hand cups the hafthead and the adze

is worked towards him in short jerky movements (Fig. 7).

Figure 6

PEDI SKIN DRESSING TECHNIQUE 155

Figure 7

After short intervals of scraping, the blade of the adze is sharpened on an

ordinary knife, a whetstone (se/otsu) or another adze blade.

Figure 8

156 ANNALS OF THE SOUTH AFRICAN MUSEUM

Figure 9

Following the scraping the preparation continues using either rough stones

(those seen were pieces of lateritic ironstone) (Fig. 8), tse/a, a name also given to

a wooden handle with rows of metal spikes along its base, or to a hollow metal

tube with a serrated edge, or aloe leaves (Fig. 9). Abrading the skin proceeded

on this occasion for more than fifteen minutes but would take much longer had

the skinworker not had as much help as he had during our stay.

Figure 10

PEDI SKIN DRESSING TECHNIQUE 157

Figure 11

The skin is unpegged and the metal scraper used again to remove any loose

scrapings. It is then carried to a near-by boulder where it is left to dry out fora

short while in the sun (Fig. 10).

The skin is then softened by rubbing it between the hands (Fig. 11). The

next step is to twist it around a tree trunk with a stick (Fig. 12) in order to wring

out all the remaining water.

Figure 12

158 ANNALS OF THE SOUTH AFRICAN MUSEUM

Figure 13

The twisted skin is pricked with a nail to allow all the water to seep out

(Fig. 13) and is then opened out and pulled hard to stretch it (Fig. 14). Then

follows a long session of braying with the hands. The left hand clasps the skin

tightly whereas the right grips it and rolls it over the fist of the left hand. This

is repeated over and over, moving from one part of the skin to the next (Fig. 15).

Figure 14

PEDI SKIN DRESSING TECHNIQUE 159

% f,

Cee

aN, i

a \

Figure 15

After the skin as been sufficiently softened it is held stretched out again and

two helpers scour it with aloe leaves or with tse/a. One scoured in a diagonal

direction whereas the other scraped in parallel movements (Fig. 16). It is then

thrown on to the boulder again, this time with the inner side facing the rock.

160 ANNALS OF THE SOUTH AFRICAN MUSEUM

Figure 17

After drying for a while, the skin is kneaded again with the hands. Finally it is

pulled taut and the aloe leaf or the tse/a rubbed over it to raise a pile (Fig. 17).

At this stage the skin is still quite damp.

OP a

Figure 18

PEDI SKIN DRESSING TECHNIQUE 161]

“3 NE I As NE 5, |

Figure 19

The next step is to smear the skin with fat from the fruit (ditshidi) of the

mochidi tree (Ximenia caffra Sond.). The small fruit is red when ripe but turns

black on boiling. After boiling it is allowed to dry out and then the kernel is

taken out and ground between grinding stones till it forms an oily paste (Fig. 18).

The skin is again pegged out, hair-side down on the layer of dried grass and the

fat is rubbed into the inner side of the skin with a wooden peg (pabolo).

After some time the hide is again brayed, rubbed and wrung with the hands

to help the fat to draw deep into the skin.

The hide is now ready to be cut but before doing this the skinworker uses

an ordinary knife to straighten it out. Then he proceeds to cut a garment to the

required shape. The article seen was a woman’s back apron or skirt, ntepa, and

the informants stated that two could be cut out of a small hide and three out of a

large hide. From the offcuts a girl’s smaller back apron could be made. The

width across the top of an apron was taken to be more or less the span of three

hands.

When the skin has been cut out and shaped it is laid on the ground and

covered with manure which is left on for about a day. Powdered dry manure is

mixed with fresh wet manure for this process (Fig. 19). This is the finai step in

the preparation, after which the garment is coloured with red ochre.

A woman’s front apron, ntetho, is always made of goat-skin which is

prepared in the same way as the ox-skin, until the cutting process is reached.

After the skin has been cut to the correct pattern it still has to be shaped.

Before shaping, however, it is decorated in the following way: the prepared skin

is moistened where the working is to begin. A household knife is used to incise

a design on the skin (Figs 20, 21). This design covers the entire surface of the

162 ANNALS OF THE SOUTH AFRICAN MUSEUM

Figure 21

PEDI SKIN DRESSING TECHNIQUE 163

Figure 22

ntetho and when worn the decorated side is on the outside (Figs 22, 23). The

knife which is used for doing these designs has to be sharpened at intervals on a

whetstone (se/otsu).

To shape the apron, strips of skin are cut out of it, again using the knife,

similar to the way a dressmaker makes darts except that in this case the ‘dart’ is

cut out and the apron sewn together again (Fig. 24). The fibre (/eshika) which is

used is made from the sinew of cattle. It is taken from the carcass while it is still

fresh. The dry sinew is then rolled to form a two-ply fibre which is, as was seen in

one case, kept in a hussif (molalachika) made from ‘bobbejaanstert’ (Vellozia

equisetoides) (Fig. 25). An awl is used for making the holes through which the

sewing strand is passed (Fig. 26).

The back apron, ntepa (Figs 27, 28), is always coloured with red ochre

(Jetsoko) which is rubbed into it and gives it a dark red glow. The front apron is

sometimes coloured red but it may according to choice have chalk rubbed into

it to bring out the incised design more clearly. This gives it a beige colouring.

This skinworker purchased red ochre from the near-by Penge Mine, but his

supply of chalk came from his family who live near Sekhukhuni (Schoonoord).

The preparation of other types of skins was said not to differ from the

technique described. A blanket (/ethebo) was seen that was made from many

different small skins, among them mongoose, springbok and hyrax.

164 ANNALS OF THE SOUTH AFRICAN MUSEUM

PEDI SKIN DRESSING TECHNIQUE

Figure 25

Figure 26

166 ANNALS OF THE SOUTH AFRICAN MUSEUM

Figure 27

The skinworker said, however, that there was a shorter method of skin-

dressing which cut out some stages of the preparation. The main stages of

soaking the skin in water, scraping it with an adze, braying with the hands and

finally cutting and shaping it, were kept.

ACKNOWLEDGEMENTS

We are indebted to the Secretary for Bantu Administration and Develop-

ment, the Magistrate, Nebo, the Lebowa Government, and Chief Sekwati for

permission to work in the area; to Johannes Mogeng Kgolone for the informa-

tion given and to him and his assistants for the very excellent demonstration of

skin dressing; and to Kgoloko Motlokwe Mompuru and Constable Titus Mdluli

who acted as interpreters.

Acknowledgement is made of the financial assistance of the Human Sciences

Research Council towards the project of which this study forms a part. Opinions

expressed or conclusions reached are those of the authors and should not be

regarded as representative of those of the Human Sciences Research Council.

PEDI SKIN DRESSING TECHNIQUE 167

Figure 28

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Harvard system (name and year) to be used: author’s name and year of publication given

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Examples (note capitalization and punctuation)

BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris

88: 100-140.

FiscHer, P.-H., DUvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des

littorines. Archs Zool. exp. gén. 74: 627-634.

Konun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee

region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the

Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L.

Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und

zentralen Siid-Afrika. 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16:

269-270.

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Scalaria coronata Lamarck, 1816: pl. 451, figs 5 a, b; Liste: 11. Turton, 1932: 80.

E. M. Shaw & H. E. Bohme

PEDI SKIN DRESSING TECHNIQUE

VOLUME 66 PART 8 NOVEMBER 1974

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CAPE TOWN

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 66 Band

November 1974 November

Part 8 Deel

A NEW SPECIES OF

MYSTACOCARIDA (CRUSTACEA)

FROM ALGOA BAY, SOUTH AFRICA

A. McLACHLAN

‘ ,

Jo Roy GRIN DEE Y

Cape Town Kaapstad

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A NEW SPECIES OF MYSTACOCARIDA (CRUSTACEA)

FROM ALGOA BAY, SOUTH AFRICA

A. MCLACHLAN

University of Port Elizabeth

J. R. GRINDLEY

Port Elizabeth Museum

(With | figure and 2 tables)

[MS accepted 15 January 1974]

CONTENTS

PAGE

Introductronns, 22) 2c) ye ow bee ce me ae 169

IMOTCTHIAIS ata toet- eet oe a) Uke (ee cem ae es 170

Systematic descriptions us ene aelO

DISCUSSION y's hte wee ene) ee eal 174

SUMMA etay ona ty Ee wero) pen vce, amet Ceres 174

FACKNMOWICUZEMIGMESH a ern Ly Ge Wee ten LS

INCICREIM COS Bens ek cae ic ass oh het a Roe rnin ok Ws

INTRODUCTION

Until the revision by Hessler (1972) of the African mystacocarids, all those

recorded on the African coasts were regarded as subspecies of Derocheilocaris

remanei Delamare Deboutteville & Chappius. D. r. katesae Noodt was recorded

from Durban, South Africa; D. r. remanei Delamare Deboutteville & Chappius

from Swakopmund, South West Africa; D. r. remanei from Luanda, Angola;

D. r. remanei from Kayar, Senegal, and D. r. remanei from the Mediterranean

(Hessler 1972).

In the light of the discovery of a distinct species, D. ingens Hessler, in

Maine, within 260 km of the known range of D. typica Pennak & Zinn, Hessler

(1972) raised the subspecies of D. remanei from Angola and Swakopmund to

specific status as D. angolensis and D. delamarei respectively. Although Hessler

(1972) was not able to examine specimens of D. r. katesae from Durban he

stated that because it was separated from other populations of D. remanei by the

Swakopmund and Angolan species it probably also warranted specific status.

Hessler (1972) based his differentiation of species on very small morpho-

logical differences; differences which were not recognized as specific differences

by Noodt (1954). Confirmation of the validity of the specific status of such

slightly different morphological forms has now come from the co-occurrence of

two closely related species in Algoa Bay, South Africa.

Ann. S. Afr. Mus. 66 (8), 1974: 169-175, 1 fig., 2 tables

169

170 ANNALS OF THE SOUTH AFRICAN MUSEUM

Here two distinct mystacocarid species, one of them undescribed, have been

taken from the same part of the same beach. This is the second recorded overlap

of distribution of two mystacocarid species. One has been identified as D.

delamarei Hessler as it appears to differ from the original description in only the

very slightest details. Eight adult specimens of this species have been found in

Algoa Bay to date. The other species recorded in Algoa Bay differs significantly

from previously described species and is here designated a new species.

MATERIALS

All the specimens described here have been obtained from the intertidal

sands of King’s Beach, the main bathing beach of Port Elizabeth (33°58’S,

25°39’E) and Bluewater Bay (33°51’S, 25°39’E). Both these beaches are in

Algoa Bay. Sand samples were collected during low and mid tides between

January and July 1973. The mystacocarids were extracted by means of an

Oostenbrink extractor (Oostenbrink 1960) as modified by Furstenberg (personal

communication). They were obtained using a sieve with 75 » mesh and were

preserved in 4% formaldehyde.

SYSTEMATIC DESCRIPTIONS

Derocheilocaris algoensis sp. n.

(Fig. 1 A-D)

Holotype—S.A.M. A13535 in the South African Museum, Cape Town.

An adult female from the mid-tide region of King’s Beach, Port Elizabeth,

South Africa.

Paratypes—S.A.M. A13536 in the South African Museum, Cape Town.

U.S.N.M. 149165 in the United States National Museum, Washing-

ton.

Numerous adult males, females and juveniles from King’s Beach, Port Elizabeth,

South Africa.

Remarks

This species is named after Algoa Bay, its type locality and the only place

where it has been recorded to date.

Occurrence

Numerous specimens of different larval stages as well as male and female

adults have been obtained from King’s Beach. These were found to occur deeper

than about 10 cm in the sand between the spring low tide and mid to high tide

marks. In some samples they appeared to be the most numerous meiobenthic

animals in the deeper sand.

A NEW SPECIES OF MYSTACOCARIDA (CRUSTACEA) FROM ALGOA BAY 171

EEN tc

BANS NS

Fig. 1.

Derocheilocaris algoensis sp. n. from King’s Beach, Algoa Bay. A—dorsal view of adult;

B—dorsal view of cephalon; C—ventral view of labrum; D—lateral view of telson showing

furcal claws and setae.

172 ANNALS OF THE SOUTH AFRICAN MUSEUM

Diagnosis of adult

(All terms are as used by Hessler 1972.)

Anteromedial lobes of antennulary portion of cephalon without medial

spines as in de/amarei (Fig. 1B); sides of antennulary portion of cephalon convex

and not converging appreciably posteriorly; posterolateral corners of anten-

nulary portion of cephalon moderately angular; constriction separating anten-

nulary portion of cephalon from rest of cephalon a moderately deep notch in

dorsal view; immediately posterior to notch sides of cephalon converge before

expanding into antennary region; anterolateral pair of spines in antennary

region of cephalon not borne on pronounced protuberances as in remanei.

Posterior part of labrum (Fig. IC) not as expanded as in D. typica and with

a slight notch in some specimens.

Lateral toothed furrows moderately developed with irregular margins and

no distinct teeth.

Distal segment of maxilla I with 8 setae.

Endopod of maxilliped with | seta on distal segment and 2 setae on penulti-

mate segment.

Supra-anal process (Fig. 1D) terminating in moderate acute process ventral

and well-developed acute process dorsal to supra-anal seta; no strongly

developed spines around base of acute processes; setules on ventral setae of

furcal claws smaller than in delamarei; supra-anal seta usually longer than

middle or dorsal furcal setae.

The ratios of the lengths of the different furcal and the supra-anal setae to

the lengths of the furcal claws were measured as by Hessler (1972) and are given

in Table 1. Figures in Table | marked with an asterisk differ significantly

(p = 0,01) from values for the same ratios in all species described by Hessler

Cis):

Table 1.

Ratios of lengths of supra-anal and furcal claw setae to furcal claw

lengths in adult D. algoensis.

No Mean S.D Range

*SSe/F 16 HO 0,09 0,84-1,18

M/F 16 0,94 0,08 0,77-1,07

D/F 16 0,88 0,11 0,68-1,05

aN 16 2.25 0,16 1,95-2,49

F = furcal claw length; SSe = supra-anal seta length; M = middle

furcal seta length; D = dorsal furcal seta length; V = ventral furcal

seta length; INo = number of specimens measured; S.D. = standard

deviation;

* indicates ratios significantly different from those for previously

described species.

A NEW SPECIES OF MYSTACOCARIDA (CRUSTACEA) FROM ALGOA BAY 173

In 15 adults measured, the body length (from the anterior tip of the cephalon

to the posterior tip of the furcal claws) averaged 0,39 + 0,02 mm with a range

of 0,37—-0,42 mm.

D. algoensis is most quickly identified by (1) the shape of the anterior

portion of the cephalon (Fig. 1B), (2) the long (but not as long as D. delamare’)

ventral furcal setae (Fig. 1D). Table 2 is a summary of some of the more

important features whereby D. r. remanei, D. delamarei, D. algoensis and

D. r. katesae can be distinguished.

Table 2.

Summary of main distinguishing characteristics for four mystacocarids recorded on the

African coasts. Compiled from Noodt (1954), Hessler (1972) and the present work.

FEATURE D.r. remanei | D. delamarei D. algoensis D. r. katesae

Antero-medial notch of U-shaped U-shaped U-shaped V-shaped

antennulary portion of with spines

cephalon

Constriction separating notched notched notched not notched

antennulary part of

cephalon from rest of

cephalon

Anterolateral spine on on on on margin =

antennary part of protuberance | protuberance

cephalon

Toothed furrows well well moderately

developed developed developed

SSe longer or shorter than shorter shorter longer —

M and D

Ventral furcal seta short very long long long

Spines on supra-anal small large small small

process

Acute process(es) on ventral ventral ventral & ventral

supra-anal process to SSe to SSe dorsal to SSe to SSe

Larvae

A number of larvae corresponding to the later larval stages described by

Hessler & Sanders (1966) for D. typica have been found during the winter

months. It is hoped that at a later stage when a complete series of larval stages

has been collected it might be possible to publish a description of the larval

development of D. algoensis.

174 ANNALS OF THE SOUTH AFRICAN MUSEUM

Derocheilocaris delamarei Hessler, 1972

Eight adults and a number of larvae were found near and below low tide at

King’s Beach and Bluewater Bay. They were taken in the surface 10 cm of the

sand during late summer. These specimens are immediately distinguishable from

D. algoensis by their very long ventral furcal setae which bear long setules. They

appear to resemble D. delamarei in practically all details. The only difference

noted is that the ventral furcal setae bear 27-37 setules whereas D. delamarei as

illustrated by Hessler (1972) bears 48 setules on the ventral furcal setae. Setal

length/furcal claw length ratios fall in the same order as recorded for D. dela-

marei (Hessler 1972). The number of setules on the ventral furcal setae of

D. delamarei as well as the shape of the cephalon in D. r. katesae may not be

reliable in the published descriptions so that the differences relating to these

features may be of limited value. Considering that 2 100 km of coastline separate

Swakopmund and Algoa Bay, it 1s surprising that D. delamarei differs so little in

form at the end of its known range. The relative rarity of specimens of D. dela-

marei in Algoa Bay might indicate that this is near the eastern limit of the

distribution of this species.

DISCUSSION

Two distinct mystacocarid species are here recorded from the same beach.

One species, D. de/amarei, appears to occur in the surface 10 cm of the sand and

has only been found in summer, while another species, D. algoensis, is plentiful

at depths greater than 10 cm in the sand. Algoa Bay is probably near the eastern

limit of the range of D. delamarei. This is the second recorded overlap between

two mystacocarid species.

These two species differ in details so small that, before the publication of

Hessler’s (1972) review, they would not have been considered sufficient to

warrant separate species status. Nevertheless, because these small differences

remain distinct, even where the species occur together, they must indicate

different species. This therefore confirms the view of Hessler (1972) that mystaco-

carids are very conservative and that small yet constant differences do constitute

specific differences. In this respect the shape of the antennulary portion of the

cephalon and the lengths of the supra-anal and furcal claw spines appear to be

especially important characteristics.

SUMMARY

An overlap of distribution of two mystacocarid species is recorded from

Algoa Bay (33°58’S, 25°39’E). One is an undescribed species, here named

Derocheilocaris algoensis sp. n. and the other has been identified as D. delamarei

Hessler. This is the second recorded case of overlap of mystacocarid species and

supports the views of Hessler (1972) that small differences between mystacocarids

do constitute specific differences.

A NEW SPECIES OF MYSTACOCARIDA (CRUSTACEA) FROM ALGOA BAY i

ACKNOWLEDGEMENTS

This work was assisted by a research grant from the University of Port

Elizabeth. We thank Dr J. P. Furstenberg of the Zoology Department of the

University of Port Elizabeth for valuable criticism.

REFERENCES

ARMSTRONG, J. C. 1949. The systematic position of the genus Derocheilocaris and the status of

the subclass Mystacocarida. Am. Mus. Novit. 1413: 1-6.

DauL, E. 1952. Reports of the Lund University Chile Expedition 1948-49; 7. Mystacocarida.

Acta Univ. lund. 48: 1-41.

DELAMARE DEBOUTTEVILLE, C. 1960. Biologie des eaux souterraines littorales et continentales.

Paris: Hermann.

*DELAMARE DEBOUTTEVILLE, C. & DE BARROS MACHADO, A. 1954. Presence de la sous-classe

des Mystacocarides sur les cétes de Angola. PublgGdes cult. Co. Diam. Angola 23: 118-124.

*DELAMARE DEBOUTTEVILLE, C. & CHAPPIUS, P. A. 1951. Présence de l’orde des Mystacocarida

Pennak et Zinn dans le sable des plages du Rousillon: Derocheilocaris remanei n. sp.

C. r. hebd. Acad. Sci., Paris 233: 437-439.

*DELAMARE DEBOUTTEVILLE, C. & CHAPPIUS, P. A. 1957. Contribution a l’étude de la faune

interstitielle marine des c6tes d’Afrique. 1. Mystacocarides, Copepodes et Isopodes. Bull.

Inst. fr. Afr. noire (A) 19: 491-500.

HEssLer, R. R. 1972. New species of Mystacocarida from Africa. Crustaceana 22: 259-273.

HEssLer, R. R. & SANDERS, H. L. 1966. Derocheilocaris typicus Pennak and Zinn (Mystoco-

carida) revisited. Crustaceana 11: 141-155.

JANSSON, B. 1966. On the ecology of Derocheilocaris remanei Delamare and Chappius

(Crustacea, Mystacocarida). Vie Milieu 17: 43-186.

Noopt, W. 1954. Crustacea Mystacocarida von Suid Afrika. Kieler Meeresforsch. 10: 243-246.

Noopt, W. 1961. Neuevo hallazgo de Derocheilocaris galvarini Dahl en Chile central

(Crustacea, Mystacocarida). /nvestnes zool. chil. 7: 97-99.

OOSTENBRINK, M. 1960. Estimate nematode populations by some selected methods. Jn:

SASSER, J. N. & JENKINS, W. R., eds. Nematology fundamentals and recent advances with

emphasis on plant parasites and soil forms: 85-102. Chapel Hill: University of North

Carolina Press.

PENNAK, R. W. & ZINN, D. J. 1943. Mystacocarida, a new order of Crustacea from intertidal

beaches in Massachusetts and Connecticut. Smithson. misc. Collns 103: 1-11.

*Not seen in the original.

INSTRUCTIONS TO AUTHORS

Based on

CONFERENCE OF BIOLOGICAL EDITORS, COMMITTEE ON FORM AND STYLE. 1960.

Style manual for biological journals. Washington: American Institute of Biological Sciences.

MANUSCRIPT

To be typewritten, double spaced, with good margins arranged in the following order:

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REFERENCES.

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BuLLouGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FIsCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris

88: 100-140.

FIscHER, P.-H., DUVAL, M. & RaArFry, A. 1933. Etudes sur les échanges respiratoires des

littorines. Archs Zool. exp. gén. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee

region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320.

Konan, A. J. 19605. Spawning behaviour, egg masses and larval development in Conus from the

Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. Jn: SCHULTZE, L.

Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und

zentralen Siid-Afrika. 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16:

269-270.

ZOOLOGICAL NOMENCLATURE

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in the synonymy list.

Example

Scalaria coronata Lamarck, 1816: pl. 451, figs 5 a, b; Liste: 11. Turton, 1932: 80.

5 ~*

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A. McLachlan & J. R. Grindley

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ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 66 Band

February 1975 Februarie

Part 9 ‘Deel

SOUTH AFRICAN CUMACEA

PART 1

FAMILY BODOTRIIDAE,

SUBFAMILY VAUNTHOMPSONIINAE

JENNIFER DAY

Cape Town Kaapstad

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SOUTH AFRICAN CUMACEA

PART 1

FAMILY BODOTRIIDAE, SUBFAMILY VAUNTHOMPSONIINAE

JENNIFER DAY

Zoology Department, University of Cape Town

(With 15 figures and | table)

[MS accepted 21 June 1974]

CONTENTS

PAGE

Introduction . 5 : ; : ; ; Sia alee

Material : s ; : : : 3 5 like

Station data . : : : ; : j ree SS

Methods j ; 5 . : Me i gis:

Structure and terminology : ; , : SSO

Systematics . 2 182

Key to the southern African species of Vaunthomp-

soniinae . : : ; ‘ é 21) eek 83

Pscudosympodommea ; 5 : : ; Sa dkesss)

Heterocuma : é , : : 5 ASS

Cumopsis . ‘ ‘ : : : F He Pll)

Hypocuma . : : : ; i : SMe ay be

Vaunthompsonia . : ; : : : 5 y= PADI

Bathycuma. : : , Lo M2OF

Distribution of the Vaunthompsoniinae : 216

Distribution of the South African Vaunthompsoniinae 217

Summary . : : : : : meee)

Acknowledgements : : ; f ; ee

References . : , ; ; ; : Me 20

INTRODUCTION

This is the first in a series of papers on the Cumacea of southern Africa

(south of 20°S). The families best represented in these waters are the Bodotriidae

and Diastylidae. There are relatively few Leuconidae, Lampropidae and

Nannastacidae, while the Pseudocumatidae are numerous, but confined almost

exclusively to estuaries, where there are perhaps two species. There are also

further specimens of the Ceratocumatidae to be described in a later paper in

the series.

Studies on the cumacean fauna of southern African waters are scanty.

Stebbing published two papers (1910, 1912) on material sent to him by the South

African Museum, mainly describing specimens collected during the voyages of

the s.s. Pieter Faure from 1898 to 1907. Zimmer (1908) included several South

African species in his descriptions of the Deutsches Tiefsee-Expedition material,

177

Ann. S. Afr. Mus. 66 (9), 1975: 177-220, 15 figs, 1 table

178 ANNALS OF THE SOUTH AFRICAN MUSEUM

and later (1921) described a few in the collection of the Zoologisches Museum,

Berlin. Since then, Fage (1951) and Jones (1956) have described a few species

from this region in the collections of the Belgian Oceanographic Expedition and

Atlantide series respectively, and Hale (1953) and Jones (1960) have published

on some of the material in the collection of the University of Cape Town.

Otherwise, the cumacean fauna of the region is poorly known.

MATERIAL

Some of the more interesting specimens were kindly loaned by Dr Brian

Kensley of the South African Museum (SAM) —largely material obtained by the

s.s. Pieter Faure, which is old and therefore frequently decalcified and damaged.

Two specimens of Vaunthompsonia natalensis were sent to me by Mr Tim

McClurg from the Natal Benthic Survey being carried out by the National

Institute for Water Research (NIWR) of the Council for Scientific and Industrial

Research (CSIR). The vast majority of specimens, however, was obtained by the

Zoology Department of the University of Cape Town (UCT) during an extensive

benthic survey of inshore waters from Liideritzbucht in South West Africa to

Inhambane in Mocgambique, the cruises being funded by the Oceanographic

Research Unit of the CSIR. I am particularly grateful for access to this material.

STATION DATA

The UCT material is numbered according to the area from which it was

obtained: WCD (West Coast Dredge) from the border of South West Africa to

Cape Agulhas, LBT from the Lambert’s Bay Transect, FAL from False Bay,

SCD (South Coast Dredge) from Cape Agulhas to the southern border of Natal,

NAD (Natal Dredge) from Natal waters, and PED (Portuguese East Dredge)

from Mocambique (Fig. 1). “Coast’ indicates material from the NIWR survey,

and ‘SAM’, South African Museum material. The depths and positions of all

SAM (Pieter Faure) stations have been approximated from charts, since the

station data are sketchy, and many depths are not given, or are inaccurate.

METHODS

All the SAM and NIWR material was obtained by dredging. In the UCT

survey, heavy-duty dredges and Van Veen grabs of area 0,2 m? were used. Some

of the LBT material was obtained using a scuba-diver-operated suction device

(Christie & Allen 1972). The instrument is a cylinder sampling an area of 0,1 m*

by 60 cm deep, which is lowered right into the substrate. Different fractions of

substrate may then be collected, enabling analysis of fractions from different

levels below the surface. The cumaceans were all obtained in the top 10 cm

fraction.

Length measurements were in all cases taken from the most anterior point

of the carapace to the posterior edge of the telsonic somite, the uropods being

omitted from these measurements.

179

PART |

SOUTH AFRICAN CUMACEA:

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180 ANNALS OF THE SOUTH AFRICAN MUSEUM

STRUCTURE AND TERMINOLOGY

The terminology used by authors who are familiar with a group is frequently

confusing to those who are dealing with it for the first time. Below is a brief

account of the structure of the Cumacea, and the terminology used to describe it.

For a fuller description, the reader is referred to Jones (1963), whose terminology

is adopted in the present series, particularly in the use of the term ‘somite’ as a

metameric segment of the body, and ‘segment’ as a podomere of the appendages.

The Cumacea are typical Malacostraca, having 19 body segments or somites,

each of which may bear a pair of appendages. The head region consists of five

somites, bearing from the anterior end two pairs of antennae, a pair of mandibles

and two pairs of maxillae. There are eight thoracic somites, the anterior three

bearing maxillipeds, and the posterior five (the pedigerous somites) bearing

pereiopods. The abdomen, of six pleon somites, may have up to five pairs of

pleopods in the male, and none in the female. The sixth abdominal somite bears

a pair of uropods. A telson is present in some families, and in others it is fused

with the last abdominal somite which is known as the felsonic somite.

A well-developed carapace is present, fused not only to all five head somites,

but also to three, four, five or six thoracic somites. It is bowed laterally to cover

the branchial chambers, and is produced anteriorly to form pseudorostral lobes,

below which is the exhalent opening of the branchial chamber. The two lobes

together are known as the pseudorostrum. Antero-laterally the carapace is

frequently notched to accommodate the first antenna—the antennal notch—with

a sharp point, the antero-lateral angle, below it. Dorsally, if visual elements are

present, they are usually fused into a single median eye, frequently pigmented,

with a variable number of lenses. The eye is situated on the eye/obe, a median

projection of the carapace anteriorly between the pseudorostral lobes. The cara-

pace may be quite smooth, but is often ornamented with tubercles, spines or

other projections, or is sculptured into median and/or lateral keels or carinae.

The free thoracic and abdominal somites may also be carinate.

APPENDAGES

Antenna I (antennule) consists of three basal segments bearing two flagella.

The main flagellum, of up to six segments, carries a number of sensory filaments

or aesthetascs, which are usually annulated. The accessory flagellum may have

up to four segments, or may be wanting.

Antenna 2 is rudimentary in the female, but may reach the entire length of

the body in the male, where it may be used for sensing the presence of the female,

or for clasping her during copulation.

Mandibles are formed from a single segment, and are normally crescent-

shaped.

Maxilla I (maxillule) is rudimentary, but may bear one or two epipodal

filaments reaching into the branchial chamber.

Maxilla 2 (maxilla) is also rudimentary.

SOUTH AFRICAN CUMACEA: PART | 181

The thoracic appendages are typical stenopodous limbs. Exopods, some of

which may be rudimentary, are always found consecutively on at least two of

these limbs from maxilliped 3 to pereiopod 4 inclusive. The maximum number

is five, and there are usually more in the male than the female. The exopods

consist of a basal segment, sometimes expanded into a laminar plate, and a

variable number of distal segments which usually carry long plumose setae.

Well-developed exopods assist considerably in swimming. The endopeod of a

thoracic appendage is typically divided into seven segments. These are named

(from the proximal end) the coxa, basis, ischium, merus, carpus, propodus and

dactyl.

Maxilliped I normally consists of a seven-segmented endopod and an

epipodite developed into a series of plates or lobes which function as the gill.

Maxilliped 2 is also normally seven-segmented. The coxa of ovigerous

females is produced to form a rudimentary oostegite, bearing filaments which

extend into the brood chamber and fan the developing embryos.

Maxilliped 3, always seven-segmented, usually bears a well-developed exo-

pod. The outer distal portion of the basis is frequently expanded, and the basis

forms a shield protecting the mid-ventral region of the thorax.

Pereiopod I, always bearing an exopod, is seven-segmented, and is frequently

elongated, reaching beyond the anterior end of the pseudorostral lobes.

Pereiopods 2 to 5 may or may not bear exopods, and are normally seven-

segmented, although in some genera the ischium of pereiopod 2 is fused with the

basis. :

The male may bear up to five pairs of biramous pleopods. These may be

rudimentary, but if fully developed consist of a basal segment, a 1-segmented

inner ramus and a 2-segmented outer ramus. The distal ends of the rami are

frequently supplied with a large number of plumose setae used in swimming.

The uropods, also biramous, consist of a l-segmented peduncle, a 2-seg-

mented exopod and an endopod which may consist of one, two or three segments.

These appendages form the forked tail characteristic of all Cumacea.

SEXUAL DIMORPHISM

The Cumacea exhibit considerable sexual dimorphism. The sexual diffe-

rences include small details such as the sculpturing and degree of armature of

the exoskeleton, but most other more basic differences can be attributed to

characters allowing greater swimming capacity in the male, and the presence of

a marsupium or brood pouch in the female. Distinguishing male from female is

important, since several characters of the males of some genera are the same as

the female characters of another. Determination of sex is not always easy,

especially in an immature animal, but the following may be helpful as a guideline:

The adult male has between zero and five pairs of pleopods, usually seti-

ferous, and the second antennae extend a considerable distance along the body.

Thoracic exopods are frequently flattened and expanded, the antero-lateral angle

rounded or obliterated, and serrations of the carapace reduced or absent.

182 ANNALS OF THE SOUTH AFRICAN MUSEUM

The subadult male has a full complement of pleopods, not yet

setiferous, and the second antennae are still developing. Sculpturing of the

exoskeleton is often midway between the condition of the adult male and the

female.

The ovigerous female is distinguished by the presence of a marsupium. The

coxae of maxilliped 3 and pereiopods | to 3 develop oostegites, forming a large

ventral brood chamber in which the eggs are visible during the later stages of

development.

The adult female differs from an ovigerous one only in the absence of a

marsupium.

The manca, the larval stage at which the animals are released from the

marsupium, is characterized by the absence of the last pair of pereiopods.

The juveniles may be recognized, apart from their small size, by the absence

of sexual dimorphism.

The majority of males and females in most collections are intermediate

between the fully adult and juvenile stages, and although some sexual differences

are present, they are not always easily determined. For example, in many

bodotriids, the lateral plates of the pedigerous somites differ even in young males

and females. However, this type of character is variable, and must be determined

anew for each genus or species. One distinguishing characteristic which holds

true for many (but not all) genera is that the male tends to have more pereiopods

bearing exopods than does the female.

SYSTEMATICS

The Cumacea, being a rather homogeneous group, have few characters

which divide them obviously into families. Thus two main schemes have been

used to separate them. The one, detailed by Stebbing in 1913, now has 27

families. The other more commonly adopted scheme has seven families.

Although neither of these systems is ideal, the latter is less cumbersome and will

be adopted in the present series of papers.

Family Bodotriidae Scott, 1901

Diagnosis

No free telson. Pleopods with an outer process to the inner ramus—usually

five pairs, but may be two (Mancocuma) or three (Leptocuma). Mandibles

normal (i.e. not broad at base). Endopod of uropod 1I- or 2-segmented. Branchial

apparatus without gill plates or supports.

The family was divided by Hale (1944) into two subfamilies, the Bodotriinae

and Vaunthompsoniinae. The Bodotriinae are characterized by having exopods

limited to the third maxillipeds and first pereiopods in both sexes. The South

African representatives of the subfamily will be dealt with in the second paper of

this series.

SOUTH AFRICAN CUMACEA: PART | 183

Subfamily Vaunthompsoniinae G. O. Sars, 1879

Diagnosis

Bodotriidae with exopods on pereiopods other than the first pair.

KEY TO THE SOUTH AFRICAN SPECIES OF VAUNTHOMPSONIINAE

It should be noted that this key will separate all species found to date in

southern African waters, but will not necessarily distinguish them from species

from other areas.

imeExapous present on pereiopods t-and Zionly (Gand Q). a5.55 2. oc ee ec. cee sc cnccececes

Pseudosympodomma africanum (Fig. 2)

SeEKONOGS pLesent.on perelopods | to. 3 (Gand 9). oooh ecicje cca cues Sos wee seabhnucece ce 2

Pee noOUs.OrperelOpods 2 and 3, TUGIMENLATY, = sls 4c) = acces vias «eccs sae eo mianae wes > 3

— Exopods of pereiopods 2 and 3 well-developed; ¢ with exopod on pereiopod 4.......... 5

3 Maxilliped 3 with basis not at all produced distally. .Cumopsis robusta sp. nov. (Figs 5, 6)

Sm MiaxiMedisuwith: Oasis produced distally... 25.2 osc aces ces cle ec ees wget ae ens 4

4 Pereiopods 2 and 3 with exopods 1-segmented......... Heterocuma africanum africanum

=SEChOpousr. ang > With -exOpods 2-sepmented.: s 2. 42..ciac- es secs ek ee ete ew es selene

Heterocuma africanum intermedium (Figs 3, 4)

Se Viaxuiped > with basis not atallproduced distally 5.5225 22. joes. ne ce dee we ws oes 6

Sewiaxihipedes. with: basis) produced distally... c.0554 500 +c ence ees 66 sles ses ocisien deecles 9

6 Eye absent; carpus of maxilliped 3 inserted half way along length of merus; merus expanded

SUIS EIST gs BARA ee a ee eres gee Hypocuma dentatum sp. nov. (Figs 7, 8)

— Eye present; carpus of maxilliped 3 not expanded, and inserted at distal end of merus....7

fesehiated mucddorsal carina presentin ?: 3: .......6.-.052 ++ see oe Vaunthompsonia cristata

NORMA GORSAL Carinaypresemt 1M feMAale. . sic oe tg wel o wliteec wie le coe ec Siele wee % aisle ere bree 8

8 Merus of maxilliped 3 longer than carpus; first segment of exopod of uropod less than half

length of first segment of endopod........ Vaunthompsonia natalensis sp. nov. (Figs 9, 10)

— Merus of maxilliped 3 shorter than carpus; first segment of exopod of uropod nearly as

longerasshrst Sepiiment Of CNGOPOd...:.......6.05+ + sa eeme +s Vaunthompsonia sp. (Fig. 11)

9 Carapace of ¢ and 2 with middorsal serrations; telsonic somite produced between uropods

LOm Ams eerMtS LEM OUI A pres ic eie cmc Slee ett Gene, SUR Ee nem, Tot ae AE UON, ete Wh vane. ola wasPaiors 10

— Carapace of at least with no middorsal serrations; telsonic somite little produced between

SAO) OS PRMD sch Sh oltsrciiy Ae cai-soh a, Saree ohsuaoaseaeh cies yanaeetens Bathycuma datum sp. nov. (Fig. 15)

10 Peduncle of uropod longer than telsonic somite; antenna 1 of adult ¢ very large, with

MAEM ESCUAC Bes ih oils a ou he Ale fade tel ecapaviahes by ottase arene. sani cece Bathycuma natalense (Figs 13, 14)

— Peduncle of uropod shorter than telsonic somite; antenna 1 of ¢ unmodified...........-

Bathycuma capense (Figs 12, 13)

Pseudosympodomma Kurian, 1954

Generic diagnosis

Eyelobe narrowly linguiform with a cluster of lenses anteriorly—more in

male than female. Pseudorostral lobes extending anteriorly but not meeting in

front of elongated eyelobe. All pedigerous somites exposed. Basis of maxilliped 3

distally produced. Pereiopods 1 and 2 with free exopods in both sexes; adult

male sometimes with a small ridge in position of exopod of pereiopod 3. Male

with five pairs of pleopods. Endopod of uropod 2-segmented.

Type species P. indicum Kurian, 1954, from India, ‘in weeds, 0,2 fm’.

184 ANNALS OF THE SOUTH AFRICAN MUSEUM

Pseudosympodomma africanum* (Stebbing, 1912)

Fig. 2

Sympodomma africanus Stebbing, 1912: 138, pl. L.

non Sympodomma africanum: Hale, 1928: 40, figs 9-10; 1944: 284, fig. 30.

Records

SAM A4331, Pieter Faure 10782: 29°49’S/31°08’E 85m 19 10,9 mm

SAM A597, Pieter Faure 17643: 34°24’S/17°58’E 370m _ 19 adult 15,7 mm PARATYPE

SAM A598 _ Pieter Faure 17643: 34°24’S/17°58’E 370m _ 1d subadult 15,7 mm

PED 23 X__ 22.8.1964 26°00’S/33°0S’E 135m _ 1 manca 4,3 mm

Note: In his description of S. africanum, Stebbing gave the depth as 805 m.

In fact, if the distance of 124 miles from Cape Natal (Pieter Faure log) is correct,

then the depth can be no more than 400 m in any direction, according to present-

day charts. The bearing also appears to be the reciprocal of the normal bearings

in the log, as “Cape Natal N 3/4 E 123 miles’ is practically in the surf zone.

Holotype designated by Stebbing (1912) as Sympodomma africanus, Pieter

Faure 17643 (as above), subadult male, 18,0 mm. British Museum (Natural

History).

Description

Adult female, length 15,7 mm, paratype. Broken in two, but otherwise

undamaged.

Carapace faintly tubercular, nearly twice as long as deep; median carina

with three very large, anteriorly-directed teeth on the anterior part (Fig. 2A).

Eye present, lenses distinct—seven above and seven below (Figs 2B, 2C).

Eyelobe elongated, pseudorostral lobes not meeting. Antennal notch fairly

shallow, semicircular; antero-lateral angle almost rectangular. All pedigerous

somites visible above. Sternite of fourth pedigerous somite with anteriorly-

directed tooth. Carapace as long as free thoracic somites. Cephalothorax equal

in length to first four abdominal somites. Pedigerous somites with faint lateral

carinae; two pairs of lateral carinae along length of abdomen. Telsonic somite

produced between uropods for nearly } its length.

Antenna | (Fig. 2D) geniculate between segments | and 2. First segment

large, slightly expanded laterally, equal in length to next two segments. Flagellum

2-segmented with two terminal setae. Accessory flagellum 2-segmented, equal in

length to first segment of flagellum.

Maxilliped 3 (Fig. 2E) with basis 24 times length of remaining segments,

widely expanded at distal end, with four spines along the outer edge and two

apical ones. Inner edge with a row of small denticles along its length. Remaining

segments almost equal in length, ischium slightly the longest. Merus expanded

slightly externally and carpus internally. Exopod small, basal segment less than

+ length of basis.

*The words ‘omma’ (Gr.), an eye, and ‘cuma’ (Gr.), an embryo, are grammatically neuter,

hence Pseudosympodomma africanum, Bathycuma natalense, etc.

SOUTH AFRICAN CUMACEA: PART | 185

Fig. 2. Pseudosympodomma africanum (Stebbing, 1912)

Adult female, paratype, 15,7 mm: A, lateral view; B, dorsal view of carapace; C, ventral view

of eyelobe; D, antenna 1; E, maxilliped 3; F, pereiopod 1; G, pereiopod 2; H, pereiopod 3;

I, telsonic somite and uropod.

Subadult male, 15,7 mm: J, lateral view; K, dorsal view of carapace; L, distal portion of

maxilliped 3; M, proximal segments of pereiopod 3.

Pereiopod | (Fig. 2F) fairly stout. Basis equal in length to next four seg-

ments. Ischium short. Propodus and dactyl long, subequal, each equal in length

to merus plus carpus. Exopod small, basal segment not much more than a

quarter length of basis.

Pereiopod 2 (Fig. 2G) stout. Basis equal in length to next four segments.

Ischium 4 as long as broad. Merus and carpus subequal. Dactyl equal to merus

186 ANNALS OF THE SOUTH AFRICAN MUSEUM

plus carpus and bearing a number of scattered spines. Basal segment of exopod

2 length of basis.

Pereiopod 3 (Fig. 2H) more slender. Basis equal in length to rest of leg.

Ischium small, half as long as broad. Merus and carpus subequal. Carpus just

longer than propodus, which is again slightly longer than dactyl. No exopod.

Peduncle of uropod (Fig. 2I) just longer than telsonic somite with 20

unequal spines along inner edge. Endopod slightly longer than exopod and

about 2 length of peduncle; first segment twice length of second, with eight small

spines on inner margin, second with four, and one terminally. Exopod with first

segment half the length of second, and unarmed. Second segment with three

plumose setae on the inner margin and four terminal spines.

Subadult male, length 15,7 mm. In poor condition, uropods and parts of

some thoracic appendages missing. As in the female, except as follows: teeth on

middorsal carina less elevated, and followed by a row of uneven tubercles, the

dorsal outline thus appearing uneven. Tubercles scattered over the rest of the

carapace (Figs 2J, 2K). Antennal notch much shallower, antero-lateral angle less

obvious. Ventral margin faintly dentate below the angle for about 3 its length.

Sternites of fourth and fifth pedigerous somites with an anteriorly-directed

tooth in the midline. Abdominal somites with three pairs of carinae.

Distal prolongation of maxilliped 3 (Fig. 2L) narrower than in female, with

four distal spines. Inner edge more spinous. Merus and carpus further expanded,

these edges dentate.

Pereiopods as in female. Pereiopod 3 (Fig. 2M) without a free exopod, but

with a slight swelling along the outer edge of the basis for about 3 its length.

Five pairs of pleopods present.

Uropods missing. Telsonic somite of holotype male broader than that of

female. Stebbing’s figure of uropods of holotype as in female of present collec-

tion, but with greater armature.

Remarks

Stebbing (1912) described the species from a single subadult male, and the

above specimens differ from his description only as follows: carapace of holotype

less tubercular; distinct individual lenses present in eye of female, but eye of

male as figured by Stebbing. Maxilliped 3 of holotype with merus and carpus

slightly more expanded, with setae and not denticles along expanded edges.

Stebbing figured the basal segment of antenna | as geniculate. The antennae of

the holotype are missing, but in the present specimens the antenna is geniculate

between the first and second segments.

In 1912 Stebbing erected a new family, the Sympodommatidae, with the single

genus Sympodomma to receive four species—Vaunthompsonia anomalum

(G. O. Sars, 1871), Heterocuma weberi Calman, 1905, Heterocuma diomedaea

Calman, 1912 and a new species, Sympodomma africanum. The main distinguish-

ing feature of the new family and genus was the presence of exopods on pereio-

pod 3 in both sexes. These were obviously present (according to descriptions in

SOUTH AFRICAN CUMACEA: PART | 187

the literature) in the first three species, but he says of S. africanum (one

subadult male) that: ‘exopods to the third pair were not satisfactorily made

out, but may be presumed, as they occur in both sexes of the allied Japanese

species.’

In addition to the holotype, I have examined four specimens which

undoubtedly belong to Stebbing’s S. africanum. One is a paratype, a subadult

female, from the type locality with the same station data and museum number as

the holotype. Another specimen, a subadult male from the same locality but with

different station data, has a swelling in the expected position of the exopod of

pereiopod 3, but this is fused to the basis along its entire length. Unfortunately

the holotype now consists of little more than an empty carapace, three badly

damaged and limbless thoracic somites and several abdominal somites, so that

the pereiopods in question could not be examined. Neither of the females nor

the manca had the slightest trace of an exopod on these limbs.

In 1954 Kurian erected a new genus, Pseudosympodomma, to receive a new

Indian species, P. indicum, which has exopods on pereiopods | and 2 only in

both sexes. He suggested that Sympodomma africanum might be referable to the

new genus. Owing to the lack of an exopod on pereiopod 3 of the present

specimens, Stebbing’s S. africanum must now be referred to the genus Pseudo-

sympodomma.

The two species of the genus may be distinguished as follows: the pseudo-

rostral lobes and eyelobes of P. indicum are relatively longer than those of

P. africanum. P. indicum has a 3-segmented accessory flagellum, while that of

P. africanum is 2-segmented. In P. indicum the exopod of maxilliped 3 is less than

a quarter the length of the basis; the prolongation of the basis is narrower and

extends well beyond the merus. The last two segments of the first pereiopod are

longer and more slender. The two segments of the endopod of the uropod are

subequal, the two rami are of equal length, and the first segment of the exopod

is only a quarter the length of the second. In the description of P. indicum, no

mention is made of the teeth on the sternites of the pedigerous somites.

The two genera Sympodomma and Pseudosympodomma are very similar in

general appearance, in distinctive features such as the large teeth or incisions on

the dorsal carina, and the presence of teeth on the thoracic sternites of several

species. Thus the genera are closely related, and the single feature distinguishing

between them is the presence or absence of exopods on pereiopod 3. In the female

this is quite clear, but in the male even this character is less distinctive due to the

small protuberances on the bases of these legs. Thus the generic diagnosis of

Pseudosympodomma has been expanded to accommodate P. africanum.

It should also be noted that Hale (1928, 1944, 1949) had some specimens

which he considered to be S. africanum. Later, being unable to obtain more

material, he erected a new species, Sympodomma ?incerta Hale, 1949. It is very

similar to P. africanum in external appearance, but due to the presence of an

obvious exopod on pereiopod 3 of the male, Hale’s query may be removed, and

the species becomes Sympodomma incertum.

188 ANNALS OF THE SOUTH AFRICAN MUSEUM

Distribution of Pseudosympodomma

P. indicum is found in shallow Indian waters (Kurian 1954), and P. africanum

from Natal to Cape Point at depths from 85 to 370 m. Thus the genus falls into

the Indo-Pacific faunal group.

Heterocuma Miers, 1879

Generic diagnosis

Vaunthompsoniinae with the first pedigerous somite visible dorsally.

Telsonic somite not produced. Eye present. Basis and merus of maxilliped 3

distally produced. Pereiopods 2 and 3 with rudimentary exopods in both sexes.

Endopod of uropod 2-segmented. Male with five pairs of pleopods.

Type species H. sarsi Miers, 1879, from Korea and Japan.

KEY TO THE SPECIES OF HETEROCUMA

l Pereiopods)2 and 3 with exopods 2-segmentediaw «42-45. 4-20 e see see eee

H. africanum intermedium (Fage, 1924)—West and South Africa

= Peretopods 2 and 3 with exopods l-segmented=. 5..-.5..---4e5s0 cee | tte ee 2

2 Pleon segments carinate sf hhc Fs ca ce oe ca Caw oe wei heed oe 6 66 ae tla 3

= Pleon: segments. not.carimate. 6.56 0s le ais Se geo cee 4

3 First segment of endopod of urepod' shorter than second: 5.32 )4.- 1-1. 1-1

H. sarsi Miers, 1879— Korea and Japan

— First segment of endopod of uropod longer than second. . H. andamani Kurian,1954—India

Endopod of uropod distinctly shorter than peduncle: .2 5... .- 1-2 4..2= eee

H. africanum africanum Zimmer, 1921 —India, West and South Africa

— Endopod of uropod just longer than peduncle.......... H. armatum Kurian, 1954—India

Heterocuma africanum intermedium (Fage, 1924) n. comb.

Figs 3 (2), 4 (3)

Heterocuma intermedia Fage, 1924: 364, fig. 1.

Heterocuma africana: Jones, 1956: 194 (?pars); 1960: 172.

Records

WCD 99 R 32705'S/1817-B. 2.7 196te 27m Sandy 1¢ adult 18,4 mm

shell

SCD SNe 34°55’S/21°26’E 21.2.1960 91m Sand 19 adult 19,1 mm

192 ovig. 24,8 mm

SCD 293 F 33°04’S/27°S7V’E 6.2.1962 84m Sandy 19 ovig. 22,4 mm

shell

FAL (False Bay) 34°S/18°E 15-77 m Sandy 15 records:

shell 2 adult g 18,2-18,4 mm;

1 subadult ¢ 19,3 mm;

23 7,9-9,4 mm;

32 ovig. 19,2-23,2 mm;

82 adult 17,6-21,6 mm;

52 7,4-11,0 mm;

5 manca 4,0-5,8 mm (1

adult g, 1 2 ovig. and 2 2

adult identified by Jones

1960)

SOUTH AFRICAN CUMACEA: PART 1 189

Fig. 3. Heterocuma africanum intermedium (Fage, 1924)

Ovigerous female, 23,2 mm: A, lateral view; B, detail of anterior end of carapace; C, dorsal

view of carapace; D, antenna 1; E, maxilliped 3; F, pereiopod 1; G, pereiopod 2; H, pereiopod

3; I, pereiopod 4; J, pereiopod 5; K, uropod.

190 ANNALS OF THE SOUTH AFRICAN MUSEUM

Holotype designated by Fage (1924) as Heterocuma intermedia; Museum

National d’Histoire Naturelle, Paris.

Description

Ovigerous female, length 23,2 mm. Carapace twice as long as deep with a

faint mid-dorsal carina. In lateral view (Fig. 3A), mid-dorsal line irregular and

faintly arched posteriorly. Antennal notch deep, antero-lateral angle acute

(Fig. 3B). Carapace just shorter than remaining free thoracic somites. Thorax

and abdomen subequal. No lateral plates or carinae on abdomen. Eye present

with single lens faintly visible on either side. Pseudorostral lobes not meeting

anterior to eyelobe (Fig. 3C).

Antenna | (Fig. 3D) with basal segment very much expanded, being twice as

long and twice as broad as second segment. Second and third segments subequal,

second broader. Flagellum 2-segmented, first twice as long and as broad as

second which carries a number of terminal setae and two aesthetascs. Accessory

flagellum minute, 1-segmented.

Maxilliped 3 (Fig. 3E) with well-developed broad distal prolongation of the

basis. Ischium normal, merus dilated externally and carpus internally. Propodus

and dactyl subequal. Basal segment of exopod less than a quarter length of basis.

Basis of pereiopod 1 (Fig. 3F) a quarter as long again as rest of leg. Ischium

and merus subequal, as are carpus and propodus. Dactyl half length of propodus.

Basal segment of exopod about a quarter length of basis.

Pereiopod 2 (Fig. 3G) very stout—basis half as long as broad. Ischium

fused with basis. Dactyl with a large number of spines. Exopod large, as long as

the basis, with a small but obvious second segment.

Pereiopod 3 (Fig. 3H) more slender, with exopod half length of basis, and

having a small second segment.

Pereiopods 4 and 5 (Figs 3I and 3J) without exopods. Third to fifth pereio-

pods all furnished with distinctive knobbed setae on the bases.

Peduncle of uropods (Fig. 3K) half as long again as telsonic somite, with one

row of 14 small spines on the inner edge. Exopod equal in length to peduncle,

with first segment half as long as second. First with a single plumose seta,

second with about 20, and four short terminal spines. Endopod # length of

peduncle, first segment about 3 length of second with nine small and one large

terminal spines on the inner edge. Second segment with 9-10 small spines on

inner and outer edges, and five terminal spines of equal length.

Note: a single female from False Bay has a particularly well-marked row of

tubercles on the mid-dorsal line of the carapace, and carinae running down the

length of the abdomen on either side of the mid-line. Otherwise it corresponds

with the above description.

Adult male, length 18,4 mm. As the female, except as follows: carapace with

much less deeply indented antennal notch, antero-lateral angle almost obsolete

(Fig. 4B). Thorax equal in length to next five abdominal somites (Fig. 4A). As is

typical for the family, the thoracic sideplates are well developed, that of the

SOUTH AFRICAN CUMACEA: PART | 191

Fig. 4. Heterocuma africanum intermedium (Fage, 1924).

Adult male, 18,4 mm: A, lateral view; B, detail of anterior end of carapace;

C, detail of eyelobe in dorsal view; D, antenna 1; E, distal segments of antenna 1.

fourth pedigerous somite having a prominent linguiform anterior projection.

The armature of the peduncle of the uropods is more pronounced, having about

50 small, closely-packed spines in two rows along the inner edge. The single

subadult male from the west coast differs from the adult males only in that the

thoracic sideplates and uropodal armature are not fully developed, and the eye

has 10 distinct lenses on the dorsal surface (Fig. 4C). Antenna 1 (Fig. 4D) is

somewhat shorter in the male, and the flagellum is 1-segmented (Fig. 4E).

Distribution

Tropical West Africa (Fage 1924, 1950, 1951; Jones 1960—part); False Bay -

and south-west coast of South Africa (Jones 1960).

Heterocuma africanum africanum Zimmer, 1921

Heterocuma africana Zimmer, 1921: 129, figs 25-27. Kurian, 1954: 294, fig. 7.

Records

NAD 86 G_ 29°10’S/31°51’E_—-_-.29.7.1964 43 m Sand 292 adult 10,4-11,2 mm;

ives 6,8 mm

Holotype designated by Zimmer, 1921 from West Africa, one subadult male.

?Zoologisches Museum, Berlin.

192 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

The single sample from Natal is of interest in that although generally the

specimens are very similar to H. africanum intermedium, they have short,

l-segmented exopods on pereiopods 2 and 3. The eyes of the single male and

both females have lenses similar to those shown in Figure 4C. Although the

females are adult, they are only about half the size of comparable specimens from

False Bay. The carapace has no trace of a carina.

Distribution

Tropical West Africa (Zimmer 1921; Jones 1956—part); India (Kurian

1954); Natal.

Remarks

Heterocuma africanum was described but poorly figured by Zimmer (1921)

from an immature specimen collected together with many juveniles from Free-

town in Sierra Leone. Fage (1924) described H. intermedium from Rio do Oro,

from Goreé (1950) and from Darsen and Annobon Islands (1951). Jones (1956)

examined many specimens from the coast of West Africa, and equated Fage’s

H. intermedium with H. africanum due to the fact that Zimmer obviously

described the latter from an immature specimen. More recently, Jones (1960)

identified several specimens (now in the present collection) from False Bay and

Dassen Island as H. africanum. These are indicated in the records above.

Undoubtedly the two forms are very similar in most respects, but they are in fact

distinguishable by the nature of the exopods of the second and third pereiopods.

In Zimmer’s specimen, they were ‘in the form of short cylinders’, while Fage

(1924) specifically mentions ‘the greater size and the division into two articles of

the exopods of the second and third pereiopods’. Kurian (1954) identified two

adult specimens (7-8 mm) as H. africanum, and both of these are figured as

having short, l-segmented exopods. The majority of the South African speci-

mens have larger, 2-segmented exopods, while the exopods of those from Natal

are smaller and 1-segmented. Jones (1956, 1960) mentioned the differences in

size of some of the West African specimens, and the fact that those from South

Africa are very large, but did not describe the exopods. In fact there is correlation

between the size of the animals and the nature of the exopods. Those with

l-segmented exopods are smaller—Zimmer’s largest was 4 mm, Kurian’s

7-8 mm, and the Natal specimens reach 11,2 mm. Those with the bigger 2-seg-

mented exopods are also larger in size—Fage’s 16 mm, and the largest of the

present specimens nearly 25 mm. The fact that 1-segmented exopods are present

in adult animals from both India and Natal shows that this is not an age-

dependent character, and it is therefore suggested that there are two distinct

forms, insufficiently different to warrant specific rank. Those of Zimmer and

Kurian, together with the NAD specimens are therefore designated H. africanum

africanum, as opposed to those of Fage and Jones, and the majority of the

specimens in the present collection, which become H. africanum intermedium.

SOUTH AFRICAN CUMACEA: PART 1 193

Note: the size differences of the specimens identified by Jones (1956) in

West Africa suggest that both forms may occur there.

Distribution of Heterocuma

H. sarsi, H. armatum and H. andamani are all shallow-water Indian Ocean

or Indo-Pacific species. The two subspecies of H. africanum extend the range to

the tropical Atlantic region.

Cumopsis G. O. Sars, 1878

Generic diagnosis

Vaunthompsoniinae with rudimentary exopods on pereiopods 2 and 3 in

both sexes. Antenna | of male with a brush of sensory setae at base of flagellum.

Basis of maxilliped produced distally very slightly or not at all, carpus not

widened. Male with five pairs of pleopods. Telsonic somite truncate posteriorly,

and not produced between the uropods. Endopod of uropod 2-segmented.

Type species C. longipes (A. Dohrn, 1869) as Cuma longipes from British Isles

and Mediterranean.

KEY TO THE SPECIES OF CUMOPSITS

PGS CEIBIO | ASE PRCSCIIG fe 5 for ciescioren so be Siw Re = Mee eee tors len Co Ree Ors wD ELCs 2

SAE TECK AT OIGSTA SEM bsterte fears CAA e eS che oitick veo ao. Sea are ebeloin oe aiele Sicis aang biel e embers 3

Pe wordistinct lateral folds; carapace flattened dorsally... .. 0.2.0. 06. 02. ce cece cence wee

C. goodsiri (Van Beneden, 1861)—Europe, Indo-Pacific

— A single lateral fold; carapace convex dorsally....... C. wafri Jones, 1956—West Africa

3 Thoracic sideplates well defined dorsally.......... C. elongata Jones, 1956—West Africa

SMoaictcasideplates: not.denned, dorsally. 2 <4 656 is wetecwe ss cleinw ce bios se hae scien eee 4

PE COnnGClcr@eNKOpod: twice lengthy Ol TAM... s-> che.cie ers Sie tsie foe cause eneie as.0 B.s1e syblavmisota sate S)

— Peduncle of uropod only slightly longer than rami.................. C. robusta sp. nov.

Seitstasepment Of exopod of uropod longer than second... ................5.-2-.0---

C. fagei Bacescu, 1956—W. France, Morocco

SEESe Seement OF Exopod ef uropod shorter than second. ..—. 4.2. 2-2: .-2:2.....+0-e0-

C. longipes (A. Dohrn, 1869)—Britain and Mediterranean

Cumopsis robusta sp. nov.

Figs 5 (2), 6 (3)

Records

LBT 5H 15.9.1970 32°04’S/18°18’E 3m _ Sand 1 Q ovig. 3,7 mm HOLOTYPE

LBT 8L 15.9.1970 32°04’S/18°18’°E Sm _ Sand 1 dg adult 4,0 mm

LBT 105F 30.1.1972 32°04’S/18°18’E LWS Sand 59 ovig. 3,7-4,8 mm

LBT 144A Sept. 1972 32°04’S/18°18’E LWS Sand 2 9 ovig. 3,7-4,2 mm

LBT 146E Sept. 1972 32°04’S/18°18’E 1m Sand 1 dsubadult 3,6 mm

EBT 147G Sept. 1972 32°04’S/18°18E 1m Sand 19 3,5 mm

CP 833A May1972 False Bay (34°S/18°E) Sand 1 9 ovig. 4,4 mm

i:9 3,2 mm

194 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 5. Cumopsis robusta sp. nov.

Ovigerous female, holotype, 3,7 mm: A, lateral view; B, dorsal view of carapace; C, antenna 1;

D, detail of distal end of antenna 1; E, maxilliped 3; F, pereiopod 1; G, pereiopod 2; H,

pereiopod 3; I, pereiopod 4; J, telsonic somite and uropod.

SOUTH AFRICAN CUMACEA: PART | 195

Holotype

Ovigerous female, deposited in the South African Museum, number

SAM A13433.

Description

Ovigerous female, holotype, length 3,7 mm. Integument lightly calcified,

with faint reticulations at high magnifications. Carapace nearly twice as long as

deep, and equal in length to remaining free thoracic somites (Fig. 5A). No trace

of lateral folds. Carapace in dorsal view (Fig. 5B) almost rectangular anteriorly

due to short unproduced pseudorostral lobes being interrupted by eyelobe. Eye

elliptical with a number of dark red pigmented rings; lenses not discernable.

Antennal notch hardly defined. Cephalothorax just longer than abdomen.

Abdominal somites without sideplates. Telsonic somite plus uropods equal in

length to the four preceding somites. Mouthparts and lower edge of carapace

dark in fresh specimens.

Three basal segments of antenna | (Fig. 5C) subequal, slight flexure between

first and second. Flagellum with two aesthetascs. Accessory flagellum small,

1-segmented (Fig. 5D).

Basis of maxilliped 3 (Fig. 5E) half as long again as rest of limb, distally

truncate, and broadly widened at its midpoint. Ischium half as long as wide.

Merus slightly flared externally. Carpus and propodus subequal. Basal segment

of exopod less than half length of basis.

Basis of pereiopod | (Fig. 5F) stout, just longer than remaining segments, of

which carpus the longest. Basal segment of exopod 2 length of basis, and slightly

expanded at the mid-point.

Pereiopod 2 (Fig. 5G) stout. Ischium fused with basis. Carpus and propodus

subequal. Dacty! equal in length to merus plus carpus, with many spines. Exopod

l-segmented, and about quarter length of basis, with four long plumose setae.

Pereiopod 3 (Fig. 5H) very stout. Basis equal in length to next four seg-

ments. Merus and carpus subequal. Exopod as in pereiopod 2.

Pereiopod 4 (Fig. 51) stout, with many setae. No exopod.

Telsonic somite square in dorsal view (Fig. 5J), unarmed. Uropods equal

in length to the four preceding somites. Peduncle just longer than rami, with —

three small spines on inner edge. First segment of endopod 14 times length of

second, with six spines on inner edge; second segment with two spines on inner

edge and two long terminal spines. Segments of exopod subequal, first unarmed,

second with two plumose setae on inner edge and two long terminal ones.

Adult male, length 4,0 mm. Differs from female as follows: carapace } as

deep as long; cephalothorax equal in length to next five abdominal somites

(Fig. 6A), which are more robust than in the female. Sideplates of pedigerous and

abdominal somites defined ventrally, that of the fourth pedigerous somite being

produced anteriorly. No antennal notch.

Antenna | (Figs 6B, 6C) very robust, with a brush of sensory setae surround-

ing the 2-segmented flagellum (Fig. 6D).

196 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 6. Cumopsis robusta sp. nov.

Adult male, 4,0 mm: A, lateral view; B, dorsal view of carapace; C, antenna 1; D, detail of

distal end of antenna 1; E, maxilliped 3; F, pereiopod 1; G, pereiopod 2; H, pereiopod 3;

I, telsonic somite and uropod.

SOUTH AFRICAN CUMACEA: PART | 197

Antenna 2 extending just beyond uropods—segments short.

Maxilliped 3 (Fig. 6E) with merus slightly expanded externally; less robust

than in female, and basis not angled.

Propodus of pereiopod | (Fig. 6F) relatively shorter.

Pereiopods 2 and 3 (Figs 6G and 6H) less stout. Merus and carpus of

pereiopod 2 shorter than propodus. Basal segment of exopod nearly half length

of basis.

Telsonic somite and uropods (Fig. 61) equal in length to the three preceding

somites. Inner border of peduncle armed with about 23 small spines. Armature

of rami greater than in female.

Remarks

The species of Cumopsis are rather similar morphologically. C. robusta may

be distinguished from C. fagei, which it closely resembles, as follows: in C. fagei

the peduncle of the uropods is almost twice the length of the endopod, while in

C. robusta the two are almost subequal. In dorsal view the eyelobe and pseudo-

rostral lobes form a straight line anteriorly in C. robusta while in C. fagei the

pseudorostral lobes extend beyond the eyelobe for a short distance. The peduncle

of the uropod is relatively broader and the rami are subequal in C. robusta, while

the exopod is the longer in C. fagei.

Distribution of Cumopsis

This genus is found almost exclusively intertidally or in the infratidal fringe.

It occurs in the British Isles, through the Mediterranean, tropical West Africa

and South Africa to Annam. C. robusta would appear to be endemic to the

south-western coast of South Africa.

Hypocuma Jones, 1973

Generic diagnosis

Vaunthompsoniinae with pseudorostral lobes produced anteriorly to meet

in front of the eyeless eyelobe. No antennal notch or angle in either sex.

Antenna | long and slender. Mandible normal. Five pairs of thoracic appendages

bearing large flattened exopods in the male, and four in the female. Merus of

maxillipeds 2 and 3 expanded, basis of maxilliped 3 not produced distally.

Pereiopod 2 7-segmented. Well-developed exopods on first four pereiopods in

male, and first three in female. Female with rudimentary exopod on pereiopod 4.

Five pairs of pleopods in male. Telsonic somite produced between uropods.

Type species Hypocuma serratifrons Jones, 1973 from 1 934 m off the Canary

Islands.

Hypocuma dentatum sp. nov.

Figs 7 (g), 8 (2)

Records

SAM A10602a, Pieter Faure 17440: 34°25’S/17°50’E about 400 m

1 subadult ¢ 5,9 mm—uropods missing. HOLOTYPE

1 adult 3 6,0 mm—last somite missing

2 adult 2—cephalothorax only

198 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 7. Hypocuma dentatum sp. nov.

Adult male, holotype, 5,9 mm: A, lateral view; B, detail of anterior end of carapace; C, dorsal

view of carapace; D, antenna 1; E, maxilliped 2; F, maxilliped 3; G, pereiopod 1; H, pereio-

pod 2; I, pereiopod 3; J, pereiopod 4; K, pleopod 2; L, dorsal view of telsonic somite; M,

lateral view of telsonic somite.

SOUTH AFRICAN CUMACEA: PART 1 199

Holotype

Subadult male, deposited in the South African Museum, number

SAM A13434.

Description

Subadult male, holotype, length 5,9 mm. Carapace twice as long as deep

with slightly upturned pseudorostrum reaching well beyond eyelobe and

constituting 4 total length of carapace (Fig. 7A). No antennal notch (Fig. 7B).

A row of well-developed denticles on anterior edges of carapace, becoming

smaller and disappearing along ventral edge. Eyelobe eyeless with a row of small

denticles dorsally (Fig. 7C). Exoskeleton generally sculptured with triangular

denticles of varying size. Carapace 13 times length of free thoracic somites, and

of almost equal length to first four abdominal somites. First pedigerous somite

visible dorsally and laterally, second to fourth with slight ventro-lateral flanging.

Abdominal somites with well-developed sideplates. Telsonic somite produced

between uropods for about 4 its length.

Antenna | (Fig. 7D) slender, elongate, and geniculate between first and

second segments. Flagellum 3-segmented, with first segment greatly elongated.

Accessory flagellum small, 2-segmented.

Maxilliped 2 (Fig. 7E) with a row of denticles on inner edge of basis; ischium

small; merus and carpus subequal, inner edge of merus expanded distally, and

bearing five denticles.

Maxilliped 3 (Fig. 7F) with basis slightly longer than combined length of

remaining segments, and not produced distally. Ischium twice as wide as long,

with two denticles on inner edge. Merus expanded externally with seven denticles

along inner and distal edges. Carpus and propodus subequal, and each 1} times

length of merus. Dactyl small. Basal segment of exopod oval, and less than half

length of basis.

Pereiopod | (Fig. 7G) relatively short. Ischium as broad as long, and slightly

expanded internally, as is the merus. Merus twice length of ischium. Carpus

cylindrical, twice length of merus. Dactyl and part of propodus missing. Exopod

very well developed, basal segment 2 length of basis.

Pereiopod 2 (Fig. 7H) largest of the legs, nearly 24 times length of basis of

pereiopod 1. Basis slightly keeled to accommodate the large exopod, which has

basal segment 3 length of basis, flattened, with a row of 9-10 denticles along the

outer and distal edges. Ischium half as long as broad. Merus three times length

of ischium. Carpus cylindrical, equal in length to ischium plus merus. Propodus

small, just more than } length of carpus. Dactyl elongate, equal in length to

ischium, merus and carpus together, with a small terminal spine and five lateral

spines on inner edge, interspersed with small denticles.

Pereiopod 3 (Fig. 71) smaller, reaching end of carpus of pereiopod 2. Basal

segment of exopod smaller and less expanded, slightly more than half length of

basis. Basis long and straight, 14 times length of rest of limb, with a row of

denticles along its length. Ischium square, merus twice its length. Carpus

200 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 8. Hypocuma dentatum sp. nov.

Adult female, paratype, carapace length 2,0 mm: A, lateral view of cephalothorax; B, dorsal

view of cephalothorax; C, antenna 1; D, maxilliped 2; E, maxilliped 3.

elongate, twice length of ischium plus merus. Propodus and dactyl small,

subequal.

Pereiopod 4 (Fig. 7J) smaller still, with basal segment of exopod ? length of

basis. Ischium square, merus slightly longer. Rest of limb missing. Pereiopod 5

missing.

Five pairs of pleopods (Fig. 7K), each with a sharp external projection on

inner ramus.

Telsonic somite (Figs 7L, 7M) produced between insertion of uropods for

3 its length, unarmed. Uropods missing.

Adult female, paratype, length of carapace 2,0 mm. Last thoracic segment

and abdomen missing in both specimens. Only bases of pereiopods retained.

Carapace similar to male, but deeper, about 1} times as long as deep

(Fig. 8A), with denticles on anterior edge. Pseudorostrum just less than a

quarter length of carapace, and more upturned than in male. No eye (Fig. 8B).

Antenna | (Fig. 8C) less elongate. First segment longest, second and third

subequal and together about ? length of first. Accessory flagellum with minute

second segment. Flagellum l-segmented, and nearly as long as basal segment.

Maxilliped 2 (Fig. 8D) with merus more expanded internally than in male.

Maxilliped 3 (Fig. 8E) with exopod #2 length of basis. Merus expanded

internally, but not as elongated as in male.

Pereiopods represented by bases only, that of pereiopod 2 largest. Articula-

tion of exopod of pereiopod 3 visible, but exopod missing.

SOUTH AFRICAN CUMACEA: PART | 201

Remarks

H. dentatum quite obviously belongs to the genus Hypocuma, and the

differences between it and H. serratifrons are not great. The two species may be

distinguished as follows: in H. dentatum the exoskeleton is covered by scattered

denticles of varying size, those on the anterior edge of the carapace being

particularly distinctive, and larger than in H. serratifrons. The general shape of

the carapace differs, mainly in the lack of a depression behind the pseudorostrum

in H. dentatum and the nature and degree of tilting of the pseudorostral lobes.

The telsonic somite protrudes between the uropods for less than 4 its length in

H. dentatum, and for nearly half in H. serratifrons. The flagellum of the first

antenna in H. serratifrons is 2-segmented in both the male and the female, and in

H. dentatum is apparently 1-segmented in the female, and 3-segmented in the

male, but since the specimens of H. dentatum are very old and quite decalcified,

it is difficult to be certain about the exact position of some sutures.

Distribution of Hypocuma

The single sample of H. dentatum was obtained from about 400 m off Cape

Point. No substrate data are given. The very considerable development of the

exopods of the male thoracic appendages indicates that it is an active swimmer.

Further collecting off the south and east coasts of southern Africa is needed to

determine the distribution range. H. serratifrons Jones is also represented by a

single sample, from 1 934 m off the Canary Islands.

Vaunthompsonia Bate, 1858

Generic diagnosis

Pseudorostrum short, eyes present. All pedigerous somites visible from

above. Telsonic somite slightly produced between uropods. Antenna | short

with l-segmented accessory flagellum. Basis of maxilliped 3 not produced

distally. Exopods present on pereiopods 1-4 in male and 1-3 in female. Male

with five pairs of pleopods. Endopod of uropod 2-segmented.

Type species Vaunthompsonia cristata (Bate, 1856), British Isles, Mediterranean,

West Indies, Indo-Pacific, South Africa.

KEY TO THE SPECIES OF VAUNTHOMPSONIA

iEcauncle of Uropods longer than telsonic somite. = .sss0c a+ 2--e22s456-- 0-20 ee ae oe 2

— Peduncle of uropods shorter than or equal to telsonic somite...............0 ee eee eee 3

2 Pereiopod 1 reaching end of carapace with part of propodus; dorsal outline not strongly

EBT He meme rere AR elie ae ite APON ACR cone meer clas Seen a V. cristata (Bate, 1856) —widespread

— Pereiopod 1 reaching end of carapace with dactyl only; dorsal outline strongly arched....

V. arabica Calman, 1907—India

Seantero-lateral borders: Of carapace settated!. =. a. ciclo te woe cee ee ob oc «nme eels eee 4

—Antero-lateral borders of carapace not serrated. ........6.5---0 ese s cee een acre n ees 6

4 Minute serrations on antero-lateral border only................ V. sp.—South Africa

— Serrations present on carapace apart from antero-lateral border..........-..-+sesee+: 5

202 ANNALS OF THE SOUTH AFRICAN MUSEUM

5 Single row of mid-dorsal serrations on carapace...... V. serratifrons Gamo, 1964—Japan

— A double row of serrations on posterior half of carapace, converging towards posterior

DORGED eee tate tae ye ne! Sen na V. meridionalis G. O. Sars, 1887— Antarctica

6 Basis of maxilliped 3 somewhat produced distally...V. inermis Zimmer, 1909 — Antarctica

= Basis, of maxilliped:3 not produced si...4 255 hb. s sco bs» he © a cet ae 7

7 Length of telsonic somite anterior to insertion of peduncle greater than length posterior to

ATUS@ HU OM (ice iaes ia: wer atte alco. airnutcbrn rs erie ou ser fren Ars ras Ua eleeRs ah oS SE St ne ne nn 8

— Length of telsonic somite anterior to insertion of peduncle shorter than or equal to length

POStETION TOMMSETUOME 6.64. Daeaiaie ata Oa Re hae eo UES waked elses eee ee 9

First segment of exopod almost equal in length to second of endopod..................

V. natalensis sp. nov.—South Africa

— First segment of exopod half length of second segment of endopod....................

V. media Zimmer, 1952—Indochina

9 Pereiopod 1 with basal segment of exopod # length of basis; first two segments of antenna 1

longer than rest-of appendage yoy. 60. oles oe cee and cas Com ata. oe 10

— Pereiopod 1 with basal segment of exopod 2 length of basis; first two segments of antenna 1

shorter thant rest of appendages 7 mine ace eee V. dawydoffi Zimmer, 1952—Indochina

10. First segment of exopod of uropod!; length of second. .....5....-2.5.. 0. eee

V. floridana Bacescu, 1971 — Caribbean

— First segment of exopod of uropod's length) ofisecond 4.2)... 40-4. 2) eee

V. nana Hale, 1944— Australia

Vaunthompsonia cristata (Bate, 1856)

Vaunthompsonia cristata: G. O. Sars, 1879: 22, pl. 23-26, figs 17-18. Stebbing, 1913: 10,

figs 5-7.

Records

One specimen from False Bay (34°S/18°E), depth 5 m

Holotype

Designated by Bate, 1856, from western France. ?British Museum (Natural

History).

Remarks

This species, represented by a single damaged male, was identified by

Jones (1960) from UCT material taken from False Bay. Dr Jones has kindly

re-examined the material, which is still in his possession, and confirms his

original identification.

Distribution

Widespread—British Isles, Mediterranean, Annam, Japan, West Indies,

South Africa, in shallow waters.

Vaunthompsonia natalensis sp. nov.

Figs 9 (2), 10 (g)

Records

SCD 272 Y 19.7.1961 34°23’S/25°5S4’°E Sand and shell 182m 17 @ ovig. 3,4mm

SCD 321 L .9.7.1962 34°15’S/25°50‘’E Fine sand 108m 1 subadult ¢ 3,0mm

Coast 4/P3 = 6.2.1973 —30°37’S/30°40’E Ts i.) lee 2,0 mm

Coast 4/Q2 6.2.1973 30°46’S/30°31’E 48m 198 1,8 mm

SOUTH AFRICAN CUMACEA: PART 1] 203

Fig. 9. Vaunthompsonia natalensis sp. nov.

Adult female, holotype, 3,4 mm: A, lateral view; B, detail of anterior end of

carapace; C, dorsal view of carapace; D, antenna 1; E, maxilliped 3; F, pereio-

pod 1; G, pereiopod 2; H, pereiopod 3; I, pereiopod 4; J, pereiopod 5; K, telsonic

somite and uropod of adult; L, telsonic somite and uropod of juvenile.

Holotype

Ovigerous female deposited in the South African Museum, number

SAM A13435.

Description

Ovigerous female, holotype, length 3,4 mm. Body compact, smooth,

without serrations (Fig. 9A). Carapace about 3 as deep as long, somewhat

vaulted anteriorly with faint mid-dorsal carina on eyelobe. Antennal notch

shallow, antero-lateral angle defined by a small tooth (Fig. 9B). Eye obvious,

heart-shaped in dorsal view, with no visible lenses (Fig. 9C). Pseudorostrum

meeting in front of the rounded eyelobe for a short distance. Carapace a quarter

204 ANNALS OF THE SOUTH AFRICAN MUSEUM

as long again as free thoracic somites, all of which are visible dorsally. Cephalo-

thorax as long as abdomen excluding uropods. Abdominal somites cylindrical,

each with a small posterior articulatory process. Telsonic somite produced

between uropods for half its length, with several fine spines between uropods,

and two fine anal setae (minutely serrated between uropods in juveniles).

Antenna | (Fig. 9D) with basal segment longest, and second segment

shortest. Flagellum 2-segmented with two terminal setae. Accessory flagellum

1-segmented.

Basal segment of maxilliped 3 (Fig. 9E) not produced. Ischium twice as

broad as long. Merus somewhat expanded distally, and slightly longer than

broad. Carpus widened distally, subequal to merus. Propodus and dactyl

nearly subequal.

Pereiopod | (Fig. 9F) fairly elongate, with dactyl and part of propodus

extending beyond carapace anteriorly. Basis equal in length to next four seg-

ments. Ischium small, square. Next four segments almost subequal. Basal

segment of exopod half length of basis.

Pereiopod 2 (Fig. 9G) relatively stout, 7-segmented. Ischium 4 as long as

wide, merus and carpus subequal. Propodus 4 length of dactyl, which bears a

number of spines. Basal segment of exopod less than half length of basis.

Pereiopods 3-5 (Figs 9H-J) similar, except for exopod of pereiopod 3.

Basis of pereiopod 5 longest.

Peduncle of uropod stout, shorter than telsonic somite (Fig. 9K) with seven

sharp setae on inner edge; juveniles with three spines on minutely serrated inner

edge (Fig. 9L). Rami of juveniles subequal. First segment of endopod of adult

twice length of second with ten small spines on inner edge; juveniles with three

spines on serrated inner edge. Second segment of adult with six small spines on

inner edge (juveniles unarmed) and two terminal spines in all three specimens.

Exopod of juveniles with first segment ? length of second, unarmed. Second

segment missing in holotype; minutely serrated on inner edge of juveniles, with

two short spines and one terminally.

Subadult male length 3,0 mm. Differs from the female as follows: generally

more robust; carapace slightly less deep (Fig. 10A). Eye with a ring of 11 lenses

around edge of pigmented area (Fig. 10B).

First segment of flagellum of antenna | longer than second (Fig. 10C).

Merus of maxilliped 3 (Fig. 10E) slightly longer and broader. Carpus not

expanded.

Last five segments of pereiopod | missing.

Five pairs of pleopods present (Fig. 10H).

Shape of telsonic somite as in female, but unserrated posteriorly (Fig. 101).

Peduncle of uropod slightly longer, with nine spines on inner edge. First segment

of exopod unarmed, half length of second; second bearing four small spines on

outer and three on inner edge, and three terminally. First segment of endopod

twice length of second, with 12 sharp spines on inner edge; second with five small

spines on inner edge, and three terminally.

SOUTH AFRICAN CUMACEA: PART | 205

Fig. 10. Vaunthompsonia natalensis sp. nov.

Subadult male, 3,0 mm: A, lateral view; B, dorsal view of carapace; C, antenna 1;

D, detail of distal end of antenna 1; E, maxilliped 3; F, pereiopod 2; G, pereio-

pod 4; H, pleopod 3; I, telsonic somite and uropod.

Remarks

This species conforms to the general facies of the genus. It differs from the

other unserrated members as follows: the carapace of the female is somewhat

deepened midway along its length; the eye is heart-shaped in dorsal view; the

merus of maxilliped 3 is slightly broadened distally, particularly in the female.

The exopods of the pereiopods are smaller and narrower than those of other

species, and the peduncles and rami of the uropods are particularly stout. The

proportions of the uropodal segments are distinctive, especially in that the first

segment of the exopod and the second of the endopod are subequal.

Distribution

Endemic to Natal from 45 to 182 m.

206 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 11. Vaunthompsonia sp.

Adult female, 4,4 mm: A, lateral view; B, detail of anterior end of carapace;

C, dorsal view of carapace; D, detail of anterior end of carapace in dorsal view;

E, antenna 1; F, maxilliped 3; G, pereiopod 1; H, pereiopod 2; I, pereiopod 3;

J, pereiopod 4; K, pereiopod 5; L, telsonic somite and uropod.

Vaunthompsonia sp.

Fig. 11

Records

WCD 95 V 3.71961 33°06’S/17°49’E 88 m Mud and rock ie 4,4 mm

Description

Adult female, length 4,4 mm. In most respects (Figs 11A—D, F, H-K)

similar to V. media Zimmer, 1952, from Annam, but differs in the length of the

basal segment of antenna | (Fig. 11E), in the relative lengths of the merus and

SOUTH AFRICAN CUMACEA: PART | 207

carpus of pereiopod | (Fig. 11G) and the telsonic somite and uropods (Fig. 11L),

described below.

Telsonic somite 3 as wide as long at its widest point, and produced between

uropods for half its length, terminating in two small anal setae. Ratio of outer

edge of telsonic somite: outer edge of peduncle is | : 2. Peduncle with nine spines

on inner edge. First segment of endopod slightly more than half length of

peduncle, with 12 fine spinules along inner edge. Second segment missing. First

segment of exovod ? as long as first segment of endopod, unarmed. Second

segment just shorter than first, with four small spines near the end.

Remarks

Since neither of the uropods of the single female is complete, and since

Zimmer described V. media on the basis of adult males, certain identification or

the erection of a new species should await further material. The specimen differs

from V. natalensis in the shape of the carapace, in the basis and carpus of

maxilliped 3, in the presence of serrations on the lower border of the carapace

and in particular in the proportions of the telsonic somite, and the peduncle and

rami of the uropods.

Distribution of Vaunthompsonia

Vaunthompsonia has the widest distribution of any of the genera under

consideration. Species occur from the British Isles (V. cristata) to South Georgia

and Kerguelen (V. meridionalis and V. inermis), Japan (V. serratifrons) and

Australia (V. nana). The genus is generally confined to waters shallower than

250 m, but V. meridionalis has been found at 315 m in South Georgia.

Bathycuma Hansen, 1895

Generic diagnosis

Pseudorostral lobes meeting anteriorly. Eye absent. First pedigerous somite

exposed. Telsonic somite produced between uropods. Basis of maxilliped 3

produced distally. Exopods present on pereiopods 1-4 in male, and 1-3 in

female. Male with five pairs of pleopods. Endopod of uropod 2-segmented.

Type species B. elongatum Hansen, 1895, from 4 980 m in the central Atlantic. ©

KEY TO THE SPECIES OF BATH YCUMA

IBC ATA ACTON |) COT SOIMULEES ssi coo tel <2 Saves shoved See ohonene. & WS rw & dun ave, 0 uilein goin nye ee Bie ome 4

= ELST SOPNTES THOR ta ey rer ee a ee renee eerie 3

2 Distal prolongation of basis of maxilliped 3 not reaching end of merus.............-.-.

B. longicaudatum Calman, 1912—California

— Distal prolongation of basis of maxilliped 3 reaching beyond end of merus.............

B. magnum Jones, 1969—Indian Ocean

Beectciopod 2 with six segments...) .5...03....2s.5- B. elongatum Hansen, 1895—Atlantic

EEC OPOM I ow lily SEVEN *SEPINEMIS iis ae a ieis aiete cls. aerate, Siuste a clese wb e slide wigis 6 eye bie = = 4

208 ANNALS OF THE SOUTH AFRICAN MUSEUM

A Segments of endopod of uropod not subequal: ....2:.....2.495.- 2 seek 5

= Segments of subequal endopod of uropod’.::..2......4 4.200545) soe 6

5 First segment of endopod of uropod twice length of second; prolongation of basis of

maxilliped 3 reaching middle of merus........ B. capense (Zimmer, 1921)—South Africa

— First segment of endopod of uropod shorter than second; prolongation of basis of maxilliped

3 reaching end Ofmenisas tae oe eee ee nee B. brevirostre (Norman, 1879)—Europe

6 Carapace without mid-dorsal serrations; segments of exopod of uropod subequal........

B. datum sp. nov.—South Africa

— Carapace with mid-dorsal serrations; first segment of exopod of uropod half length of

SOCOM 5 o.5ehavsileite leile'ss toe: Svs Suan toner d aye lane a oraneveneee eben ks: © ARONA ciate tence ee Z

7 Uropods more than twice length of telsonic somite; first segment of endopod distinctly

longerthanlSecondianas serine B. natalense Stebbing, 1912—South Africa

— Uropods less than half length of telsonic somite; first segment of endopod just longer than

SECON SE Mee de ns, seme Caner eat rte B. longirostre Calman, 1905— Malaya

Bathycuma capense (Zimmer, 1921)

Figs 12 (2), 13A—D (3)

Vaunthompsonia capensis Zimmer, 1921: 131, figs 28-29.

Records

SAM A10602c, Pieter Faure 17440: 34°20’S/18°50’E about 400 m 1¢

7,2 mm

WCD 77 A 8.10.1960 34°17’S/17°53’E 320m Sandand 1 2 adult

mud 10,9 mm

1 ¢ subadult

10,2 mm

SCD 405 E ? 34°10’S/20°12’E about 200m Sand 49

8,6-10,2 mm

2 3 subadult

9,9-10,2 mm

1 juv.

4,6 mm

Holotype designated by Zimmer (1921) from Agulhas Bank 35°19’S/20°12’E

140 m. Zoologisches Museum, Berlin.

Description

Adult female, length 10,2 mm. Body elongate, carapace short, vaulted, a

little more than 14 times as long as deep (Fig. 12A), mid-dorsal carina bearing

numerous small teeth anteriorly. Pseudorostral lobes meeting in front of very

small eyeless eyelobe for 3/5 length of carapace (Fig. 12C). Antennal notch not

deep, but dorso-ventrally excavated with several minute teeth above (Fig. 12B).

Antero-lateral angle acute, defined by a large tooth with several small serrations

along the ventro-lateral margin. Carapace just longer than rest of thorax.

Pedigerous somites all exposed, second, third and fourth with slight flanges

visible dorsally. Cephalothorax just shorter than next five abdominal somites

which are cylindrical in cross-section. Telsonic somite produced between uropods

for 4 its length.

SOUTH AFRICAN CUMACEA: PART 1 209

Fig. 12. Bathycuma capense (Zimmer, 1921)

Adult female, 10,2 mm: A, lateral view; B, detail of anterior end of carapace; C, dorsal view -

of carapace; D, antenna 1; E, detail of distal end of antenna 1; F, maxilliped 3; G, pereiopod 1;

H, pereiopod 2; I, pereiopod 3; J, telsonic somite and uropod.

Antenna | (Fig. 12D) robust, first segment geniculate and longer than next

two together. Second segment just shorter than third. Flagellum (Fig. 12E)

3-segmented, nearly as long as second basal segment, with two terminal setae.

2-segmented accessory flagellum half as long as first of flagellar segments, with

two plumose terminal setae.

Maxilliped 3 (Fig. 12F) setiferous. Basis more than twice length of remaining

segments, with a distal prolongation reaching half way along merus. Other seg-

210 ANNALS OF THE SOUTH AFRICAN MUSEUM

ments subequal, ischium slightly longest and stoutest. Basal segment of exopod

more than quarter length of basis.

Pereiopod | (Fig. 12G) elongate, with propodus and dactyl exceeding tip of

pseudorostrum. Basis setiferous. Ischium broader than long. Carpus equal to

ischium plus merus. Propodus and dactyl long and slender. Basal segment of

exopod # length of basis.

Pereiopod 2 (Fig. 12H) fairly stout, basis nearly as long as last four seg-

ments. Ischium distinct but very short. Merus and carpus subequal, stout, each

with a strong distal spine. Propodus 3 length of merus. Dactyl relatively short,

twice length of propodus, with a few small spines. Basal segment of exopod 2

length of basis.

Pereiopod 3 (Fig. 121) more slender. Basis longer than remaining segments

together, of which carpus is longest. Exopod as in pereiopod 2.

Peduncle of uropods shorter than telsonic somite (Fig. 12J) with nine

uneven spines along inner edge. Exopod just longer than endopod and just

shorter than peduncle. First segment of exopod less than half length of second,

unarmed. Second segment with four small spines on both edges, and two

terminally. First segment of endopod more than twice length of second, with

seven small spines on inner edge. Second segment with four spines on inner edge

and two terminally.

Subadult male, length 10,2 mm. As in female, except: carapace more

rectangular in outline; pseudorostral lobes + total length of carapace (Fig. 13A).

Antennal notch much shallower, angle obtuse with finer teeth below (Fig. 13B).

Length of cephalothorax somewhat less than length of next fiveabdominal somites.

Antenna | (Fig. 13C) with first segment not geniculate, accessory flagellum

longer, flagellum 2-segmented, terminating in two aesthetascs.

Thoracic appendages as in female, but pereiopod | longer, with carpus

exceeding tip of pseudorostrum.

Armature of uropods more extensive (Fig. 13D).

Remarks

Zimmer (1921) described B. capense as Vaunthompsonia capensis from a

single subadult male, figuring only the entire animal and a uropod. Although he

did not accept the genus Bathycuma, considering it at most to be a subgenus of

Vaunthompsonia, it is quite obvious that Zimmer’s specimen belongs to Bathy-

cuma as now accepted. I am unable to find any significant differences between

Zimmer’s description and the present material.

Bathycuma natalense Stebbing, 1912

Figs 13E-J (¢), 14 (9)

Bathycuma natalense Stebbing, 1912: 135, pl. 49.

Records

SST 66C. 21.7.1972 34°23’S/21°26’E 15m Sandandshell 1Qadult 9,0mm

1Q 6,5 mm

Ld 6,1 mm

SOUTH AFRICAN CUMACEA: PART | 214

Fig. 13. Bathycuma capense (Zimmer, 1921)

Subadult male, 10,2 mm: A, lateral view; B, anterior end of carapace; C, antenna 1; D, telsonic

somite and uropod.

Bathycuma natalense Stebbing, 1912

Adult male, 11,2 mm: E, lateral view; F, detail of anterior end of carapace; G, dorsal view of

carapace; H, antenna 1; I, detail of distal segments of antenna 1; J, telsonic somite and uropod.

Db ANNALS OF THE SOUTH AFRICAN MUSEUM

Holotype designated by Stebbing (1912): Pieter Faure 12605, 30°10’S/31°03’E

410 m, 2 adult males. British Museum (Natural History).

Description

Adult female, length 9,0 mm. Carapace elongate, less than twice as long as

broad, serrated mid-dorsally along eyelobe (Fig. 14A). Pseudorostrum relatively

long, meeting in front of small triangular eyelobe for ¢ of total length of carapace

(Fig. 14E). Antennal notch (Fig. 14B) semicircular, fairly small but well-

defined by a large antero-lateral tooth with a few serrations running along

ventro-lateral border. Carapace longer than free thoracic somites which are

slightly flanged laterally. Cephalothorax just longer than next five abdominal

somites, which are cylindrical in cross-section and lack carinae. Telsonic somite

produced between uropods for nearly 4 its length.

Antenna | (Fig. 14C) fairly long and slender. Basal segment geniculate,

next two subequal, each 3 length of basal segment. Flagellum (Fig. 14D)

2-segmented, first twice length of second, which bears two aesthetascs. Accessory

flagellum 2-segmented and 3 length of first flagellar segment.

Maxilliped 3 (Figs 14F and 14G) with basis three times length of remaining

segments. Distal prolongation almost reaching end of merus, with four small

denticles on inner edge just below insertion of ischium. Ischium just longer than

wide with single denticle on inner distal edge. Merus slightly longer than

ischium, somewhat expanded externally with two denticles at apex of expansion.

Carpus and merus subequal, propodus and dacty! subequal, shorter than carpus.

Basal segment of exopod less than 4 length of basis.

Basis of pereiopod | (Fig. 14H) long and slender. Ischium short with a single

spine on slightly expanded inner edge. Merus twice length of ischium. Carpus

and rest of limb missing in this and all other specimens. Basal segment of

exopod § length of basis.

Basis of pereiopod 2 (Fig. 141) equal in length to last four segments.

Ischium as long as broad, merus 23 times length of ischium. Carpus equal to

ischium plus merus. Dactyl fairly stout and equal in length to carpus. Basal

segment of exopod more than half length of basis.

Basis of pereiopod 3 (Fig. 14J) longer than rest of leg. Ischium slightly

broader than long, merus 1} times length of ischium. Carpus longer than

ischium and merus together, and twice length of propodus and dactyl together.

Basal segment of exopod ? length of basis.

Pereiopods 4 and 5 without exopods.

Peduncle of uropod longer than telsonic somite (Fig. 14K), with about 13

spines arranged in two rows along inner edge. Exopod } length of peduncle,

first segment unarmed, half length of second, which has eight small spines on

outer and five on inner edge, and three terminally. First segment of endopod

slightly longer than first segment of exopod, with four spines on inner edge.

Second segment missing.

Adult male, length 11,2 mm. As the male is in poor condition, Stebbing’s

SOUTH AFRICAN CUMACEA: PART 1 213

Fig. 14. Bathycuma natalense Stebbing, 1912.

Adult female, 9,0 mm: A, lateral view; B, detail of anterior end of carapace; C, antenna 1;

D, detail of distal end of antenna 1; E, dorsal view of carapace; F, maxilliped 3; G, distal

segments of maxilliped 3; H, pereiopod 1; I, pereiopod 2; J, pereiopod 3; K, telsonic somite

and uropod.

type is redescribed where it differs from the female. The whole animal (Figs 13E

and G) is refigured, as the type specimens do not correspond well with his

original figures.

Carapace as in female, but lacking antennal notch and antero-lateral angle

(Fig. 13E). Twice as long as deep, and more vaulted anteriorly with serrations on

ventral edge of pseudorostral lobes.

214 ANNALS OF THE SOUTH AFRICAN MUSEUM

Antenna | (Fig. 13H) unusual, extremely robust, with numerous setae on

the accessory flagellum (Fig. 131). Basal segment geniculate. Flagellum

3-segmented, first segment small with a number of setae; second twice length of

first, third just longer than first with a single terminal spine. Accessory flagellum

1-segmented with numerous long setae.

Pereiopods as in female.

Telsonic somite (Fig. 13K) with minute serrations laterally, posterior to

insertion of uropods. Uropods and first segment of endopod relatively longer.

Remarks

In most respects, this species is similar to other members of the genus, but is

distinguished by the greater length of the peduncle of the uropods, and the very

robust first antennae of the male. B. natalense and B. capense are similar to each

other in general appearance, and are probably closely related. In addition to the

differences in the first antennae of the adult males, however, they are also

distinguished by the relative lengths of the uropods and the different shapes of

the carapace.

The depth of the type specimen (about 400 m) is fairly typical for the genus,

but the shallow depth of 15 m for the SST sample seems inexplicable. Records,

logs and labels have been checked and seem to be correct. The greatest depth

reached in this particular transect was 200 m, so that even had a labelling error

occurred, the depth is surprisingly shallow. Further collecting is required to

determine whether in fact B. natalense does normally occur at such shallow depths.

Bathycuma datum sp. nov.

lee JS)

Records

SAM A10602b, Pieter Faure 17440: 34°25’S/18°50’E 400 m Pe 7,7 mm

Holotype

Unique adult female, deposited in the South African Museum, number

SAM A13436.

Description

Adult female, \ength 7,7 mm. Body smooth, integument very finely

squamous, Dorsal outline of carapace gently arched, not carinate or serrate

(Fig. 15A). Pseudorostrum very short, meeting just in front of minute, triangular

eyeless eyelobe (Fig. 15C). Antennal notch relatively deep, antero-lateral angle

with a small tooth (Fig. 15B). Carapace 2 as deep as long, and just longer than

remaining free thoracic somites. Terga of second, third and fourth pedigerous

somites slightly elevated, the second most obviously. Cephalothorax just shorter

than abdomen. Telsonic somite only slightly produced between uropods.

Antenna | (Fig. 15D) short, basal segment robust, ? as broad as long with a

row of six small denticles along outer edge. Second segment + length of first, and

broader than long. Third segment nearly twice length of second. Flagellum

SOUTH AFRICAN CUMACEA: PART | 215

Fig. 15. Bathycuma datum sp. nov.

Adult female, holotype, 7,7 mm: A, lateral view; B, detail of anterior end of

carapace; C, dorsal view of carapace; D, antenna 1; E, maxilliped 3; F, pereiopod

1; G, pereiopod 2; H, pereiopod 3; I, telsonic somite and uropod.

l-segmented with two terminal aesthetascs. Accessory flagellum 1-segmented,

and half as long as flagellum.

Mandible normal, with narrow base.

Basis of maxilliped 3 (Fig. 15E) (including distal prolongation) twice length

of remaining segments. Prolongation of basis narrow, extending not quite to

distal end of merus. Ischium nearly twice as long as broad, merus slightly longer

than ischium, slightly expanded internally. Basal segment of exopod less than

3 length of basis.

216 ANNALS OF THE SOUTH AFRICAN MUSEUM

Pereiopod | (Fig. 15F) elongated, basis just shorter than remaining seg-

ments. Ischium square with two small denticles distally. Merus twice length of

ischium. Carpus equal to ischium plus merus. Propodus and dactyl subequal.

Basal segment of exopod less than 4 length of basis.

Basis of pereiopod 2 (Fig. 15G) just longer than next four segments.

Ischium 1% times as broad as long, merus twice this length. Merus and carpus

subequal, propodus small. Dactyl almost as long as merus plus carpus, seti-

ferous, with four very stout terminal spines. Basal segment of exopod more than

half length of basis.

Pereiopod 3 (Fig. 15H) with basis longer than remaining segments. Ischium

square, merus twice its length. Propodus and dactyl almost subequal, each 2

length of carpus. Basal segment of exopod less than half length of basis.

Pereiopods 4 and 5 similar, but basis of pereiopod 4 longer.

Peduncle of uropods (Fig. 151) 15 times length of telsonic somite and just

longer than exopod, with seven evenly spaced spines along inner edge. Endopod

just shorter than exopod, segments subequal, first with six fine spines on inner

edge, second with six small spines on inner edge and two long terminal spines.

First segment of exopod $ length of second, unarmed. Second with four plumose

setae and one spine on inner edge, and two spines terminally.

Remarks

This species may be distinguished from others of the genus by the absence

of a serrated dorsal carina, the short, stout first antenna with a 1-segmented

flagellum, and the relatively short prolongation of the telsonic somite between

the uropods.

Distribution of Bathycuma

As the name Bathycuma indicates, most of the species in this genus inhabit

waters of a considerable depth—usually more than 1 000 m. The genus is quite

widely distributed in the North Atlantic, Mediterranean, Indian and Pacific

Oceans, generally being circumtropical, with certain elements from higher

latitudes. The three South African species would appear to have been derived

from the Indian Ocean group, rather than having spread from the North

Atlantic, since no species have been found on the west coast of Africa. All three

are endemic, B. capense to the west coast, B. natalense to the south and east

coasts, and B. datum to the Cape west of the Peninsula. The known depths at

which all three species occur are unusually shallow for the genus.

DISTRIBUTION OF THE VAUNTHOMPSONIINAE

In general the distribution of each species in the subfamily is rather narrow.

This is partly due to rather scanty collecting in many areas, but even taking this

into account, of the 65 species listed in Jones (1969) and the present work, only

one, Vaunthompsonia cristata, is found in the Atlantic, Indian and Pacific

Oceans, one in the Pacific and Indian Oceans (Heterocuma sarsi), one in the

SOUTH AFRICAN CUMACEA: PART | 217

Atlantic and Pacific Oceans (Cumopsis goodsiri), and one in the Indian and

Atlantic Oceans (Heterocuma africanum)—a total of less than 7°% of the species

in the subfamily occurring in more than one ocean.

Zimmer (1941) has called the Bodotriidae a ‘negatively amphipolar’

family. This is borne out by the fact that none of the Vaunthompsoniinae has

been found at latitudes greater than 70°. Table | details the distribution of the

subfamily. Each species may be represented more than once, as each record of a

species from widely differing areas has been included.

TABLE 1

Distribution of Vaunthompsoniinae according to depth and latitude. Data mainly from

Jones (1969).

Latitude Shore-S m 5-200 m 200-2 000 m >2 000 m Total number

N of 70°N — — — — —

70°N-50°N 1 4 — — 5

50°N-20°N 3 13 5 1 22

20°N-20°S 1 13 3 1 18

20°S-—50°S D 36 4 Z. 44

50°S-70°S _- 2 = 1 3

S of 70°S — — — — —

Total 7 (5 sp) 68 (47 sp) 12 (10 sp) 5 (5 sp) 92 (67 sp)

Only eight of the 92 records are from latitudes greater than 50°. The majority

of records are from 20° to 50° N and S, fewer being found in the tropics, which is

generally true for the Cumacea. The predominance of records in the south

temperate latitudes may be due to the very extensive work done by Hale in

Australia. 21 of the 44 records for these latitudes are his, and as few other areas

have been as thoroughly worked, the data are somewhat biased in this direction.

However the bias is offset to some extent by the fact that only one member of the

subfamily has been described from the whole of South America, where very little

collecting has been done. Assuming that more records will eventually come from

this region, it may be concluded that the Vaunthompsoniinae are a family of

temperate latitudes, predominating in the south.

Again, 81% of the records are from depths less than 200 m, indicating that

the subfamily belongs predominantly to the shelf fauna, with some elements,

particularly members of the genus Cumopsis, now occupying the infratidal

fringe, and some, particularly Bathycuma and Gaussicuma, the bathyal and

abyssal zones.

DISTRIBUTION OF THE SOUTH AFRICAN VAUNTHOMPSONIINAE

The southern African coast is washed on the western side by the cold

northward-flowing Benguela Current (surface 15°C, bottom 10°C), and the warm

Mocambique Current flowing southwards along the east coast (surface 25-27°C,

bottom 21°C). The southern coast has the fast-flowing warm Agulhas Current

running from north-east to south-west (surface 25°C, bottom 12-14°C), and a

218 ANNALS OF THE SOUTH AFRICAN MUSEUM

narrow counter-current close inshore. The Cape Peninsula marks the westward

extent of the warm tongues of Agulhas water which occasionally penetrate into

False Bay (surface 15°C, bottom about 12°C). The boundary between Agulhas

and Benguela water is not stationary, so that False Bay may also receive cold

Benguela water at other times of the year. Thus generally animals from the west

coast are cold-water forms, and those from the south and east coasts (including

False Bay), warm-water forms. It is frequently found that animals occurring in

fairly shallow water on the cold west coast inhabit deeper waters on the south

coast due to the increased temperature of the Agulhas water (Day et al. 1970).

Seven of the eleven species and subspecies of Vaunthompsoniinae in these

waters are confined to one or other coast, suggesting that their distribution may

be largely temperature-dependent.

The only inter- and infra-tidal species, Cumopsis robusta, which belongs to a

predominantly intertidal genus, has only been found in False Bay and on the west

coast, and is therefore endemic. The anatomically related Heterocuma is a shelf

genus. H. africanum intermedium occurs in fairly large numbers in False Bay

(maximum depth just less than 90 m) to a depth of 66 m, and has also been

found in 27 m on the west coast. The same species has also been recorded twice

from the south coast in 84 to 91 m—an example of shallower depth range on the

west coast. It has also been recorded from shallow waters off tropical West

Africa. It is thus a warm-water Atlantic form, and is the only species represented

by more than 12 specimens in the present collection. H. africanum africanum has

only been found off Natal in 43 m, as well as in the Indian Ocean and tropical

West African waters.

Pseudosympodomma africanum is endemic, being found at a depth of 370 m

off the Cape Peninsula, and also at 85 and 135 m off the east coast.

The Bathycuma/Hypocuma]Vaunthompsonia group is divided not only

taxonomically but also ecologically, Bathycuma generally occurring at very great

depths, the two species of Hypocuma at 400 and | 934 m and Vaunthompsonia at

less than 250 m. Bathycuma, represented by three species and 17 specimens, can

be divided into a group living in fairly deep water off the Cape (B. datum and

B. capense), and B. natalense with the rather peculiar depth distribution from

15 to 400 m off Natal. The depth range of all three species is rather shallow, since

Bathycuma is generally a typically bathyal genus. Hypocuma dentatum, repre-

sented by only four specimens in a single sample, appears to be a deep-water

endemic form. Although three species of Vaunthompsonia are present, they are

represented by only six specimens, so that numerically, members of the genus are

only scantily represented in these waters. The three species do not overlap in

range, and the distribution of each necessarily appears narrow due to the paucity

of specimens. V. natalensis has been found only on the south and east coasts,

V. cristata in False Bay, and V. sp. on the west coast.

Thus the fauna can be divided into three groups:

1. Cold-water forms occurring on the west coast only—Hypocuma dentatum,

Bathycuma datum and Vaunthompsonia sp.

SOUTH AFRICAN CUMACEA: PART | 219

2. Warm-water forms occurring on the south and east coasts only— Bathycuma

natalense, Vaunthompsonia natalensis and Heterocuma africanum africanum.

3. Forms occurring around the coast, or including False Bay in their range—

Pseudosympodomma africanum, Bathycuma capense, Vaunthompsonia cristata,

Heterocuma africanum intermedium and Cumopsis robusta.

The present collection contains at least two thousand specimens of the

Bodotriidae. All but 77 of these are Bodotriinae, representing an estimated

18-20 species, giving a specimen : species ratio of at least 100 : 1. The 77 Vaun-

thompsoniinae represent 11 species, giving a very high ratio of 7 : 1. Thus the

subfamily exhibits a very high diversity in these waters. There is also a high rate

of endemism. Of the ten named species and subspecies, only three—Vaun-

thompsonia cristata, Heterocuma africanum africanum and H. a. intermedium—

are not endemic. Thus 70°% of the species are endemic, with a link to the tropical

West African fauna in the form of the two subspecies of H. africanum, while

V. cristata is cosmopolitan.

SUMMARY

Nine species of the southern African members of the subfamily Vaun-

thompsoniinae are described and figured. Of these, Hypocuma dentatum,

Cumopsis robusta, Bathycuma datum and Vaunthompsonia natalensis are new

species. Heterocuma africanum africanum and H. a. intermedium are newly

designated subspecies. The females of Bathycuma capense, B. natalense and

Pseudosympodomma africanum are described for the first time, P. africanum also

being allocated to a different genus. A specimen of Vaunthompsonia is briefly

described and figured, but not named. It is found that the subfamily is represented

in southern Africa by ten species in six genera.

The general distribution of the Vaunthompsoniinae is discussed, and a

more detailed account is given of the distribution of its southern African

members. It is concluded that this subfamily in southern African waters has a

high rate of endemism and a high diversity, but a very low frequency of

occurrence.

ACKNOWLEDGEMENTS

I should like to thank Dr R. R. Given of the Allan Hancock Marine Founda-

tion, California, for his help, advice and encouragement; Dr M. Bacescu of the

Musée d’Histoire Naturelle, Bucharest, for providing specimens of Cumopsis

fagei and literature, and Dr N. S. Jones of the Marine Biological Station, Isle of

Man, for specimens of C. fagei. I am also very grateful to Dr Brian Kensley of

the South African Museum, Cape Town, for material and for tracing Pieter

Faure station data, and Mr Tim McClurg of the National Institute for Water

Research, Durban, for material. My special thanks go to Professor J. H. Day of

the Department of Zoology of the University of Cape Town, for discussions of

taxonomic problems, and for his constructive criticism of the manuscript.

220 ANNALS OF THE SOUTH AFRICAN MUSEUM

REFERENCES

BAcescu, M. 1956. Cumopsis fagei n. sp. Cumacé nouveau, provenant des eaux du littoral

Frangais de la Manche. Vie Milieu 7: 357-365.

BatTE, S. 1858. Jn: KINAHAN, J. R. On the genus Scorpionura (J. V. Thompson MSS.). Nat.

Hist. Rev. 5: 202-205.

CHRISTIE, N. D. & ALLEN, J. C. 1972. A self-contained diver-operated quantitative sampler for

investigating the macrofauna of soft substrates. Trans. R. Soc. S. Afr. 40: 299-307.

Day, J. H., Frecp, J. G. & PENRITH, M. J. 1970. The benthic fauna and fishes of False Bay,

South Africa. Trans. R. Soc. S. Afr. 39: 1-108.

Face, L. 1924. A propos d’une espéce nouvelle du genre Heterocuma. Bull. Mus. natn. Hist.

nat., Paris. (2) 30: 364-367.

Face, L. 1950. Sur un nouveau Cumacé de la céte occidentale d’Afrique. Eocuma cadenati

n. sp. Bull. Mus. natn. Hist. nat., Paris (2) 22: 450-452.

Face, L. 1951. Cumacés. Result. scient. Expéd. océanogr. Belge Eaux Cét. Afr. Atlant. Sud.

3(1): 1-9.

HANSEN, H. J. 1895. Isopoden, Cumaceen und Stomatopoden der Plankton-Expedition.

Ergebn. Plankton-Exped. 2: 1-105.

HALE, H. M. 1928. Australian Cumacea. Trans. R. Soc. S. Aust. 52: 31-48.

HAtg, H. M. 1944. Australian Cumacea. No 8. The family Bodotriidae. Trans. R. Soc. S. Aust.

68: 225-285.

HA tg, H. M. 1949. Australian Cumacea. No 15. The family Bodotriidae (continued). Rec. S.

Aust. Mus. 9: 107-125.

HALE, H. M. 1953. Two new Cumacea from South Africa. Trans. R. Soc. S. Aust. 76: 45—50.

Jones, N. S. 1956. Cumacea from the west coast of Africa. Atlantide Rep. 4: 183-212.

JONES, N. S. 1960. Cumacea from South Africa. Ann. Mag. nat. Hist. (13) 2: 171-180.

JONES, N. S. 1963. The marine fauna of New Zealand: Crustacea of the order Cumacea. Bull.

N. Z. Dep. scient. ind. Res. 152: 1-80.

JONES, N. S. 1969. The systematics and distribution of Cumacea from depths exceeding 200 m.

Galathea Rep. 10: 99-180.

Jones, N. S. 1973. Some new Cumacea from deep water in the Atlantic. Crustaceana. 25:

297-319.

KurIAN, C. V. 1954. Notes on the Cumacea (Sympoda) in the Zoological Survey of India.

Rec. Indian Mus. 52: 275-311.

Miers, E. J. 1879. On a collection of Crustacea made by Capt. H. C. St. John in the Corean

and Japanese seas. Proc. zool. Soc. Lond. 1879: 18-23.

SARs, G. O. 1878-9. Middelhavets Cumaceer. Arch. Math. Naturv. 3-4: 1-196.

STEBBING, T. R. R. 1910. Sympoda. Ann. S. Afr. Mus. 6: 409-419.

STEBBING, T. R. R. 1912. South African Crustacea. Part 6. The Sympoda. Ann. S. Afr. Mus.

10: 129-176.

STEBBING, T. R. R. 1913. Cumacea. Tierreich 39: 1-210.

ZIMMER, C. 1908. Die Cumaceen der ,,Deutschen Tiefsee-Expedition”. Wiss. Ergebn. dt.

Tiefsee-Exped. ‘Valdivia’ 8: 155-196.

ZIMMER, C. 1921. Mitteilung tiber Cumaceen des Berliner Zoologischen Museums. Mitt. zool.

Mus. Berl. 10: 115-149.

ZIMMER, C. 1941. Cumaceen. Bronn’s KI. Ordn. Tierreichs. 5 (1, Book 4): 1-222.

ZIMMER, C. 1952. Indochinische Cumaceen. Mitt. zool. Mus. Berl. 28: 5-35.

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REFERENCES

Harvard system (name and year) to be used: author’s name and year of publication given

in text; full references at the end of the article, arranged alphabetically by names, chronologi-

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Examples (note capitalization and punctuation)

BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris

88: 100-140.

FISCHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des

littorines. Archs Zool. exp. gén. 74: 627-634.

Kon, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee

region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320.

Konan, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the

Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51. Z

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L.

Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und

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Jennifer Day

SOUTH AFRICAN CUMACEA

PART 1

FAMILY BODOTRIIDAE,

SUBFAMILY VAUNTHOMPSONIINAE

eo 41C0

_ VOLUME 66 PART 10 FEBRUARY 1975

CAPE TOWN

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

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February 1975 Februarie

Part 10 #£Deel

A NEW SPECIES AND RECORDS OF SCUTELLIDIUM

(COPEPODA HARPACTICOIDEA) FROM SOUTH

AFRICA, AND A WORLD KEY TO THE GENUS

G. M. BRANCH

Cape Town Kaapstad

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

A NEW SPECIES AND RECORDS OF SCUTELLIDIUM (COPEPODA

HARPACTICOIDEA) FROM SOUTH AFRICA, AND A WORLD KEY

TO THE GENUS

G. M. BRANCH

Zoology Department, University of Cape Town

(With 24 figures)

[MS accepted I July 1974]

CONTENTS

PAGE

Introduction 5 : ; ; 5 : : 3 mel

Methods . : ; : : : : : : ee 74 |

Scutellidium macrosetum sp. nov. . ; : ‘ 3 ‘e221

S. ringueleti : ; : : : : : ; i Meo

Key to the world species of Scutellidium : 5 : by 510)

Annotations to key : : , : P , SAR

Summary . ; : . ; : ; : ae DOD

References : ; ? : : ; ; : 2282

INTRODUCTION

The genus Scutellidium Claus, 1866 now comprises 25 species, of which

S. sarsi and S. digitatum are of uncertain position (and are not included in the

key given here). These are all listed or described by Lang (1948, 1965), Vervoort

(1964), Hicks (1971) and Branch (1973).

The present collection, made at Kalk Bay in the Cape Peninsula of South

Africa (34°8’S/18°27’E), includes three species: S. patellarum (Branch 1973),

S. ringueleti and a new species, described below as S. macrosetum sp. nov.

METHODS

All material was obtained from alcohol washings of intertidal algae,

preserved in 70° alcohol and then dissected and mounted in a glycerine-

formalin mixture. Drawings are based on camera lucida projections. The follow- —

ing abbreviations have been used: Al and A2, Antenna | and 2. End., endo-

podite; Exp., expodite; PI-5 swimmings legs 1-5; Segmt., segment. Student’s

t test has been used to compare mean lengths, and where relevent, 95° con-

fidence limits have been given for mean values.

Scutellidium macrosetum sp. nov.

Female

Live material pale yellow-brown. Length (rostrum to furcae): 0,738 mm,

range: 0,690 to 0,780 mm based on 15 specimens. Body compressed dorso-

ventrally. Prosoma oval but truncated posteriorly (Fig. 1). Posterior corners

Ann. S. Afr. Mus. 66 (10), 1975: 221-232, 24 figs.

221

222 ANNALS OF THE SOUTH AFRICAN MUSEUM

of prosomal segments forming rounded epimera: those of the 3rd segment

expanded and about twice the size of the other segments (Fig. 2). Prosoma

naked. Urosome 5 segmented, the genital ‘double segment’ very swollen and

almost twice the width of the other segments (Fig. 3). Tufts of setae on the

posterior—lateral margins of all but the last segment. Furcal rami about as

long as wide, with two very long and four shorter setae. Rostrum (Fig. 4)

rounded and naked except for two pairs of slender filaments (which are

frequently invisible).

Al 9 segmented and stout: similar to that of S. armatus (Wiborg, 1964),

except that segment 4 is shorter and wider. Segment | stout and well developed;

segment 2 about 23 x length of segment 3; segments 4-7 short and broad,

telescoped together, making the setal arrangement difficult to determine

accurately (Fig. 5). The basal segment has a characteristic strongly plumose seta

(Fig. 5a), one of the readily identifiable features of the species. Some of the

setae on segment 2 are also setose. The relative lengths and armature of the

segments appear in Table |.

TABLE 1. Details of A,

Segment 1 2 8) 4 5 6 i 8 9 Total

Length pw 93) 198) 42 13 es 5-7) US 19 42 32

2 >Relative length 29 30 13 4 y ifs) 2 535 13 =100

Setae (IP a 8 esas) Lye vA 3 1 2 2 4 36

Length pu 60 80 23 34 23 38 11 26 295

6 pRelative length “20 27 dss 5S) nS 13 4 9 100

Setae 1 oe tol 9 S+A 2 1 2 5 36

A2 basipodite with a single seta (Fig. 6). End 1 naked except for terminal

spinules, end 2 with setae and spines (see Fig. 6). Exp. weakly segmented, with

7 setae, the terminal 2 naked and the others shortly plumose.

Mandible (Fig. 7) and maxilla (Fig. 8) almost identical to S. armatus.

Maxilla 2 (Fig. 9) with a single segmented syncoxa bearing a shortly plumose

seta. No basal endite or setae on the syncoxa (but as these are difficult to dissect

out intact they may have been lost); a small distal endite with two setae.

Maxilliped (Fig. 10) 3 jointed, the basal segment with a long seta, the

2nd joint not as expanded as in most other species, and the terminal joint with

a stout toothed claw and 3 setae.

P| (Fig. 11) exp. and end. very broadly expanded, exp. about same length

as end. segment 1. Exp. 1 nearly as broad as end. 1; exp. 2 about half the size,

with 2 long internal setae and one short external seta. Exp. 3 characteristically

extremely small, with 4 small penicillate setae and one very long seta. End.

1 with a large inner seta arising just below the midpoint, and a semi-elliptical

row of setules. End. 2 about half the length and with a single short inner seta.

End. 3 small, carrying 2 penicillate setae which conceal a small straight plumose

seta. Basis with a broad outer seta and a small curved inner seta.

P2 (Fig. 12): end. 2 almost as long as wide. A slender outer seta on the

basis. End. 3 with only 4 setae.

NEW SPECIES AND RECORDS OF SCUTELLIDIUM (COPEPODA HARPACTICOIDEA) 223

Zoo il

ee Zs

Figs 1-8. Scutellidium macrosetum sp. nov. 2

1. Dorsal view of °. 2. Epimera of segments 2, 3 and 4.

3. Ventral view of urosome. 4. Rostrum.

5. Antenna 1. 6. Antenna 2.

7. Mandible. 8. Maxilla 1.

Both scales indicate 100 pz.

ANNALS OF THE SOUTH AFRICAN MUSEUM

224

Figs 9-15. S. macrosetum °

ao)

iD)

(hs

= —

—

tot 8)

aif

Star

—

(ee

The scale indicates 100 wu.

NEW SPECIES AND RECORDS OF SCUTELLIDIUM (COPEPODA HARPACTICOIDEA) 225

P3 and P4 as in Figures 13 and 14.

P5 (Fig. 15) with a bilobed basoendopodite, inner lobe with 3 setae, outer

with one longer seta. Exp. with a characteristic terminal setae, which is very

swollen at the base and distally tapered and bipectinate. (The specific name

macrosetum is derived from this). In addition there are 4 marginal setae of

approximately the same length (in one animal the 3rd marginal was half the

length of the other setae), and six parallel rows of setules.

Male

Similar to 2 but smaller, length 0,645 mm, (range 0,615—0,675, based on

3 specimens). Urosome much more slender and clearly 5 segmented. First

segment with dorso-lateral lobes (Fig. 16) which project posteriorly, partly

covering P5 and P6. Segments 2-4 each overlap the subsequent segment

ventrally, forming a spinule fringed flange (Figs 16, 17).

Apart from Al, P5 and P6 all appendages very similar to those of the 9.

Al (Fig. 18) 8 segmented, the basal segment also bearing a strongly plumose

seta. Segment lengths and armature given in Table | and Figure 19a-c.

P5 (Fig. 20) basoendopodite not as well defined as in the female: inner

lobe small, with | seta flanked by two small setules. Outer lobe larger, with

1 seta and a row of setules. Exp. again with a large terminal seta which is

swollen at the base and bipectinate. Outer margin with 4 setae of roughly equal

length, and a fringe of setules. (This is the only species in the genus in which

the ¢ has 5 setae on P5 exp.)

P6 a conical projection (Fig. 21) with a single stout blunt seta and two

slender setae about twice the length.

Affinities

S. macrosetum is closely allied to S. armatus (Wiborg), the only member of

the genus with which it can be confused. The easiest distinguishing factor is that

S. macrosetum has 5 setae on PS in both sexes while S. armatus has 4. (The

possibility that S. armatus was incorrectly described and had simply lost one

seta, can be discounted as both Wiborg (1964) and Pallares (1969: pl. 1, figs

2, 11), describe both sexes as having only 4 setae on P35.)

Other differences are summarized below.

S. macrosetum S. armatus

1. Inner seta, P1 basis Slender and curved Broad and ‘knife-shaped’

2. Relative width of Pl End. ike il 2:1 (Pallares 1969)

ifpeelvexp. | 214: 1 (Wiborg 1964)

3. Relative length of Pl exp. 34:1 21

DAPivexp: 35

4. Shape of P5 2 terminal seta Base bulbous, terminal Broad and curved tapering

part tapering gradually

5. 2 P5 no. of setae 5 4

6. 3 P5 no. of setae 5 4

emrlvexp: 25, setae 3: broad inner, stubby 2: broad inner and stubby

outer and _ terminal outer (Pallares 1969)

long and slender.

8. P5 2. 6 parallel rows of setules present absent

226 ANNALS OF THE SOUTH AFRICAN MUSEUM

Figs 16-21. S. macrosetum 3

16, 17. Dorsal and ventral views of the urosome.

18. Antenna 1. 19a, b, c. Segments 3, 4, 5 of antenna 1.

ZO RS: 21 PG;

Scale indicates 100 », and applies to all the figures except Fig. 21 which is twice the size.

NEW SPECIES AND RECORDS OF SCUTELLIDIUM (COPEPODA HARPACTICOIDEA) 227

WAL

Figs 22-23

22. Scutellidium ringueleti 3 PS.

23. S. ringueleti 2 P2.

S. macrosetum can be distinguished by the following combination of

features: P5 exp. with 5 setae, the terminal seta swollen at the base and pectinate;

furcal rami as long as wide; PI end. 1 about 2 x length of end. 2; P2 end. 2

nearly twice as long as wide; Pl exp. about same length as Pi end. 1; basal

segment Al with a very strongly setose seta.

Material

Dissected holotype (2) and allotype (¢) and intact paratypes (15 29 and -

33S) have been deposited in the South African Museum, Cape Town. (Cata-

logue numbers S.A.M. A13599, A13600, A13601).

Scutellidium ringueleti Pallares, 1969

Material

55 99 (length 0,55-0,95 mm) and 14 3g (length 0,55-0,80 mm) collected

at low tide level on Gigartina stiriata and Bifurcaria brassicaeformis.

Known distribution

Ria Deseado, Patagonia.

228 ANNALS OF THE SOUTH AFRICAN MUSEUM

Frequency

5 (7 . Oamecie

Length mm

Fig 24. Size and colour variations in S. ringueleti, in relation to foodplant. Shading indicates

a purple coloration.

f: size distribution of 29 collected on Gigartina.

a—d: 2 colour variations. ae ish

h: size frequency of 3.

g: size distribution of 292 on Bifurcaria.

NEW SPECIES AND RECORDS OF SCUTELLIDIUM (COPEPODA HARPACTICOIDEA) 229

Discussion

The present material agrees almost exactly with the excellent description

given by Pallares (1969), except for a few minor particulars: the terminal seta

of ¢ P5 much longer—about 2,5 x length of the exp. (Fig. 22); 9 PS terminal

seta, two thirds length of adjacent seta; maxilliped end. | with a proximal

marginal spine (often not visible); outer seta on basis of P2 extending just

beyond tip of exp. (Fig. 23), but rather variable in length.

Many of these features are intermediate between those of S. ringueleti

and the closely allied S. spinatum from New Zealand. Both these species have

been described in detail and the differences between them are slight (see Hicks

1971: 92). The intermediate nature of the South African material makes it

even more difficult to separate the species, and it seems probable that

S. spinatum is synonymous with S. ringue/eti. A direct comparison of the material

is required.

Colour varieties

The South African material contains several colour varieties of S. ringueleti:

Light yellow brown (58%); brown with a deep purple strand across the last

free metasomal segment and the first urosomal segment (32°); brown with

the last 3 metasomal segments and the epimera deep purple (3%); entirely

purple (7%).

These are of interest because Monk (1941) has used similar colour variations

to recognize varieties and species. Scutellidium arthuri var. magna Monk, 1941

is distinguished from the typical form by its greater size (1,4 mm as compared

with 0,91 mm) and the presence of a deep purple band on the ‘first free segment

of the metasome and on the urosome.’ Lang (1965) considers the variety invalid.

Monk has also described a closely allied species S. purpurocincta, distinguished

from S. arthuri by its colour, ‘purple on the 3 free segments of the metasome’

(Monk 1941: 96), and the proportionally longer third segment of Al. Lang

(1965: 151) also doubts the validity of this species.

As in Monk’s case, the present purple varieties of S. ringueleti are sig-

nificantly larger than the ‘typical’ brown form: 0,85 + 0,02 compared with

0,69 + 0,04 (significantly different at p < 0,001).

Further samples of S. ringueleti show that the purple and brown varieties

were associated with different foodplants, respectively Gigartina stiriata (Rhodo-

phyta) and Bifurcaria brassicaeformis (Phaeophyta). Figure 24 illustrates the

relationship between copepod size and colour, associated with these two algae.

Clearly such colour and size variations are phenotypic and related to the food-

plant: they are valueless in distinguishing taxonomic varieties. Curiously, the

males displayed no such variations and were all small and light brown. This

parallels Monk’s case again, for he found no males corresponding to the

purple forms, S. arthuri magna or S. purpurocincta.

In the light of this the variety magna cannot be justified, and further doubt

is cast on the validity of S. purpurocincta.

230

20.

2) I

22,

ANNALS OF THE SOUTH AFRICAN MUSEUM

KEY TO THE WORLD SPECIES OF SCUTELLIDIUM

Furcal rami as long as wide or wider than long 4

Furcal rami longer than wide . ee er ae Oe oe 2

Exp. PS long: 4 x longer than wide. . ase” acs S. leaneniece (Brady, 1910)

Exp. P5 shorter: less than 3 x longer than wide i 3

Exp. PS with transverse rows of spinules; Al segment 2 1} 25 % length of segment 3

S. antarcticum (Lang, 1936)

Exp. P5 without transverse rows of spinules; Al segment 2 and 3 subequal

S. major (T. Scott, 1912)

External seta on P2 basis strongly increased in size: more than 3 length of exopod . 5

External seta on P2 basis, normal: less than}. length of exp... . . . . 13

Pi'end. 1 roushly same lencthvasiend’2 . =) = 5) \3)-- (2) ) ae 6

Pi end. 1 at least 14 x lengthofend.2. . 7

P2 seta on basis docs not reach tip of exp.; maxilliped end. ] with my marginal spines

and 3 rows of spinules. . . S. spinatum Hicks, 1971

P2 seta on basis extends beyond tip of ee: manilliped end. 1 with no marginal spines

and 1 row of spinules .. . SS. ringueleti: Pallares, 1969

Outer spines of P2 end. setiform, and ie to DEX length of segments bearing them

S. patellarum Branch, 1973

Outer spines of P2 end: short; nonmal 9. > =. se 8

+ Pl end: 1. abouts > Clength of.end..2 oo 3 % ~~ 2 oe ele 9

Pivends 132 0or morerc< lengthiotends 2.1.) sa. ee oe 10

P1 exp. 2 strongly swollen internally, seta arising from centre ‘of swelling

S. plumosum Brady, 1899

P1 exp. less strongly swollen, and seta arising distal to swelling

S. intermedium® (Nicholls, 1941)

. Plend.1, 3 x length ofend.2 . . . .+. . ... S.StrigosumPallatesee

Pivend..lee2,.<slengthyvotende2i.) Vw U5 Ween See eee ene ne - ot

ay le2 CNC: 21dS LON S*AS WAGE nn, | ice ke ee lenis aces arse ie ‘Ss. acer Poppe, 1884

P2 end. 2 longer than wide . . ore 12

. Al segments 2 and 3 similar in length (proportions 13: 11) Si pubpuroenete: Monk, 1941

Al segment 2 obviously longer than 3 (proportions about 17 : 11)

S. dentipes* Vervoort, 1964

2 Pivexp., same densth ase endl) =). ae Bie hie bar 1920)

Pl exp. 1 shorter thanend.1. . aie 14

P5 terminal seta much wider than other Sai, aad petoce wot a 15

PS temminaliseta notas above, .-'. a co ae eee 16

bereXps With 4 Setae iieiw woe gt.” Cue gee ae ee ee sf ar ene (Wiborg, 1964)

P5 exp. with S:setae gh a Sig ae Se

. P34 end. 2 with l-seta =. .- s 6 ws bw es 4). SS. Aippolytes (Kaoyer aa

P3—4 end. 2 with2 setae . . re Ce eh cy |

. P5 exp. with at least 3 terminal setae : Poe a) Ve en a 3 18

P5 exp. with 1 terminal seta (others marginal) WR ee ey: |

. P5 basoendopodite inner lobe with 3 setae . . . .. . Ss. idyoides (Brady, sai

P5 basoendopodite inner lobe with 2 setae

5 eS) exp. inserting with a broad base, internal and Stionell walls neatly parallel, Ne) in

exp. is elongate; Pl end. 2, inner seta arises nearer distal end of segment

S. cockburni Fairbridge, 1944

P5 exp. narrow at base and apex, margins convex and exp. oval; Pl end. 2 inner seta

arises nearer proximal end of segment . . . _§. australe (T. Scott, 1912)

P5 exp. with 2 short thick setae: 1 terminal and 1 external, and with marginal spinules

and transverse rows of spinules . . . . . . . . S. fuciculum (T. Scott, 1912)

P5 exp. with 5 distinct setae 21

P5 terminal seta short, <4 length of exp.: coxa “of maxilliped without a " spine

S. longicauda (Philippi, _

P5 terminal seta longer at least > 4 length exp.; exp. length < 3 x width .

P5 terminal seta about 4 length of decent seta, and swollen at base; coxa of ee

With-an internal: sping "aS act) a eee S. deseadensis® Pallares, 1966

NEW SPECIES AND RECORDS OF SCUTELLIDIUM (COPEPODA HARPACTICOIDEA) 231

P5 terminal seta elongate, same length as adjacent seta, not swollen at base; coxa of

maxilliped with internal spine. . Fp S. loureiroi® Jakobi, 1954

PS terminal seta elongate, slightly longer than adjacent seta, not swollen at base.

(Maxilliped undescribed.) No external seta at base of P2 end. (?) S. lamellipes®? Monk,

1941

ie Spelesmuities, spines. orsetaes =, 6 2 Nb a eh atte oe oe 2

Bxomeomvitn 4 spines or setae. . «fw. st a ee : 3

Prone swith oS Spines Or setae: 9. e. e ‘ 5 macr oon sp. nov.

Peeriecniaeieaimost 2 <lengthend:2 ° .. . . . ag. longicauda (Philippi, 1846)

Pi end. 1 not much longer thanend.2 . . eg Ss. intermedium (Nicholls, 1941)

3. Outer spines P2 exp. setiform and about 13 to 2 « length of the segments bearing them

S. patellarum Branch, 1973

Outer spines P2 exp. short. . PP) ee ee ey Nee meee 4

4. Pl end. | roughly same length as end. 2 PT TT PIAL Od ce Qa ot ee eee 5

Pmcieamuaticast sx Jensth of end: 2. 6.0. % Goon si aeshalhe, be os: 9

Meme @mpne- setae on P5'exp. lancet-shaped = 2. 8 3). woe) a eee 6

All setae on PS exp. normal . SOP a td A Ai |S ee em 1 4

6. Pl end. 1 almost same length as exp. 1 hs of oo ae eS. eustheum(Bnan, 1920)

Pmcidasnaoout ls < length ofexp. 1. . . << = ca 2 S.australe(X. Scott, 1912)

(zwes—dVend. 2with i seta... . ee) fee en Se ippalyress(Kioyer. 1862)

P3—-4 end. 2 with 2 setae bh Gs ania Bete 3 5: In MURR AOR 8 8

8. PS exp. with rows of small spinules at base:

P2 seta on basis not reaching beyond tip of exp. . . . . SS. spinatum' Hicks 1971

P5 exp. without rows of spinules at base;

P2 seta on basis reaches beyond tip of exp. . . 9. ringueleti* Pallares 1969

9. External seta on basis of P2 elongated, at least 4 lene ORexp)? Mey shane at eee 10

Peaetitavsera.on basis of P2short? 2 4 < #8 shiecs bee : 11

MOeertvend isabout 13 x length ofend2, . ... ... % S. pleco Brady. 1899

Bigendealabout 2 < length of end. 2 := >... . << .S. arthuri Poppe; 1884

Fagcndemabout 3 < lengthiofend.2 9... = #28 . 4S: eo Pallares, 1969

iret) 2 < length ofexp.2 .< =: .« . «. .« «> S. armatus (Wiborg, 1964)

P1 exp. 1 about same length asexp.2 . . . . . JS. deseadensis® Pallares, 1969

‘LS. lamellipes®’ Monk, 1941

ANNOTATIONS

1. As described above (p. 229) these two spp. may be synonymous.

2. In Vervoort’s (1964) key, S. plumosum is described with a normal

external seta on P2 basis, but Pallares (1969: pl. 5, fig. 1) shows it as strongly

developed. I have followed Pallares.

3. P2 of S. intermedium has not been described but as this species is very

closely allied to S. plumosum (Nicholls, 1941) I assume that the external seta

on P2 basis is similarly elongated (see note 2).

4. S. purpurocincta is possibly synonymous with S. arthuri (see discussion

p. 229). S. dentipes may also be synonymous with S. arthuri, as Vervoort

himself admits.

5. S. deseadensis, S. loureiroi, S. lamellipes are possibly synonymous.

Jakobi’s description is not adequate to make comparisons, and Monk’s

description incomplete. S. /amellipes can only be separated from S. deseadensis

by ‘having no spine on the second basis’ of P2 (Monk 1941: 97) but as Lang

(1965) points out, this statement is probably incorrect: in several species

232 ANNALS OF THE SOUTH AFRICAN MUSEUM

(including S. deseadensis) this spine lies in a higher plane than the exp. and can

be overlooked while focusing on the exp.

6. S. deseadensis and S. lamellipes are probably synonymous (see also

note 5). Vervoort (1964: 106, couplet 6) describes S. lamellipes as having P1

end. | and 2 of about the same length (which would separate the two species)

but this statement is doubtful, as Monk does not describe or figure this appen-

dage, and describes S. /amellipes as being ‘very similar to S. longicauda’.

SUMMARY

A description is given of Scutellidium macrosetum sp. nov., which is allied

to S. armatus but distinguishable by having five setae on P5 exp. A new record

and colour variations are described for S. ringueleti. A key is given to the known

species of Scutellidium.

REFERENCES

BRANCH, G. M. 1973. Scutellidium patellarum n. sp., a harpacticoid copepod associated with

Patella spp. in South Africa, and a description of its larval development. Crustaceana

26: 179-200.

Hicks, G. R. E. 1971. Some littoral harpacticoid copepods, including five new species from

Wellington, New Zealand. N.Z. JI mar. freshw. Res. 5: 86-119.

JAKOBI, H. 1954. Especies novas de Harpacticoide (Copepoda-Crustacea) encontradas em

algas marinhas de litoral Parana, Santa Catarina. Bolm Inst. Oceanogr. S. Paulo 5:

189-211.

LANG, K. 1948. Monographie der Harpacticiden. 1-2. Lund: Ohlsson.

LANG, K. 1965. Copepoda Harpacticoidea from the California Pacific Coast. K. svenska

Vetensk Akad. Handl. 10 (2): 1-560.

Monk, C. R. 1941. Marine harpacticoid copepods from California. Trans. Am. microsc.

Soc. 60: 75-99.

NICHOLLS, A. G. 1941. Littoral Copepoda from South Australia. (1) Harpacticoida. Rec.

S. Aust. Mus. 6: 381-427.

PALLARES, R. E. 1969. El género Scutellidium en la ria Deseado (Crustacea; Copepoda).

Physis. B. Aires 29: 51-72.

VERVOORT, W. 1964. Free living Copepoda from Ifaluk Atoll in the Caroline Islands. Bull.

U.S. natn. Mus. 236: 1-431.

WipBorG, K. F. 1964. Marine copepods of Tristan da Cunha. Results Norw. scient. Exped.

Tristan da Cunha 5 (51): 1-44.

NOTE ADDED IN PRESS

Mielke (1974) has described a $ Scutellidium hippolytes with 5 setae on PS exp. (in

contradiction to Lang’s (1948) description). However, this species is easily distinguish-

able from S. macrosetum by having only 1 seta on P3-4 end. 2.

Mieke, W. 1974. Eulitorale Harpacticoidea (Copepoda) von Spitsbergen. Microfauna

Meeresboden 37: 161-210

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Style manual for biological journals. Washington: American Institute of Biological Sciences.

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REFERENCES

Harvard system (name and year) to be used: author’s name and year of publication given

in text; full references at the end of the article, arranged alphabetically by names, chronologi-

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Examples (note capitalization and punctuation)

BULLOuGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris

88: 100-140.

FISCHER, P.-H., DUvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des

littorines. Archs Zool. exp. gén. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee

region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320.

Koun, A. J. 19605. Spawning behaviour, egg masses and larval development in Conus from the

Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51. :

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L.

Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und

zentralen Siid-Afrika. 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16:

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G. M. Branch

A NEW SPECIES AND RECORDS OF SCUTELLIDIUM

(COPEPODA HARPACTICOIDEA) FROM SOUTH

AFRICA, AND A WORLD KEY TO THE GENUS

VOLUME 66 PART 11 MARCH 1975 ISSN 0303-2515

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A REVISION OF MASSY’S CHECKLISTS OF ‘SOUTH AFRICAN’

CEPHALOPODA

MARTINA A. ROELEVELD

South African Museum, Cape Town

[MS accepted 11 July 1974]

CONTENTS

PAGE

Introduction .. delle tela? 2)

Southern African species listed by Massy (4925). ete ~ 236

Gonatus fabricii . . . Saget 2810)

() Teleoteuthis caribea’. ~:— . =o eee = 236

ycoteuthis-diadema’ <<) 2.2.. 8 We es 236

Asthenoteuthion planctonicum . . . . . . 237

IPICFY SIOLCULNIS C1QV OL = ch n'a es ah ee

Octopodoteuthis larvae 5 ae ek ae

Benthoteuthisimegalops: oe ee | 2ST

IBLGCRIOLCUTNISITUSEH (2016) Mee Shh Pee i ee et ed

Srenoteuthis Dartrani..\., os ne ee eT

Symplectoteuthis oualaniensis . . . . . . ~~ 237

iocranchia:reinhardt) = en ee 238

EVIL ODSISIZV ONG, i ube SOM Ne) Stn ee ee (LOS

MECOCHIGNCYCIUT GT fhe nc sees ee eee ao

iMevalocranchia maxima... 5 oe en es 289

Eoligoireyniaudts Wa 8 ia ee ee ees Ae 239

Loligo indica . . . 3 Sie er Ne Doo

Sepioteuthis laienionaas Se ee AN Cia eT Oo

Sepioteuthis mauritiana: \. :-".. <~ . | 239

Sepioteuthis major. Role eae RDO

Sepia vermiculata = Maracbie Sek ag ene” Bho

SCDIAZONZIVGNICA’, 2) | uk A Oe ee AO

Sepia acuminata? oF. | a ee ee DAD

Sepiaipapiliala CS So NI ees 240

Sepia natalensis. 20). i. eee ee 240

Sepia insignis. . ere i. Deed

Sepia (Doratosepion) pura eee ya eA

Sepia (Doratosepion) burnupi . . . . . . ~~ 244i

Sepia (Doratosepion) incerta. . . . . . . 24i1

Sepia (Doratosepion) confusa . . . . . . 241

ICTHISCDINSHVDICUS) 0 in re tee et 2A

Velodonaiiovata v2. Se 6) a a2

iPolypus eranulatus «4°30. ise oe. 242

(POL PUSIROLRIGUSe a iy eRe ie 2aS

ROW DASEVAIGIVIE (a) 0h Teer ee a” ee

(POLVPUSICADEHSIS| | se wa VEU Se ne eS

Polypusfontanianas yay Ne Nic 2 OS A 248

GISIOBUSINGICHS | 2 rR oe ot aes

VAT COnQUIDATEO! Ue Nad Gs hs Ne ee Got

Argonauta tuberculata’’. 9. . . . . + ,. + 244

233

Ann. S. Afr. Mus. 66 (11), 1975: 233-255

234 ANNALS OF THE SOUTH AFRICAN MUSEUM

PAGE

Amphitretus pelagicus . 244

Non-southern African species listed by Massy 4925). 244

Southeast Atlantic .. 244

Hensenioteuthis niciaehonens 4° in Oe ee Beas

Polypus verrucosus . de Shed Te ACL rae re meee

Southern Ocean (south of 45°S) gh ae ab gt Oe areas

Poly pusdleyisi, to Ge ois eau es Rie eee

Central Indian‘ @cean> 2) 8 ee ee es

Architeuthis:sancti-paulie 2) 1 | ee es

Enoploteuthisshoylei 2 Vs ues OR AS

Loligo tricarinata. . . ac ee ay bee eas

Sepioteuthis SATS Lat De ON is eee

Sepiola stenodactyla . 5 i. 0s 3 ue So ee AS

IPOlyPUS‘GFGNEG” 2 ee se ee ge

Northwestdndian Ocean - =| «+. = «= «ye 245

Abraliopsisimorist 5 vo ot. ee Os ee AS

Calliteuthisihovlei., -2 0 os) 4 8 te, 2

Mastigoteuthis glaucopis. . . . . . . . 246

Liocranchiawaldivie. 35 <6) se. ce, 28 a) ee

Corynomma speculator . . . . . . . . 246

Rossiaimastigophora spin i eh yaee eee ea

SCPIGiSINGAIENSIS o5) sa, oe, ek hey see ee

Sepia. venusia’ 3) es es ee ee

Polypushorsti 5. ws sy a ele ee ee

Polypus:herdmant.- 3... 3) So) saan ae

Polypus arboreseens . « . 2 s.e 8 = 4 2A

Opisthoteuthis MeCAUSOIAES & on. se Mam ee

Species listed in error . iy Ls euih- aaa > Bae eae

Grimalditeuthis bonplandi : i DAT

Southern African species omitted by Massy (1925) ee?” 1)

Moroteuthis sp. . . . eee,

Abraliopsis. gilehristi ..°.. 3. . ea eee

Octopodoteuthopsis sp. «3: |) sa sae eee

Histioteuthis bonnellit.. <=, 24) et ee ee

Todaropsiseblanae sk 4 se ae Ce ee ee

Chiroteuthis sp. ac. 5.) 9 2s Ae ee

Mastigoteuthisssp. ~ . +). cas) al ee eee

Taoniusjpellucida . .. «2-2. Soe pee eae

Liocranchia intermedia . : .°. «. «» « » 248

Cranchiaescabra.” << Re 8 ee eee

Galiteuthisanrmaia. 7. s (sa =») see ee

Anomalocranchia impennis . . . . . . «~~ + 249

Euprymna sp. , Ly aegltcte) Meena eee

Heteroteuthis hawatiensts Cb es ss dT yeni ee

IROSSIG"ENIOIRANICG.. ko el TA Nas Cogs aban een

Sepiatuberculata...%:" «. Nat wees ee eee

Sepia HCGOnIS. “3. 3 oS a So ieee Soe ee

SepICH@ieyGnea > se is el 1 en

Spinulawspinula. «| s.. a. << dete tee eo

OGLODUSRUIBQEIS bw ee) ee

Octopus: Schulizels:0 «5 AE) Se ee ee

Eledone nigra: . os 8 Ae

areonaia higns: <.. os 0 Ws} ee ee

Eledonella.pyemaea... 2 0 CY nd a ee

Cirroteuthis gilchristt. <a) ane ee a

REVISION OF MASSY’S CHECKLISTS OF ‘SOUTH AFRICAN’ CEPHALOPODA 235

PAGE

Grimpoteuthis spp. . i 4, Bo ne ie xe Bae Re EON

Loligo burnupi [species fein nets. ewes (ee ee

Sepiaayinis (species dubia 5°. . «4s. «. 254

Sequel tothe 925ichecklist; 3%... : 2% BAY cote) Zot

Summary . : Pee ty I LS eM ese SOS

Acknowledgements ND ish cee. Sebi 3 1s ee OR MERRY ct RCS

IRCLENENICES ttae cb Alpha ccna Whee aT Raoele eae peo

INTRODUCTION

The works of A. L. Massy and G. C. Robson are important milestones in the

knowledge of southern African cephalopods. Robson published four papers in

1924 on the Cephalopoda collected by Dr J. D. F. Gilchrist when conducting

the South African Marine Survey from the S.S. Pickle, and a short additional

paper in April 1925. In January 1925, apparently without seeing Robson’s

papers of 1924 (possibly due to a delay in publication), Massy published her

first paper on cephalopods of this region and included a checklist of South

African Cephalopoda (pages 204-206), but unfortunately followed Hoyle’s

(1886) definition of this area. Hoyle’s ‘South African Region’ (1886: 217 and

map) extends from about Hollam’s Bird Island on the west coast of southern

Africa to the Red Sea, and includes Madagascar, Mauritius, the Tristan and

Prince Edward Island groups, and the Kerguelen and Heard Islands. Massy’s

checklist covers this same region, but the more modern conception of the

southern African region includes the area south of 20°S (Day 1967: vii). For

pelagic and oceanic species the area has arbitrarily been restricted here to 20° to

45°S and 10° to 40°E. This excludes all the island groups and much of the east

African coast.

It was considered that a separation into regions of the cephalopods listed by

Massy (1925) will be of some use in compiling future checklists and distribution

maps. Since future authors will wish to check on the localities of the species

listed, and on the validity of the separation, before quoting their distributions,

all relevant published records are given, as far as they could be traced. These are

presumably the records upon which Massy based her checklist, although this

cannot be verified, as she gave no details. Where possible, original references

were checked for exact localities, but in a few cases the original references were -

not available; for references quoted at second hand, the source is given in

brackets. Within the zoogeographic regions, the species are presented in the

order and with the names and comments given by Massy (in boldface type),

including her sometimes inconsistent abbreviations of authors’ names and

linking of dual authors by ‘&’, ‘e.’ and ‘et’. The year of publication of the species

and, where necessary, synonyms, have been added (in lightface type).

The species listed by Massy have been divided into two major groups:

species that occur in the southern African region (modern concept —20° to 45°S

and 10° to 40°E) and those that occur outside this region. The non-southern

African species have been allocated to four zoogeographical regions: southeast

236 ANNALS OF THE SOUTH AFRICAN MUSEUM

Atlantic, Southern Ocean (south of 45°S), central Indian Ocean and northwest

Indian Ocean.

A third group of species, those recorded from the southern African region up

to 1925 but not listed by Massy, is also included, so that a complete list of

cephalopod species known from the southern African region up to 1925 is now

available.

Finally, the taxonomic papers published by Robson (1926) and Massy (1927,

1928), and ending with Massy’s list of additions to her checklist (1928: 89-90),

are discussed. No attempt has been made to bring the checklist up to date

beyond 1928, as new records still come to hand with sufficient frequency to show

that the cephalopod fauna of this region is as yet incompletely known. Recent

checklists have been published by Voss (1962, 1967) and Barnard (1974).

SOUTHERN AFRICAN SPECIES LISTED BY MASSY (1925)

Gonatus fabricii (Lichtenstein, 1818)

Gonatus sp. Steenstrup, 1882: 149 (translation) — one specimen ex

Diomedea exulans, a different form to the arctic species of Gonatus, and

fragments, probably of the same species, from 40°S, 15°E.

According to Clarke (1966: 152), G. fabricii is ‘a widely distributed species

from the colder regions of the north Atlantic and north Pacific...’ and “is

quite distinct from all other species except G. antarcticus which is often

considered a southern variety of G. fabricii.’ According to Voss (personal

communication) G. fabricii is most likely a complex of species, and there

appears to be more than one species in the Atlantic; until distinctions have

been made, G. fabricii must serve as a catch-all category.

(?) Teleoteuthis caribea (Lesueur, 1821) [= Onykia carriboea Lesueur, 1821]

Teleoteuthis caribaea: Pfeffer, 1912: 49 — one specimen from 28°40’S,

10°45’E, one from 35°45’S, 22°06’E and one from the Cape of Good Hope.

Thiele, 1920: 444 — one juvenile from 28°46’S, 10°17’E.

According to Voss (personal communication), the genus Onyk-a is in a

somewhat similar situation to that of Gonatus, i.e. that O. carriboea is a

complex of species.

Onychoteuthis banksi (Leach, 1817)

Onychoteuthis banksii: Pfeffer, 1912: 70 — three specimens from the

Cape of Good Hope. Odhner, 1923: 7 — _ one specimen washed on board

off Table Bay.

Lycoteuthis diadema (Chun, 1900)

Thaumatolampas diadema: Chun, 1910: 59 — 19 in vertical net to 3 000 m

from Valdivia Sta. 89, southern part of Benguela Current, 31°21’S, 15°01’E;

12 in vertical net to 1500 m from Valdivia Sta. 118, west wind drift,

40°31’S, 15°06’E.

REVISION OF MASSY’S CHECKLISTS OF ‘SOUTH AFRICAN’ CEPHALOPODA 237

? Lycoteuthis sp. A Robson, 19246: 2 — 19 in 324 fm (595 m) from

Pickle Sta. 156, 30°00’S, 31°22’E.

Asthenoteuthion planctonicum Pfeffer, 1912 [= Lycoteuthis diadema (Chun,

1900) ]

‘Enoploteuthid larvae’ Chun, 1910: 106 — two larvae from Valdivia

Sta. 91, southern part of Benguela Current (no latitude or longitude could

be traced for this station); four larvae from Valdivia Sta. 102, Agulhas

Bank, 34°31’S, 26°00’E. Pfeffer (1912: 770) thought that these larvae

pertained to A. planctonicum.

Pterygioteuthis giardi Fischer, 1896

Pterygioteuthis Giardi: Chun, 1910: 131 — one larva from Valdivia

Sta. 117, southern region of Agulhas Current (the position of this station

could not be traced).

Octopodoteuthis larvae (Chun, 1910) [= Octopoteuthis sp. |

Octopodoteuthis larvae Chun, 1910: 144 — two larvae in vertical net to

1 800 m from Valdivia Sta. 102, Agulhas Current, 34°31’S, 26°00’E.

Benthoteuthis megalops Verrill, 1885 [= Bathyteuthis abyssicola Hoyle, 1885]

Benthoteuthis megalops: Chun, 1910: 185 — _ one specimen in vertical net

to 2500 m from Valdivia Sta. 115, origin of Benguela Current, 36°23’S,

17°38’E. Thiele, 1920: 453 — one specimen in vertical net to 2500 m

from north of Prince Edward Island, 43°04’S, 36°22’E.

Brachioteuthis riisei (Steenstrup, 1882)

Tracheloteuthis Riisei: Hoyle, 1905: 93 — one specimen from 34°40’S,

27° E:

Stenoteuthis bartrami (Lesueur, 1821) [= Ommastrephes bartrami Lesueur, 1821]

Ommastrephes Bartramii: Gray, 1849: 62 — four specimens from Cape

of Good Hope.

? Stenoteuthis Bartrami: Pfeffer, 1912: 465 — one specimen from ‘Cap’.

Stenoteuthis bartrami: Thiele, 1920: 455 — one juvenile from 28°30'S, -

38°13’E. Massy, 1925: 206 — one specimen, washed up on shore after a

storm, Durban.

According to Tomlin (1923: 40) Orbigny recorded this species as taken by

Dussumier at the Cape. |

Symplectoteuthis oualaniensis (Lesson, 1830)

Ommastrephes oualaniensis: Gray, 1849: 63 — one specimen from ? Cape

of Good Hope.

Loligo oualaniensis: Tryon, 1879: p.?2. — specimens from Cape of Good

Hope (according to Gibbons 1888: 202; Smith 1903: 355; Voss 1962: 246)

238 ANNALS OF THE SOUTH AFRICAN MUSEUM

Liocranchia reinhardti (Steenstrup, 1856)

Liocranchia reinhardti a Robson, 19246: 6 — 29, 1? in 220 fm (404 m)

from Pickle Sta. 100, 30°00’S, 31°19’E.

The only previous record that could be found, possibly for this region, is given

by Pfeffer (1912: 673) as ‘Stidatl. Ozean (Putze vend., Mus. GGttingen)’.

Pyrgopsis zygena (Vér.) [Vérany, 1851]

? Pyrgopsis rhynchophorus Rochebrune, 1884: 23 — _ specimen(s) from

Agulhas Bank (according to Clarke 1966: 223).

? Pyrgopsis pacifica: Robson, 1924b: 5 — 129 in 180 fm (330 m) (not

100 fm as stated by Robson) from Pickle Sta. 258, 29°52’S, 31°15’E.

The systematics of the genus Pyrgopsis are confused; six species have

been named (P. zygaena (Vérany, 1851); P. schneehageni (Pfeffer, 1884);

P. rhynchophorus Rochebrune, 1884; P. pacifica (Issel, 1908); P. lemur

Berry, 1920 and P. atlantica Degner, 1925), which have been synonymized

in various combinations by different authors. Pfeffer (1900: 193) synony-

mized P. rhynchophorus Rochebrune, 1884, Loligopsis schnehageni [sic]

Pfeffer, 1884 (= Pyrgopsis schneehageni (Pfeffer, 1884)) and Taonius

richardi Joubin, 1895 (= Galiteuthis armata Joubin, 1898, according to

Clarke (1966: 238)) with Zygaenopsis zygaena (Vérany, 1851), and lists one

of the localities as ‘Siidspitze Afrikas’ (? Rochebrune’s record).

Chun (1910: 354) synonymized all the species referable to Pyrgopsis

with Euzygaena (= Pyrgopsis) pacifica (Issel, 1908). Pfeffer (1912: 656-664)

kept all the species separate, but thought that P. zygaena (Vérany, 1851) is

probably synonymous with P. rhynchophorus Rochebrune, 1884. Clarke

(1966: 221-223) synonymized P. zygaena (Vérany, 1851) with P. pacifica

(Issel, 1908), but retained the others as separate species.

Recent authors (Clarke 1966: 221; Young 1972: 82; Voss, personal

communication) believe that at least some of the Pyrgopsis species are

probably larval forms of Leachia and other closely related cranchiid genera.

Young (1972: 80) has shown that this is true for at least one species, Leachia

dislocata Young, 1972.

Hoyle (1886: 217) listed P. rhynchophorus Rochebrune, 1884 under the

‘South African’ region, but not P. zygaena (Vérany, 1851). Massy (1925:

204) listed only the latter, presumably because she considered these two

species to be synonymous. In fact, prior to Robson’s (19245) records of

P. pacifica (which Massy did not apparently see before compiling her list),

Rochebrune’s record from the Agulhas Bank is the only southern African

record of Pyrgopsis.

Leachia cyclura Lesueur, 1821

Leachia cyclura Lesueur, 1821: 90 — one specimen from ? ‘Pacific

Ocean, 37°S, 33°E’.

Loligopsis cyclura Orbigny, 1848: 322 — specimen(s) from Cape Agulhas.

REVISION OF MASSY’S CHECKLISTS OF ‘SOUTH AFRICAN’ CEPHALOPODA 239

Megalocranchia maxima Pfeffer, 1884

Megalocranchia maxima Pfeffer, 1884: 24 — one specimen from Cape of

Good Hope. This same specimen was described as Desmoteuthis maxima

(Pfeffer 1900: 192) and again as Megalocranchia maxima (Pfeffer 1912: 712).

Hoyle (1886: 217) listed this species as Taonius (?) maximus.

Loligo reynaudi d’Orb. [Orbigny, 1845] [? = Loligo vulgaris Lamarck, 1798]

Loligo reynaudii: Orbigny, 1848: 315 — specimen(s) from Cape of Good

Hope. Gray, 1849: 73 — one specimen from Cape of Good Hope. Hoyle,

1910: 263 — 36 from Angra Pequena (= Liideritzbucht). Hoyle, 1912:

280 — 139, 129, 13¢ and 1 damaged in 35 fm (64 m) from Scotia Sta.

480, 8 miles (13 km) N of Dassen Island. Thiele, 1920: 440 — _ one speci-

men from Valdivia Sta. 101, Algoa Bay, 33°51’S, 25°29’E. Massy, 1925:

207 — 16, 19 in 35-40 fm (64-73 m) from Cape Henderson and 192 from

Durban.

Loligo indica Pfeffer, 1884 [= Loligo duvaucelii Orbigny, 1835]

Loligo indica: Robson, 1924d: 654 — 19 in 36 fm (66 m) from Pickle Sta.

167, Natal coast, 29°31’S, 31°24’E. Massy, 1925: 208 — 2d, 32 from

Durban; 1d from Natal coast.

Sepioteuthis loliginiformis (Ruppell & Leuckart, 1828)

Sepioteuthis loliginiformis: Martens, 1879: 727 — specimen(s) from

Inhambane.

Sepioteuthis mauritiana Quoy e. Gaimard, 1832 [= Sepioteuthis Jessoniana

Lesson, 1830]

Sepioteuthis lessoniana Férussac et d’Orbigny (= S. mauritiana Quoy et

Gaimard): Joubin, 1898: 26 — specimen(s) from the Cape.

Massy presumably had not seen Joubin’s paper, since she listed S. mauri-

tiana (? on the basis of Quoy & Gaimard’s record from Mauritius) but not

S. lessoniana, whilst Joubin had already synonymized the two species, of

which S. Jessoniana has priority.

Sepioteuthis major Gray, 1828 (Insufficiently characterized) [? = Thysanoteuthis —

rhombus (Troschel, 1857)]

Sepioteuthis major Gray, 1828: 3 — specimen(s) from Cape of Good

Hope (according to Gray 1849: 83).

Sepia vermiculata Quoy e. Gaimard, 1832 = hierredda Rang, 1837 [= Sepia

officinalis vermiculata Quoy & Gaimard, 1832, non = Sepia officinalis hierredda

Rang, 1837]

Sepia vermiculata Quoy & Gaimard, 1832: 64 — specimen(s) from Cape

of Good Hope (according to Gibbons 1888: 202; Bartsch 1915: 250;

Smith 1903: 356, 1916: 20). Robson, 1924b: 12 — 1¢ in 38 fm (70 m)

240 ANNALS OF THE SOUTH AFRICAN MUSEUM

from Pickle Sta. 386, 29°25’S, 31°37’E; and 1¢ from Natal coast. These

same specimens were later (Robson 1924d: 639) described as Acanthosepion

vermiculata. Massy, 1925: 209 — 19 in 40 fm (73 m) from Cape Hender-

son; 1¢ from Natal coast; 12 from Congella, Durban; 1 ¢ from Durban;

192 in 135 fm (248 m) from Natal coast.

Sepia hierredda (non Rang): Orbigny, 1841: 268 — specimen(s) from

Cape of Good Hope. Smith, 1916: 20 — mentions specimens in the

British Museum from Port Elizabeth, Cape Colony, and Tongaat Beach,

Natal. In the previous paragraph, however, he states his belief that S. hier-

redda is synonymous with S. vermiculata, as suggested by Orbigny.

Sepia jousseaumi Rochebrune, 1884: 117 — specimen(s) from Cape of

Good Hope (according to Smith 1903: 356, 1916: 22; Bartsch 1915: 250;

Adam 1941: 108).

Sepia Filliouxi: Joubin, 1898: 24 — 1,4 from the Cape.

Sepia zanzibarica Pfeffer, 1884

Sepia zanzibarica: Smith, 1916: 21 — one shell from Tongaat, Natal.

Tomlin, 1923: 40 — specimen(s) from Isipingo.

Sepia acuminata E. A. Smith, 1916 (Shell only known)

Sepia acuminata Smith, 1916: 21 — shells from Port Elizabeth and

Tongaat Beach, Natal. One shell from Tongaat, mentioned by Smith as

being different from the others, is S. hieronis (Robson, 1924). Robson,

1924b: 12 — 29 in 192 fm (352 m) (not 95 fm as stated by Robson) from

Pickle Sta. 95; 29° 52:8, 3-17 E:.

Rhombosepion acuminata Robson, 1924d: 643 — 29 from Pickle Sta. 95

(as above) and 2 in 160 fm (294 m) from Pickle Sta. 103, 29°54’S, 31°15’E.

Sepia papillata Quoy e. Gaimard, 1832

Sepia papillata Quoy & Gaimard, 1832: 61 — specimen(s) from Cape of

Good Hope (according to Krauss 1848: 133; Smith 1916: 22). Massy, 1925:

211 — 13 from 16 miles (26 km) NE of Bird Island.

Sepia tuberculata (non Lamarck): Orbigny, 1848: pl. 17, figs 13-15 — ?

specimen(s) from Cape of Good Hope. Steenstrup, 1875: 479 — 14, 19,

from the Cape. Hoyle, 1910: 265 — 39 from Angra Pequena (Liideritz-

bucht).

Spathidosepion tuberculatum Rochebrune, 1884: 94 — 1¢ without shell

from Cape of Good Hope (according to Adam 1944: 226).

Sepia natalensis sp. noy. [= Sepia simoniana Thiele, 1920]

Sepia tuberculata (non Lamarck): Gray, 1849: 101 — one shell from

Cape of Good Hope.

Sepia papillata (non Quoy & Gaimard): Smith, 1916: 22 — shells from

Port Elizabeth and Tongaat Beach, Natal.

Sepia simoniana Thiele, 1920: 436 — 3, 22 from Simons Bay; one shell

REVISION OF MASSY’S CHECKLISTS OF ‘SOUTH AFRICAN’ CEPHALOPODA 24]

from Olifant River mouth. Odhner, 1923: 7 — one specimen in 40 fm

(73 m) from 10 miles (16 km) from Cape Barracouta.

Sepia natalensis Massy, 1925: 212 — 12in 35-40 fm (64-73 m) from Cape

Henderson; 29 from 16 miles (26 km) NE of Bird Island; 12 from Durban.

Sepia insignis E. A. Smith, 1916 (shell only known)

Sepia insignis Smith, 1916: 25 — _ two shells from Tongaat Beach, Natal.

Sepia (Doratosepion) australis Quoy e Gaimard, 1832

Sepia australis Quoy & Gaimard, 1832: 70 — _ specimen(s) from Agulhas

Bank to 90 miles (145 km) from Cape Agulhas (according to Krauss 1848:

133; Smith 1916: 24). Hoyle, 1912: 281 — 19in35 fm (64 m) from 8 miles

(13 km) N of Dassen Island. Smith, 1916: 24 — shell(s) from Port

Elizabeth. Robson, 19246: 11 — 19, 1?¢ in 117 fm (215 m) from Pickle

Sta. 2, 33°03’S, 17°42’E; 63, 62 in 112 fm (206 m) from Pickle Sta. 7,

32°32'S, 17°42’E; 23, 89 in 100 fm (184 m) from Pickle Sta. 9, 32°37’S,

17°41’E; 33 in 68 fm (125 m) (not 67 fm as stated by Robson) from Pickle

Sta. 20, 32°20’S, 17°55’E; 13, 22 in 85 fm (156 m) from Pickle Sta. 44,

33°20’S, 17°49’E; 29 in 121 fm (222 m) from Pickle Sta. 54, 33°09’S, 17°36’E.

These specimens were later described as Rhombosepion capense (Robson

1924d: 641).

Sepia capensis: Orbigny, 1848: 278 — _ specimen(s) from Agulhas Bank.

Krauss, 1848: 133 — one specimen from Kalk Bay. Thiele, 1920: 438

— one specimen from Valdivia Sta. 114, Simons Bay, 34°20’S, 18°36’E;

one specimen from Valdivia Sta. 101, Algoa Bay, 33°51’S, 25°29’E; five

specimens from Valdivia Sta. 105, Agulhas Bank, 35°29’S, 21°03’E; one

specimen from Valdivia Sta. 106, Agulhas Bank, 35°27’S, 20°56’E.

Sepia (Doratosepion) australis Massy, 1925: 214 — 13 from 16 miles

(26 km) NE of Bird Island.

Hoyle (1886: 217) listed this species for South Africa as Sepia capensis.

Sepia (Doratosepion) burnupi Hoyle, 1904

Sepia burnupi Hoyle, 1904: 27 (partim) — two shells from Umkomaas,

Natal. Smith, 1916: 23 — shells from Tongaat Beach, Natal.

Sepia (Doratosepion) incerta E. A. Smith, 1916 :

Sepia burnupi Hoyle, 1904: 27 (partim) — _ two shells from Port Elizabeth.

Sepia incerta Smith, 1916: 23 — shells from Tongaat Beach, Natal.

Sepia (Doratosepion) burnupi (non Hoyle) Massy, 1925: 215 — 1¢ in

38 fm (70 m) from Cape Henderson; | ¢ from Natal coast.

Sepia (Doratosepion) incerta Massy, 1925: 219 — 29 from Cape Hender-

son.

Sepia (Doratosepion) confusa E. A. Smith, 1916

Sepia burnupi Hoyle, 1904: 27 (partim) — one shell from Port Elizabeth.

Sepia confusa Smith, 1916: 24 — shell(s) from Tongaat Beach, Natal.

242

ANNALS OF THE SOUTH AFRICAN MUSEUM

Robson, 19246: 12 — 29 in 158 fm (290 m) from Pickle Sta. 102, 29°51’S,

31°16’E; 19 in 36 fm (66 m) from Pickle Sta. 167, 29°31’S, 31°24’E; 23 in

192 fm (352 m) from Pickle Sta. 95, 29°52’S, 31°17’E; 1g in 180 fm (330 m)

from Pickle Sta. 107, 29°49’S, 31°18’E. These specimens were later

described as Doratosepion confusum (Robson 1924d: 647).

Sepia (Doratosepion) confusa Massy, 1925: 221 — 14 from Natal coast.

Anatomy of the ‘tail’ of the male specimens described by Robson (1924) and

Massy (1925) was described (as Doratosepion confusa) by Massy & Robson

(1923: 435) and Carleton & Robson (1924: 259).

Hemisepius typicus Steenstrup, 1875 [= Sepia (Hemisepius) typica (Steenstrup,

1875) ]

Hemisepius typicus Steenstrup, 1875: 468 — 19 from Table Bay. Hoyle,

1912: 281 — 29 in 8-10 fm (15-18 m) from Saldanha Bay. Chun, 1915:

411 — 14, 19 from Valdivia Sta. 100, St Francis Bay, 34°09’S, 24°59’E.

Velodona togata Chun, 1915

Velodona togata var. a Robson, 1924a: 206 — 2 in 220 fm (404 m) from

Pickle Sta. 99, 29°55’S, 31°21’E; 12 in 250 fm (459 m) from Pickle Sta. 396,

29°55’S, 31°22’E; 192 in 240 fm (440 m) from Pickle Sta. 162, 30°03’S,

31°12’E. These specimens were later described as Velodona togata var.

capensis (Robson 1924d: 644).

Strictly speaking this species should be listed with the non-southern

African species, as Massy had not seen Robson’s papers when she compiled

the checklist, and the only previous record of V. togata was from the East

African coast (Chun 1915: 479); but it is felt that this would only add to the

confusion already caused by the checklist itself.

Polypus granulatus (Lam.) [Lamarck, 1798] [= Octopus vulgaris Cuvier, 1797]

Octopus granulatus: Hoyle, 1886: 80 — one young specimen in 10-20 fm

(18-37 m) from Simons Bay, Cape of Good Hope.

Polypus granulatus: Thiele, 1915: 487 — 19 from Cape Town. Thiele,

1920: 436 — 1 juvenile from Simonstown. Massy, 1925: 222 —— 1° from

Park Rynie; 12 from Indian Ocean; 19 from Tongaat Beach, Natal; one

mutilated specimen from H. Reich’s coll.; 1¢ from 16 miles (26 km) NE of

Bird Island; 19, 1¢ from Scottburgh, Natal; |g from Durban.

Polypus rugosus: Robson, 1924d: 669 — 29 in 38 fm (70 m) from Pickle

Sta. 388, Natal coast, 29°27’S, 31°37’E; 13 in 40 fm (73 m) from Pickle

Sta. 389, Natal coast, 29°27’S, 31°39’E.

Robson (1929: 63) and Pickford (1955: 152) have placed Octopus granulatus

Lamarck, 1798 in synonymy with Octopus rugosus (Bosc, 1792). Massy

(1925: 222) was of the same opinion, as she listed Polypus rugosus (Orbigny,

1835) as a synonym of Polypus granulatus (Lamarck, 1798). She did not,

however, recognize Bosc’s inadequate description of Sepia rugosa (1792) as

valid, and so upheld Octopus granulatus Lamarck as having priority, but at

REVISION OF MASSY’S CHECKLISTS OF ‘SOUTH AFRICAN’ CEPHALOPODA 243

the same time giving priority to Polypus over Octopus. The priorities of these

generic names are discussed by Robson (1929: 56), and Pickford (1955: 151)

has discussed the complexities of the synonymies of Octopus vulgaris,

O. rugosus and O. granulatus. On re-examining Robson’s specimens

Pickford (1955: 158) found that “The majority of specimens from the eastern,

western, central and south Atlantic, and from West and South Africa which

Robson referred to this species (i.e. O. rugosus) are probably correctly

assigned to O. vulgaris. It cannot be established that O. rugosus is a valid

species.’

Polypus horridus (d’Orb.) [Orbigny, 1826] [= Octopus horridus Orbigny, 1826]

Octopus argus Krauss, 1848: 132 — one specimen from Natal.

Octopus horridus: Gibbons, 1888: 201 — specimen(s) from Algoa Bay.

Polypus valdivie Chun, 1915 [= Bathypolypus valdiviae (Thiele, 1915)]

? Octopus capensis Eydoux & Souleyet, 1852: 11 — one juvenile from

Cape of Good Hope, insufficiently described. Thiele (1915: 485) and Robson

(1932: 303) believed that this was probably identical with Bathypolypus

valdiviae.

Polypus valdiviae Thiele, 1915: 485 — 1, 32in 500 m from Valdivia Sta.

103, Agulhas Bank, 35°11’S, 23°02’E.

Bathypolypus grimpei Robson, 1924a: 208 — 14, in 470 fm (862 m) from

Pickle Sta. 405, Natal coast, 30°00’S, 31°35’E; 3, 42 in 380 fm (697 m)

from Pickle Sta. 164, Natal coast, 30°06’S, 31°31’E. Robson later decided

(1932: 303) that the differences between B. valdiviae and B. grimpei were

insufficient to warrant the erection of a new species for the latter.

Bathypolypus ? valdivie: Robson, 1924d: 669 — 14 in 220 fm (404 m)

from Pickle Sta. 63, Cape Town, 33°42’S, 17°36’E.

‘Undetermined polypods’ Robson, 1924d: 670 — _ one specimen in 260 fm

(477 m) from Pickle Sta. 46, west of Cape Town, 33°25’S, 17°24’E; one

specimen in 260 fm (477 m) from Pickle Sta. 294 (?), 25°48’S, 33°38’E.

Robson (1932: 303) placed these under B. valdiviae.

Polypus capensis (Eydoux et Souleyet, 1852) [? = Bathypolypus valdiviae

(Thiele, 1915)]

Octopus capensis Eydoux & Souleyet, 1852: 11 — one juvenile trom

Cape of Good Hope, insufficiently described.

Polypus fontanianus (d’Orb.) [Orbigny, 1835] [= Octopus fontanianus Orbigny,

1835]

Polypus fontanianus: Massy, 1925: 224 — 19 from Natal coast. Robson

(1929: 187) erected the genus Joubinia for this species and Polypus campbelli.

He thought (1929: 189) that Massy’s specimens represented a well-marked

local variety, and named them Joubinia fontaniana var. africana. Adam

(1938: 223) substituted Robsonella for Joubinia, as the latter was shown ts

244 ANNALS OF THE SOUTH AFRICAN MUSEUM

be preoccupied. Pickford (1955: 163) re-examined Robson’s specimens of

Robsonella fontaniana and concluded that the genus Robsonella is probably

not valid, and that the species assigned to it should be returned to the genus

Octopus.

Cistopus indicus Rapp, MS [= Cistopus indicus (Orbigny, 1840) ]

Octopus Indicus: Martens, 1879: 727 — _ specimen(s) from Inhambane,

Mocambique.

Argonauta argo Linn. [Linnaeus, 1758]

Argonauta argo: Orbigny, 1840: 158 — Rang (1829) collected this species

on the Agulhas Bank, and Reynaud in False Bay (no reference to Reynaud

given). Krauss, 1848: 133 — _ shells in fairly large numbers on south coast

of Cape Colony, from False Bay to Algoa Bay. Hoyle, 1886: 69 — one

shell from Cape of Good Hope. Gibbons, 1888: 201 — specimen(s) from

East London. Sowerby, 1892: 1 — _ this species is frequently found on the

shores of South Africa. Bartsch, 1915: 2 — one specimen from Port

Alfred.

Argonauta tuberculata Shaw, 1791 [= Argonauta nodosa Solander, 1786]

Argonauta nodosa: Humphrey, 1797: 6 — specimen(s) from Cape of

Good Hope (according to Smith 1903: 355).

Argonauta tuberculata: Orbigny, 1826: 138 — _ specimen(s) from Cape of

Good Hope (according to Robson 1932: 200). Krauss, 1848: 133 — Rang

found this species at Cape Agulhas (no reference to Rang given).

[?] Argonauta bottgeri Maltzan, 1881

Sowerby (1892: 1) lists this species for South Africa and refers to his earlier

paper (1889) where, however, A. boettgeri is not mentioned! Bartsch (1915:

230) also lists this species for South Africa, but gives no locality.

Amphitretus pelagicus Hoyle, 1885

Amphitretus pelagicus: Thiele, 1915: 532 — _ one specimen in vertical net

to 1 800 m from Valdivia Sta. 102, Agu!has Current, 34°31’S, 26°00’E.

NON-SOUTHERN AFRICAN SPECIES LISTED BY MASSY (1925)

SOUTHEAST ATLANTIC

Hensenioteuthis melancholicus (Chun, 1906) [= Sandalops melancholicus Chun,

1906]

Sandalops melancholicus: Chun, 1910: 379 — _ two specimens in vertical

net to 1 000 m from NE of Tristan da Cunha. The position was 32°08'S,

8°28’W (Thiele 1920: 458).

REVISION OF MASSY’S CHECKLISTS OF ‘SOUTH AFRICAN’ CEPHALOPODA 245

Polypus verrucosus (Hoyle, 1885) [= Octopus vulgaris Cuvier, 1797]

Octopus verrucosus Hoyle, 1885: 222 — 23 from Inaccessible Island,

Tristan da Cunha (according to Hoyle 1886: 79).

SOUTHERN OCEAN (SOUTH OF 45°s)

Polypus levis (Hoyle, 1885) [= Benthoctopus levis (Hoyle, 1885)]

Octopus levis Hoyle, 1885: 229 — 23, 22in75 fm (137 m) from Challenger

Sta. 151, off Heard Island, 52°59'30’S, 73°33’30"W (this should be

73°33'30’E) (according to Hoyle 1886: 98).

Thiele’s (1915: 486; 1920: 435) specimen from Kerguelen was considered by

Robson (1932: 227, 233) to be a distinct species, Benthoctopus thielei.

CENTRAL INDIAN OCEAN

Architeuthis sancti-pauli (Vélain, 1877)

Mouchezia sancti-pauli Vélain, 1877: 83 — one specimen from St Paul

Island.

Clarke (1966: 98) has since recorded specimens of Architeuthis spp. from

the stomachs of sperm whales caught near Durban and Saldanha Bay.

Enoploteuthis hoylei Pfeffer, 1884 [= Abraliopsis hoylei (Pfeffer, 1884) ]

Enoploteuthis hoylei Pfeffer, 1884: 17 — 1 from the Mascarenes.

Loligo tricarinata Gray, 1849 (Insufficiently characterized)

Loligo tricarinata Gray, 1849: 73 — two shells from Isle of France

(Mauritius).

Sepioteuthis madagascarensis Gray, 1849 (Insufficiently characterized)

Sepioteuthis madagascariensis Gray, 1849: 80 — one specimen from

Madagascar.

Sepiola stenodactyla Grant, 1833 [= Euprymna stenodactyla (Grant, 1833)]

Sepiola stenodactyla Grant, 1833: 84 — one specimen from Mauritius.

Polypus aranea (d’Orb.) [Orbigny, 1826] [= Octopus filamentosus Blainville,

1826]

Octopus filamentosus Blainville, 1826: 188 — specimen(s) from Mauritius

(according to Robson 1929: 143).

Octopus aranea: Orbigny, 1840: 57 — specimen(s) from Mauritius.

Martens, 1879: 727 — _ specimen(s) from Mocgambique.

NORTHWEST INDIAN OCEAN

Abraliopsis morisii (Vér.) [Vérany, 1837]

Abraliopsis Morisii: Chun, 1910: 78 — 13 in 977 m from Valdivia Sta.

254, Indian North Equatorial Current, 0°29’S, 42°47’E; 12 in 1 134 m from

246 ANNALS OF THE SOUTH AFRICAN MUSEUM

Valdivia Sta. 256, Indian North Equatorial Current, 1°49’N, 45°29’E;

specimen(s) from stomach of Coloconger raniceps caught in 628 m from

Valdivia Sta. 265, Indian North Equatorial Current, 6°24’N, 49°31’E.

According to Voss (personal communication) Chun’s detailed illustrations

involve probably three nominal species, A. hoylei, A. lineata and a new

species (Voss, in manuscript).

Calliteuthis hoylei (Goodrich, 1896)

Calliteuthis Hoylei: Chun, 1910: 170 — 19 in vertical net to 2 000 m from

Valdivia Sta. 235, Indian South Equatorial Current near Amirante Islands,

4°34’S, 53°42’E.

N. Voss (1969: 784) considered Chun’s specimen to pertain to Histioteuthis

dofleini (Pfeffer, 1912) and not to Histiopsis Hoylei Goodrich, 1896, a

species dubia.

Mastigoteuthis glaucopis Chun, 1908

Mastigoteuthis glaukopis Chun, 1908: 88 — _ one specimen from proximity

of East African coast. Chun, 1910: 233 — one specimen in | 213 m from

Valdivia Sta. 261, Indian North Equatorial Current, 4°63’S, 48°37’E

(probably the same specimen).

Liocranchia valdiviz Chun, 1906

Liocranchia Valdiviae: Chun, 1910: 337 — 19 in vertical net to 2 000 m

from Valdivia Sta. 237, tongue of Indian South Equatorial Current,

4°45’S, 48°58’E; 13 in vertical net to 2500 m from Valdivia Sta. 239,

tongue of Indian South Equatorial Current, 5°42’S, 43°36’E; 1g in 1 362m

from Valdivia Sta. 258, Indian North Equatorial Current, 2°58’N, 46°50’E.

Corynomma speculator Chun, 1906

Corynomma speculator: Chun, 1910: 367 — _ one specimen in vertical net

to 2000 m from Valdivia Sta. 237, tongue of Indian South Equatorial

Current, 4°45’S, 48°58’E.

Rossia mastigophora Chun, 1915

Rossia mastigophora Chun, 1915: 405 — 14, 22 in 638 m from Valdivia

Sta. 253, Indian North Equatorial Current, 0°27’S, 42°47’E.

Sepia singalensis Goodrich, 1896 [= Sepia pharaonis Ehrenberg, 1831]

Sepia singalensis: Hoyle, 1907: 459 — 29 from Zanzibar.

Sepia venusta Pfeffer, 1884 [2 = Sepia pharaonis Ehrenberg, 1831)

Sepia venusta Pfeffer, 1884: 12 — one specimen from Zanzibar.

Polypus horsti (Joubin, 1898) [= Octopus cyanea Gray, 1849]

Polypus horsti: Hoyle, 1907: 451 — four specimens from Zanzibar.

REVISION OF MASSY’S CHECKLISTS OF ‘SOUTH AFRICAN’ CEPHALOPODA 247

Polypus herdmani Hoyle, 1904 [= Octopus herdmani (Hoyle, 1904)]

Polypus herdmani: Hoyle, 1907: 454 — 19, 1? in 10 fm (18 m), 19 in

unknown depth, from Zanzibar.

Polypus arborescens Hoyle, 1904 [= Octopus arborescens (Hoyle, 1904) ]

Polypus arborescens: Hoyle, 1907: 454 — 19 in 5 fm (9 m), 492, 13 in

10 fm (18 m) from Zanzibar.

Opisthoteuthis medusoides Chun, 1915 [= Opisthoteuthis medusoides Thiele,

1915]

Opisthoteuthis medusoides Thiele, 1915: 538 — two specimens in 400 m

from Valdivia Sta. 243, Dar-es-Salaam, 6°39’S, 39°31’E.

O. medusoides is the only species of Opisthoteuthis that is bell-shaped, the

others being more or less flat. In this connection Bidder (1968: 1038) has

made an interesting comment: ‘the “‘species”’ figured by Chun (1914) under

the name “‘medusoides”’ should probably be regarded not as a true species,

but as evidence of the medusa-like swimming... by some chance of fixa-

tion, a group of specimens were fixed in the “‘systole’’ of the swimming

phase.’ The swimming of cirrate octopods is also discussed by Roper &

Brundage (1972). The author of O. medusoides was actually Thiele, who

wrote part of Chun’s Valdivia report, as stated on page 536 of that publica-

tion, and the year of publication was 1915, not 1914.

SPECIES LISTED IN ERROR

Grimalditeuthis bonplandi (Vér.) [Vérany, 1837] [= Grimalditeuthis bomplandi

(Vérany, 1837)]

Grimalditeuthis bonplandi: Pfeffer, 1900: 188 — lists this species as

recorded from the southern tip of Africa, but according to Pfeffer (1912:

635) this locality is incorrect. According to Clarke (1966: 214) this species

has been recorded from the North and Southwest Atlantic.

SOUTHERN AFRICAN SPECIES OMITTED BY MASSY (1925)

Although Massy apparently did not see Robson’s papers (1924a-d), and

certainly not that of 1925, which was published three months after her checklist,

Robson’s records are included here for completion of the checklist to 1925.

Moroteuthis sp. Robson, 1924

Moroteuthis sp.A Robson, 19246: 2; 1924d: 595.

12 in 135 fm (247 m) from Pickle Sta. 530, 33°34’S, 17°23’E.

Abraliopsis gilchristi (Robson, 1924)

Abralia gilchristi Robson, 19246: 3; 1924d: 601.

13 in 240 fm (439 m) from Pickle Sta. 48, 33°36’S, 19°30’E;

12 in 280 fm (512 m) from Pickle Sta. 81, 33°28’S, 17°19’E.

248 ANNALS OF THE SOUTH AFRICAN MUSEUM

Octopodoteuthopsis sp. Robson, 1924

Octopodoteuthopsis sp. A Robson, 19245: 4.

Octopodoteuthopsis sp. Robson, 1924d: 606.

One specimen in 900 fm (1 646 m) from Pickle Sta. 357 (not Sta. 347, as

stated by Robson 1924d), 32°36’S, 16°00’E.

Histioteuthis bonnellii (Férussac, 1835)

Histioteuthis bonelliana: Robson, 1924b: 4; 1924d: 608.

12 in 470 fm (860 m) from Pickle Sta. 343, 30°10’S, 14°33’E;

1g in 958 fm* (1 752 m) from Pickle Sta. 542, 31°41’S, 17°09’E.

According to N. Voss (1969: 845), Robson’s specimens pertain to H. macro-

hista N. Voss, 1969.

Todaropsis eblanae (Ball, 1841)

Todaropsis eblanae: Robson, 1924b: 5; 1924d: 614.

23, 32 in 112 fm (205 m) and 2¢ in 126 fm (230 m) from Pickle Sta. 7,

32732, 9517-428

23, 72 in 100 fm (183 m) from Pickle Sta. 9, 32°37'S, 17°41'E;

13, 12 in 67 fm (123 m) from Pickle Sta. 20, 32°20’S, 17°55’E;

13 in 85 fm (155 m) from Pickle Sta. 44, 33°20’S, 17°49’E;

23, 12 in 67 fm (123 m) from Pickle Sta. 54, 33°09’S, 17°36’E;

1g, 22 in 104 fm (190 m) (not 108 fm as stated by Robson 19245) from

Pickle Sta. 58, 32°26’S, 17°32’E.

Chiroteuthis sp. Robson, 1924

Cheiroteuthis (Doratopsis-stage) A Robson, i924c: 591.

One larva in plankton net at 100 fm (183 m) off Durban, 29°55’S, 31°22’E.

Mastigoteuthis sp. Robson, 1924

Mastigoteuthis sp. A Robson, 19246: 5; 1924d: 617.

One specimen in 820 fm (1 500 m) from Pickle Sta. 277, 29°42’S, 31°35’E.

Taonius pellucida (Chun, 1910)

Desmoteuthis pellucida Chun, 1910: 357 — 19 in vertical net to 1 000 m

from Valdivia Sta. 90, Benguela Current, 33°20’S, 15°58’E.

Pfeffer (1912: 716) repeated the description of Chun’s specimen under the

name Megalocranchia pellucida.

Liocranchia intermedia Robson, 1924

Liocranchia intermedia Robson, 1924b: 6; 1924d: 632.

13 in 260 fm (475 m) from Pickle Sta. 174, Natal coast, 29°40’S, 31°41’E.

Cranchia scabra Leach, 1817

Cranchia scabra: Robson, 1924b: 10; 1924d: 624.

1 22 in 820 fm (1 500 m) from Pickle Sta. 277, Natal coast, 29°42’S, 31°35 EB.

* According to the station list (Gilchrist 1922: 47) the depth at this station was 580 fm

(1061 m).

REVISION OF MASSY’S CHECKLISTS OF ‘SOUTH AFRICAN’ CEPHALOPODA 249

Galiteuthis armata Joubin, 1898

Galiteuthis armata: Robson, 19246: 8; 1924d: 627.

1 ?9 in 900 fm (1 646 m) from Pickle Sta. 357, Cape Town, 32°36’S, 16°00’E.

Anomalocranchia impennis Robson, 1924

Anomalocranchia impennis Robson, 1924b: 9; 1924d: 628.

One specimen in 418 fm (764 m) from Pickle Sta. 540, Cape Town, 33°30’S,

byt 2E.

Euprymna sp. Hoyle, 1912

Euprymna sp. Hoyle, 1912: 280 — one young specimen in 8-10 fm

(15-18 m) from Scotia Sta. 482, Saldanha Bay.

Heteroteuthis hawaiiensis dagamensis Robson, 1924

Heteroteuthis hawaiiensis Berry, var. dagamensis Robson, 19246: 11;

1924d: 632.

19 in 350 fm (640 m) from Pickle Sta. 176, Natal coast, 29°43’S, 31°49’E;

19 in 610 fm (1 116 m) from Pickle Sta. 251, Cape Town, 33°43’S, 17°24’E.

Rossia enigmatica Robson, 1924

Semirossia sp. A Robson, 19246: 10.

Rossia enigmatica Robson, 1924d: 635.

19 in 220 fm (402 m) from Pickle Sta. 63, Cape Town, 33°42’S, 17°36’E;

1g im 151 fm (276 m) from Pickle Sta. 6, Cape, 32°32’S, 17°18’E;

1 22 in 260 fm (475 m) from Pickle Sta. 46, 33°25’S, 17°24’E.

In the second paper (Robson 1924d) only the first two specimens above were

listed for R. enigmatica; the third specimen is mentioned at the end of the

description as being in poor condition, but probably also pertaining to this

species.

Rossia sp. Robson, 1925: 450.

22 from ‘Cape area’. Voss (1962: 253) re-examined these specimens and

considered them to be conspecific with R. enigmatica.

Sepia tuberculata Lamarck, 1798

Sepia tuberculata: Orbigny, 1848: 277 — specimen(s) from Cape of

Good Hope.

Spathidosepion tuberculatum Rochebrune, 1884:93 — 19and | shell from

Cape of Good Hope (according to Adam 1944: 226).

Spathidosepion papillatum (non Quoy & Gaimard) Rochebrune, 1884:

p? — 19 from Port Dorey (according to Adam 1944: 226). This locality

is probably incorrect (Adam, personal communication); S. tuberculata is

endemic to South Africa.

Gray (1849: 101) lists S. tuberculata from the Cape of Good Hope, but

apparently synonymizes S. papillata with it; the only specimen Gray had,

one shell, pertains to S. simoniana, according to Adam & Rees (1966: 109).

250 ANNALS OF THE SOUTH AFRICAN MUSEUM

According to Monttort (see Smith, 1916: 25) Lamarck’s types were from

the Cape of Good Hope.

Hoyle (1886: 217) listed this species from ‘South Africa’, but followed

Orbigny in considering S. papillata to be a synonym. Massy recognized that

the two species are distinct, but listed only S. papillata, although S. tuber-

culata had been recorded from South Africa.

Sepia hieronis (Robson, 1924)

Sepia acuminata Smith, 1916: 21 (partim) — one shell from Tongaat

Beach, Natal.

Sepia sp. A Robson, 19245: 13.

Rhombosepion hieronis Robson, 1924d: 645.

Two specimens in 117 fm (214 m) from Pickle Sta. 2, 33°03’S, 17°42’E;

227i 112 tm (205 m) from Pickle Sta. 7, 32-32 8, 17 42 E:

1g in 126 fm (230 m) from Pickle Sta. 8, 32°33’S, 17°29’E;

one specimen in 150 fm (274 m) from Pickle Sta. 33, 32°54’S, 17°25’E.

Sepiella cyanea Robson, 1924

Sepiella cyanea Robson, 1924b: 13; 1924d: 648.

12 in 40 fm (73 m) (not 60 fm as stated by Robson 1924d) from Pickle Sta.

389, 29°27'S,, 31 -39'E;

13 in 28 fm (51 m) from Pickle Sta. 476, 29°17’S, 31°33’E.

Spirula spirula (Linnaeus, 1758)

Spirula peronii: Krauss, 1848: 134 — many shells found after storm on

beach at Plettenberg Bay. Gibbons, 1888: 202 — specimens from Table

Bay, Algoa Bay and Natal. Sowerby, 1892: 1 — occasionally found on

beach, Port Elizabeth. Bartsch, 1915:3 — onespecimen from Port Alfred.

Spirula solandri: Bartsch, 1915: 251 — listed for South Africa, but no

locality or reference given.

Octopus vulgaris Cuvier, 1797

Octopus vulgaris: Krauss, 1848: 132 — _ three specimens from Natal coast-

Polypus vulgaris: Thiele, 1920: 436 — one specimen from Simons Bay.

Octopus schultzei (Hoyle, 1910)

Polypus schultzei Hoyle, 1910: 261 — 14 from Angra Pequena (Liideritz-

bucht).

Eledone nigra (Hoyle, 1910)

Moschites nigra Hoyle, 1910: 262 — 14, 62 from Angra Pequena

(Liideritzbucht).

Robson (1932: 279) placed this species in the genus Pareledone, but Voss

(personal communication) has re-examined the type and other specimens

in the Berlin Museum and found that the species is a typical eledonid

belonging to the genus Eledone (of which Moschites is a synonym).

REVISION OF MASSY’S CHECKLISTS OF ‘SOUTH AFRICAN’ CEPHALOPODA 251

Benthoctopus berryi Robson, 1924

Benthoctopus berryi Robson 1924d: 658.

12 in 1 200 fm (2 195 m) from Pickle Sta. 524, SW Cape Town, 33°31’S,

16°39’E (according to the station list (Gilchrist 1922: 44) the depth at this

station was | 220 fm (2 232 m)).

Argonauta hians Solander, 1786

Argonauta kochiana: Bartsch, 1915: 230 — listed for South Africa, but

no locality or reference given.

Eledonella pygmaea Verrill, 1884

Eledonella massye Robson, 1924a: 202; 1924d: 672.

12 in 1 014 fm (1 854 m) from Pickle Sta. 87, SW Cape Town, 33°55’S,

706 E.

Cirroteuthis gilchristi Robson, 1924

Cirroteuthis gilchristi Robson, 1924a: 204; 1924d: 677.

13 in 1 400 fm (2 560 m) from Pickle Sta. 526, SW Cape Town, 33°17’S,

16°24’E.

Grimpoteuthis spp. (Robson, 1924)

Cirroteuthis A Robson, 1924d: 680. -

Three specimens in 1 000 fm (1 828 m) from Pickle Sta. 525, Cape Town,

33°28’S, 16°40’E.

Eight specimens in 1 400 fm (2 560 m) from Pickle Sta. 526, Cape Town,

33°17'S, 16°24’E.

Cirroteuthis B Robson, 1924d: 682.

One specimen in 790 fm (1 443 m) from Pickle Sta. 85, 33°46’S, 17°12’E.

Robson later (1932: 155, 156) placed these specimens in the genus Grimpo-

teuthis.

Loligo burnupi Smith [species dubia]

Bartsch (1915: 241) lists this species from Port Shepstone; it is not known

to what species or reference this refers.

Sepia affinis Orbigny, 1826 [species dubia]

According to Smith (1916: 26), Montfort (1805) recorded this species from

the Cape of Good Hope, but Smith comments that ‘Sepia affinis, d’Orbigny,

is not a Sepia, but evidently belongs to the genus Sepioteuthis, and although

said by d’Orbigny and others to be the same as Sepioteuthis sepoidea [sic]

(Blainville, 1823), it is not likely to be identical considering the remoteness

of the localities—West Indies and South Africa’.

SEQUEL TO THE 1925 CHECKLIST

In 1926 Robson published a short additional paper on decapod cephalopods

collected by the S.S. Pickle, including two species not covered in his previous

papers on the collection, and constituting new records for southern Africa:

Tetronychoteuthis sp. and Histioteuthis bonnellii.

252 ANNALS OF THE SOUTH AFRICAN MUSEUM

The following year Massy (1927) described a collection of South African

Cephalopoda made by the S.S. Pieter Faure. This collection included two new

species, Rhombosepion robsoni (= Sepia robsoni) and Sepia (Doratosepion)

joubini, and one new record, Inioteuthis japonica (? = Inioteuthis capensis Voss,

1962); the remaining species had already been recorded from southern Africa

(1925 checklist and/or Robson 1924a,b,d).

In 1928 Massy described another collection of South African cephalopods,

all but one of which had already been recorded from the region (checklist 1925).

The remaining species was Sepiella obtusata (= S. cyanea, already recorded

from South Africa by Robson 19246,d). She also gave a list of additions to the

checklist of 1925, consisting of the species described by Robson (1924a,b,d), but

omitting Sepiella cyanea. She did not, however, include in this list of additions

her own species and record described in 1927, nor did she mention the species

described by Robson (1924c, 1926).

The additions to the checklist (for which the necessary synonyms have

already been given) listed by Massy (1928) are as follows:

Moroteuthis sp. Robson, 1924

Lycoteuthis sp. Robson, 1924

Abralia gilchristi Robson, 1924

Octopodoteuthopsis sp. Robson, 1924

Histioteuthis bonelliana (Férussac, 1835)

Todaropsis eblanae Ball, 1841

Mastigoteuthis sp. Robson, 1924

Liocranchia reinhardti Steenstrup, 1856, var. Robson, 1924

Liocranchia intermedia Robson, 1924

Cranchia scabra Leach, 1817

Pyrgopsis pacifica (Issel, 1908)

Galiteuthis armata Joubin, 1898

Anomalocranchia impennis Robson, 1924

Loligo spp. Robson, 1924

Heteroteuthis hawaiiensis (Berry, 1909), var. dagamensis Robson, 1924

Rossia enigmatica Robson, 1924

Rhombosepion hieronis Robson, 1924

Velodona togata Chun, 1915, var. capensis Robson, 1924

Benthoctopus berryi Robson, 1924

Bathypolypus grimpei Robson, 1924

Cirroteuthis gilchristi Robson, 1924

Cirroteuthis sp. Robson, 1924

Eledonella massyae Robson, 1924

The species omitted by Massy (1928) but recorded from southern Africa up

to 1928 are:

Cheiroteuthis sp. Robson, 1926 (= Chiroteuthis sp.)

Histioteuthis bonelliana (Férussac, 1835) (ex Robson 1926) (= H. bonnellii)

REVISION OF MASSY’S CHECKLISTS OF ‘SOUTH AFRICAN’ CEPHALOPODA 253

Tetronychoteuthis sp. Robson, 1926

Inioteuthis japonica Verrill, 1881 (ex Massy 1927) (? = J. capensis Voss, 1962)

Sepia (Doratosepion) joubini Massy, 1927

Rhombosepion robsoni Massy, 1927 (= Sepia robsoni (Massy, 1927))

Sepiella cyanea Robson, 1924

SUMMARY

Since Massy’s checklist of South African Cephalopoda covers a very much

larger area than that now considered to be the southern African zoogeographical

region, the species listed by Massy have been separated into South African and

non-South African. The locality records upon which Massy’s checklist was

probably based are given. For completion the species recorded from the southern

African region up to 1925 but not listed by Massy (1925) are also given. Finally,

the works of Massy and Robson on South African cephalopods, published up

to 1928 and culminating in Massy’s list of additions to the 1925 checklist, are

briefly discussed.

ACKNOWLEDGEMENTS

I should like to thank Drs N. A. H. Millard and M. A. Cluver, both of the

South African Museum, for advice on the manuscript.

REFERENCES

Asterisked references (*) were not seen in the original.

ApaM, W. 1938. Robsonella nom. nov. fiir Joubinia Robson, 1929 (Cephalopoda, Octopoda).

Zool. Anz. 121: 223-224.

ApaM, W. 1939. Cephalopoda. I. Le genre Sepioteuthis Blainville, 1824. Siboga Exped. Monogr.

55a: 1-34.

ApaM, W. 1941. Résultats scientifiques des croisiéres du navire-école belge ‘Mercator’. III (4).

Cephalopoda. Mém. Mus. r. Hist. nat. Belg. (2) 21: 83-161.

ApaM, W. 1944. Révision de Il’ ‘Etude monographique de la famille des Sepiadae’ d’A. T. de

Rochebrune (1884). Mém. Mus. natn. Hist. nat., Paris (n.s.) 18: 219-242.

ApaM, W. & Rees, W. J. 1966. A review of the cephalopod family Sepiidae. Scient. Rep. John

Murray Exped. 11: 1-165.

BARNARD, K. H. 1974. Contributions to the knowledge of South African marine Mollusca.

Part VII. Revised fauna list. Ann. S. Afr. Mus. 47: 663-781.

BarTscu, P. 1915. Report on the Turton collection of South African marine mollusks, with

additional notes on other South African shells contained in the United States National

Museum. Bull. U.S. natn. Mus. 91: 1-305.

Bipper, A. M. 1968. Some problems of cephalopod locomotion. Symp. Ser. mar. biol. Ass.

India 3: 1029-1052. ee

*BLAINVILLE, H. M. D. pe. 1826. In: Cuvier, F., ed. Dictionnaire des sciences naturelles. Paris:

Levrault.

CARLETON, H. M. & Rosson, G. C. 1924. On the histology and function of certain secondary

sexual organs in the cuttlefish Doratosepion confusa. Proc. R. Soc. (B) 96: 259-271.

CHUN, C. 1908. Uber Cephalopoden der Deutschen Tiefsee-Expedition. Zool. Anz. 33: 86-89.

Cuun, C. 1910. Die Cephalopoden. I. Teil: Oegopsida. Wiss. Ergebn. dt. Tiefsee-Exped.

‘Valdivia’ 18: 1-401.

CHUN, C. 1915. Die Cephalopoden. II. Teil. Myopsida, Octopoda. Wiss. Ergebn. dt. Tiefsee-

Exped. ‘Valdivia’ 18: 405-552.

254 ANNALS OF THE SOUTH AFRICAN MUSEUM

CLARKE, M. R. 1966. A review of the systematics and ecology of oceanic squids. Adv. mar. Biol.

4: 91-300.

Day, J. H. 1967. A monograph on the Polychaeta of southern Africa. Part 1. Errantia. London:

British Museum (Natural History).

Eypoux, J. F. & SOuLEYET, F. L. 1852. Voyage autour du monde exécuté pendant les années 1836

et 1837, sur la corvette la Bonite, commandée par M. Vaillant. Zoologie. 2: 7-36. Paris.

GrBBons, J. S. 1888. Partial list of the South African Mollusca. Trans. S. Afr. phil. Soc. 4:

201-219.

GILcuRIST, J. D. F. 1922. Fisheries and marine biological survey. Report No. 2. For the year

1921. Rep. Fish. Mar. Biol. Surv. Un. S. Afr. 2: 1-84.

GRANT, R. E. 1833. On the anatomy of the Sepiola vulgaris, Leach, and account of a new

species (Sep. stenodactyla, Grant,) from the coast of Mauritius. Trans. zool. Soc. Lond.

1: 77-86.

*GRAY, J. E. 1828. Spicilegia zoologica. 1. London: Treuttet & Wiirtz.

Gray, J. E. 1849. Catalogue of the Mollusca in the collection of the British Museum. 1. Cephalo-

poda Antepedia. London: British Museum (Natural History).

*Hoy_e, W. E. 1885. Diagnoses of new species of Cephalopoda collected during the cruise of

H.M.S. ‘Challenger’. Part I. The Octopoda. Ann. Mag. nat. Hist. (5) 15: 222-236.

Hoy Le, W. E. 1886. Report on the Cephalopoda collected by H.M.S. Challenger during the

years 1873-1876. Rep. Voy. Challenger 1873-76 16: 1-245.

Hoy Le, W. E. 1904. Sepia burnupi n. sp. from Natal. J. Conch., Lond. 11: 27-28.

Hoye, W. E. 1905. The marine fauna of the west coast of Ireland. IJ. On specimens of

Tracheloteuthis and Cirroteuthis from deep water off the west coast of Ireland. Rep. Sea

Inland Fish. Ireland 1902/03 (2): 93-98.

Hoy .e, W. E. 1907. The marine fauna of Zanzibar and East Africa, from collections made by

Cyril Crossland in 1901-1902. —The Cephalopoda. Proc. zool. Soc. Lond. 1907 (2): 450-461.

Hoy _e, W. E. 1910. Mollusca: Cephalopoda. In: SCHULTZE, L. Zoologische und anthropologische

Ergebnisse einer Forschungsreise im westlichen und zentralen Siidafrika 4: 261-268. Jena:

Fischer. Denkschr. med-naturw. Ges. Jena 16: 261-268.

Hoy _e, W. E. 1912. The Cephalopoda of the Scottish National Antarctic Expedition. Trans.

R. Soc. Edinb. 48: 273-283.

*HuMPHREY, G. 1797. Museum Calonnianum. Part 1. London.

JouBIN, L. 1898. Sur quelques céphalopodes du Musée Royal de Leyde et description de trois

espéces nouvelles. Notes Leyden Mus. 20: 21-28.

Krauss, F. 1848. Die siidafrikanischen Mollusken. Stuttgart: Ebner & Seubert.

LesuEur, C. A. 1821. Descriptions of several new species of cuttle-fish. J. Acad. nat. sci. Philad.

2: 86-101.

MARTENS, E. v. 1879. Ubersicht der von ihm [Hr. W. Peters] von 1843 bis 1847 in Mossambique

gesammelten Mollusca. Mber. K. preuss. Akad. Wiss. Berlin 1879: 727-749.

Massy, A. L. 1925. On the Cephalopoda of the Natal Museum. Amn. Natal Mus. 5: 201-229.

Massy, A. L. 1927. The Cephalopoda of the South African Museum. Ann. S. Afr. Mus. 25:

151-167.

Massy, A. L. 1928. On the Cephalopoda of the Natal Museum. Part II. Ann. Natal Mus. 6:

89-96.

Massy, A. L. & Rosson, G. C. 1923. On a remarkable case of sex-dimorphism in the genus

Sepia. Ann. Mag. nat. Hist. (9) 12: 435-442.

*MONTFORT, P. D. pe. 1805. Histoire naturelle des mollusques. Jn: BUFFON, G. L. L. DE.

Histoire naturelle. Nouv. éd. (suite). Paris.

OpunerR, N. Hj. 1923. Contributions to the marine molluscan faunas of South and West

Africa. Géteborgs K. Vetensk.-o. VitterhSamh. Handl. (4) 26 (7): 1-40.

*ORBIGNY, A. D. D’. 1826. Tableau méthodique de la classe des céphalopodes. Annis Sci. nat.

7: 96-169.

OrsIGny, A. D. p’. 1840, 1841, 1848. In: Férussac, A. DE. & ORBIGNY, A. D’. 1835-48. Histoire

naturelle générale et particuliére des céphalopodes acétabuliféres vivants et fossiles. Paris:

Bertrand.

PFEFFER, G. 1884. Die Cephalopoden des Hamburger Naturhistorischen Museums. Abh. Geb.

Naturw. Hamburg 8: 63-90.

PFEFFER, G. 1900. Synopsis der oegopsiden Cephalopoden. Mitt. naturh. Mus. Hamb. 17:

145-198.

REVISION OF MASSY’S CHECKLISTS OF ‘SOUTH AFRICAN’ CEPHALOPODA 255

PFEFFER, G. 1912. Die Cephalopoden der Plankton-Expedition. Ergebn. Atlant. Ozean Plankton-

exped. Humboldt-Stift. 2: 1-815.

PICKFORD, G. E. 1955. A revision of the Octopodinae in the collections of the British Museum.

Bull. Br. Mus. nat. Hist. (Zool.) 3: 151-167.

Quoy, J. R. & GAIMARD, J. P. 1832. Mollusques. In: Zoologie du voyage de I’ Astrolabe, pendant

les années 1826-1829. 2. Paris: Tastu.

*RANG, S. 1829. Sur l’animal de l’Argonaute. Férussac, Bull. Sci. Nat. 17: 132-134.

*Rapp. [Unpublished manuscript, referred to in Orbigny 1840.]

Rosson, G. C. 1924a. On new species &c. of Octopoda from South Africa. Ann. Mag. nat.

Hist. (9) 13: 202-210.

Rosson, G. C. 19245. Preliminary report on the Cephalopoda (Decapoda) procured by the

S.S. ‘Pickle’. Rep. Fish. mar. biol. Surv. Un. S. Afr. 3 (Spec.,Rep. 9): 1-14.

Rosson, G. C. 1924c. On a new Doratopsis-stage of Cheiroteuthis from S. E. Africa. Ann. Mag.

nat. Hist. (9) 13: 591-594.

Rosson, G. C. 1924d. On the Cephalopoda obtained in South African waters by Dr. J. D. F.

Gilchrist in 1920-21. Proc. zool. Soc. Lond. 1924: 589-686.

Rosson, G. C. 1925. On a new species of Rossia from South Africa. Ann. Mag. nat. Hist. (9)

15: 450-454.

Rosson, G. C. 1926. The Cephalopoda obtained by the S.S. ‘Pickle’. Supplementary report.

Rep. Fish. mar. biol. Surv. Un. S. Afr. 4 (Spec. Rep. 8): 1-6.

Rosson, G. C. 1929. A monograph of the recent Cephalopoda. Part 1. Octopodinae. London:

British Museum (Natural History).

Rosson, G. C. 1932. A monograph of the recent Cephalopoda. Part II. The Octopoda (excluding

the Octopodinae). London: British Museum (Natural History).

*ROCHEBRUNE, A. T. DE. 1884a. Etude monographique de la famille des Loligopsidae. Bull. Soc.

Dhilomath. Paris (7) 8: 7-28.

*ROCHEBRUNE, A. T. DE. 18845. Etude monographique de la famille des Sepiadae. Bull. Soc.

Dhilomath. Paris (7) 8: 74-122.

Roper, C. F. E. & BRUNDAGE, W. L. 1972. Cirrate octopods with associated deep-sea

organisms: new biological data based on deep benthic photographs (Cephalopoda).

Smithson. Contr. Zool. 121: 1-46.

SmiTH, E. A. 1903. A list of species of Mollusca from South Africa, forming an appendix to

G. B. Sowerby’s ‘Marine shells of South Africa’. Proc. malac. Soc. Lond. 5: 354-402.

SmiTH, E. A. 1916. On the shells of the South African species of Sepiidae. Proc. malac. Soc.

Lond. 12: 20-26.

Sowersy, G. B. 1889. Further notes on marine shells of South Africa, with descriptions of new

species. J. Conch., Lond. 6: 147-159.

SOWERBY, G. B. 1892. Marine shells of South Africa. London: The Author.

STEENSTRUP, J. 1875. Hemisepius, en ny slaegt af Sepia-blaeksprutternes familie med bemaerk-

ninger om Sepia-formerne i almindelighed. K. danske Vidensk. Selsk. Skr. (5) 10: 465-482.

STEENSTRUP, J. 1882. Notae teuthologicae. 1. Overs. danske Vidensk. Selsk. Forh. 1882:

143-168. (English translation: Vots@eE, A. et al. The cephalopod papers of Japetus Steen-

strup. Copenhagen: Danish Science Press, 1962.)

THIELE, J. 1915. In: CHUN, C. Die Cephalopoden. II. Teil: Myopsida, Octopoda. Wiss. Ergebn.

dt. Tiefsee-Exped. ‘Valdivia’ 18: 472-475, 485-487, 490-494, 531-533, 534-538. :

THIELE, J. 1920. Die Cephalopoden der deutschen Siidpolar-Expedition 1901-1905. Dt. Siidpol.-

Exped. 16: 433-465.

TOMLIN, J. R. LE B. 1923. On South African marine Mollusca with descriptions of several new

species. J. Conch., Lond. 17: 40-52.

*TRYON, G. W. 1879. Cephalopoda. Man. Conch. 1: 1-316.

VELAIN, C. 1877. Remarques générales au sujet de la faune des iles St. Paul et Amsterdam, suivies

d’une description de la faune malacologique des deux iles. Archs Zool. exp. gén.6: 1-144.

Voss, G. L. 1962. South African cephalopods. Trans. R. Soc. S. Afr. 36: 245-272.

Voss, G. L. 1967. Some bathypelagic cephalopods from South African waters. Ann. S. Afr.

Mus. 50: 61-88.

Voss, N. A. 1969. A monograph of the Cephalopoda of the North Atlantic. The family Histio-

teuthidae. Bull. mar. Sci. 19: 713-867.

Youn, R. E. 1972. The systernatics and areal distribution of pelagic cephalopods from the

seas off southern California. Smithson. Contr. Zool. 97: 1-159.

INSTRUCTIONS TO AUTHORS

Based on

CONFERENCE OF BIOLOGICAL EDITORS, COMMITTEE ON FORM AND STYLE. 1960.

Style manual for biological journals. Washington: American Institute of Biological Sciences.

MANUSCRIPT

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REFERENCES

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in text; full references at the end of the article, arranged alphabetically by names, chronologi-

cally within each name, with suffixes a, b, etc. to the year for more than one paper by the same

author in that year.

For books give title in italics, edition, volume number, place of publication, publisher.

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volume number, part number (only if independently paged) in parentheses, pagination.

Examples (note capitalization and punctuation)

BuLLouGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris

88: 100-140.

FISCHER, P.-H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des

littorines. Archs Zool. exp. gén. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee

region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 1960b. Spawning behaviour, egg masses and larval development in Conus from the

Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L.

Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und

zentralen Siid-Afrika. 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16:

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—_ =F

ff &

a - eo

| :

‘

ce Lv =

MARTINA A. ROELEVELD

A REVISION OF MASSY’S CHECKLISTS

OF ‘SOUTH AFRICAN’ CEPHALOPODA

VOLUME 66 PART 12 MARCH 1975 —— ISSN 0303-2515

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ADDITIONS TO THE SOUTH AFRICAN

SPECIES OF PHTHIRIINAE AND USIINAE

(DIPTERA: BOMBYLIIDAE)

WITH KEYS TO ALL THE KNOWN SPECIES

A. id. MESSE

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ADDITIONS TO THE SOUTH AFRICAN SPECIES OF PHTHIRIINAE

AND USIINAE (DIPTERA: BOMBYLIIDAE) WITH KEYS TO ALL THE

KNOWN SPECIES

A. J. HESSE

South African Museum, Cape Town

(With 3 figures)

[ MS. accepted 19 July 1974]

CONTENTS

PAGE

Introduction : : : : E : 257

Descriptions and keys. : 3 : 257

Summary . : i 3 : : : 307

Acknowledgements . : : : : 308

References . : : , ; : ; 308

INTRODUCTION

The acquisition of additional material by more intensive field work and

collecting, on the part of the South African Museum since my revision of

the South African Phthiriinae (Hesse 1938: 822-866), necessitates the descrip-

tions of quite a number of new species, the recording of undescribed sexes of

species described from one sex only, and the correction of observations made

on inadequate material.

It is also to be noted that since my revision of this subfamily, in which the

genera Phthiria Meigen, Apolysis Loew and Oligodranes Loew were accommo-

dated, Hull (1973: 194, 212) has transferred the two genera Apolysis and

Oligodranes to the subfamily Usiinae which was formerly primarily based

on the genus Usia from the Mediterranean Region and North Africa. The

former subfamily Phthiriinae of authors thus becomes restricted to the genus

Phthiria.

DESCRIPTIONS

Subfamily Phthiriinae Becker

Genus Phthiria Meigen

Phthiria: Hesse, 1938: 824. Hull, 1973: 195-199.

Phthiria laeta Bezzi

Phthiria laeta Bezzi, 1921: 96. Hesse, 1938: 831, fig. 248a.

The acquisition of additional 29 and the as yet undescribed ¢ of this

species throws an entirely different light on the identity of the species and

necessitates not only additional observations to facilitate its recognition, but

257

Ann. S. Afr. Mus. 66 (12), 1975: 257-308, 3 figs.

258 ANNALS OF THE SOUTH AFRICAN MUSEUM

also the correction of a taxonomic error made in allocating to it a variety

(xerophila) which actually belongs to an entirely different species.

The species, both 3 and 9, may be recognized as follows:

Body on the whole less humped and broader than in other species, mainly

black in g, only narrowish hind margins of sternites pallid; broadish hind

margins of tergites on sides in ¢ appearing greyish white, due to greyish white

tomentum; knees in ¢ yellowish; following parts in 2 very pale yellow or

yellowish white: the broadish occipital margin behind eyes continuous on each

side with a broadish streak on each side of frons, extending down entire genae

(excepting only for a black spot or streak on each side of antennae), the entire

buccal cavity, entire head below (except for a short, central, basal, abbreviated,

dark streak), the shoulders, sides of thorax above, slightly more than basal

half of scutellum, broad hind margins of tergites (broader on sides), entire

pleurae (except for dark lower part of mesopleuron, dark spot on anterior

lower part of pteropleuron, a dark spot on anterior upper part of sterno-

pleuron, the black lower parts or halves of sterno- and hypopleurae, and

some infusions around base of halteres), the very broad, or almost entire,

hind margins of sternites, and the coxae; legs in @ slightly more ochreous

yellow, only hinder parts of trochanters, extreme apices of tibiae, and apices

of tarsal joints, or sometimes apical halves of latter, darkened.

Vestiture with the hairs relatively sparse, a little denser in dg, entirely

white in 3, slightly more sericeous yellowish on head above, occiput above

and thorax above anteriorly in 9; sparse scaling, where present, silvery whitish

in 4g, and yellowish or golden above in 9.

Head with the anterior and genal part not tumidly prominent; antennal

joint 1 very short, subequal to joint 2; joint 3 (cf. Hesse 1938, fig. 248a) spindle-

shaped, in profile slightly broadened at about middle above and again below

nearer apex, but not humped, its lower apical process slightly thicker and pro-

jecting slightly more than upper one, the two together forming an almost

symmetrical bifid process; proboscis about 2-2,8 mm, more slender in 0.

Scutellum markedly broad and transverse, more so than in most other

species, broadly rounded behind, its width being a little more than twice its

length.

Wings greyish hyaline, with yellowish brownish, brownish to reddish-

brownish veins; knobs of halteres entirely pallid in 9, darkened above in ¢.

Legs with the basal joint of hind tarsi only about or a little more than

half length of hind tibiae.

Length of body: about 44,5 mm

Length of wing: about 4,5-5 mm

Distribution

Transvaal (¢ allotype from Johannesburg (Zumpt, 24 December 1950));

Natal.

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE 259

Fig. 1. Left side views of antennae and heads of some new species of Phthiria. (Antennae to

the same scale, the heads to a smaller scale.)

A. 9 hesperia; B. S nitidigena; CC. 2 flaviscuta; D. 92 conocephala; E. ¢ ovalicornis;

F. 3 brunnescens; G. 2 namaquensis.

Phthiria flaviscuta sp. nov.

Fig. 1C

This species may almost be considered as only a distinct western Cape

or southern Karoo variety of /aeta. In the absence of the ¢ these 99 are how-

ever placed as a separate species. Compared with the 2 /aeta they agree and

differ in the following respects:

Body slightly smaller, with the same pattern of yellow, disposed in the

same way, but the entire scutellum discally yellow and the yellow hind margins

of tergites distinctly narrower, occupying less than apical halves of tergites

on sides; legs with more darkened parts, the upper surfaces of front femora

and sometimes a subapical infusion on inner or upper apical parts of the others,

260 ANNALS OF THE SOUTH AFRICAN MUSEUM

outer faces of all the tibiae, and the greater part of tarsi darkened. The left

antenna is shown in Figure 1C.

Vestiture with the hairs and scaling, especially on thorax above and ter-

gites, distinctly longer and denser, and distinctly more yellowish or golden;

fine hairs and scaling on frons denser, more golden.

Wings very similar, but relatively shorter; knobs of halteres with a distinct

dark spot above and tending to be darkened above in outer part.

The species appears to be slightly variable; some specimens have no lateral

extension of the black on frons in front, and in others the legs are much darkened.

From 4 99 in the South African Museum.

Length of body: about 2,5-4 mm

Length of wing: about 3,5-4,5 mm

Distribution

Little Karoo: Oudtshoorn-Zebra (Museum expedition, October 1951)

(holotype); Vanwyksdorp (October 1937); Spitskop near Meiringspoort

(Museum staff, November 1935). South-western Cape: Franschhoek (Barnard,

December 1932).

Phthiria hesperia sp. nov.

Fig. 1A

This species, represented by a 9 specimen only, may eventually prove

to be only a western form of flaviscuta, but in view of the absence of the 3 and

its distinct type of third antennal joint, it is referred to a separate species which

differs from the 99° of Jaeta and especially flaviscuta in the following respects:

Antennal joint 3 (cf. Fig. 1A) ending in an upper process which is much

farther away from the apex, distinctly much more subapical in position and

conspicuously less developed and thinner than the relatively stout lower apical

process, but both these processes however more developed than in /aeta and

flaviscuta.

Central black fascia on frons not extending fascia-like laterally on frons

in front as in /aeta; yellow on sides of thorax anteriorly above as broad as in

flaviscuta and broader than in /aeta; entire scutellum yellowish as in former

species; legs with the front femora and the other femora more extensively

yellowish than in flaviscuta, and tibiae on the whole also more extensively

yellowish, not so darkened on outer surfaces; hairs on legs apparently shorter

and less developed than in flaviscuta; knobs of halteres without a distinct dark

spot or infusion laterally above.

From a @ holotype in the South African Museum.

Length of body: 3,76 mm

Length of wing: about 3,88 mm

Length of proboscis: about 2 mm

Distribution

Western Cape: Leipoldtville (Museum expedition, November 1956).

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE 261

Phthiria pulla Bezzi

Phthiria pulla Bezzi, 1922: 78. Hesse, 1938: 847.

Phthiria fallax Hesse, 1938: 836, fig. 250 (n. syn.).

Phthiria flavigenualis Hesse, 1938: 837, fig. 251 (n. syn.).

Phthiria laeta var. xerophila Hesse, 1938: 831, fig. 248b (n. syn.).

A long series of both 39 and 929 of a species of Phthiria from the Little

Karoo and also the Great Karoo, obtained since the publication of my revision,

has convinced me that the 33 of the species in question are in fact the species

pulla of Bezzi which he described from a 3 specimen from Willowmore.

As I suspected in 1938 (p. 838) there is no doubt that my flavigenualis,

described from 4 34, is identical with pulla. The single 3 specimen, described

as fallax, appears to be merely a colour variety and should also now be con-

sidered as a synonym of pulla.

The long series of 29 in this collection obviously belong to the same species

as the gd and constitute the as yet undescribed 9 sex of pulla. Moreover the

2° of my /aeta var. xerophila also do not differ specifically from the above-

mentioned newly-acquired 99. In this case there is also no doubt that my

laeta var. xerophila (both 3 and 9) represents a slight colour variety of pulla.

A more complete description of this slightly variable species pulla, as based

on both sexes, is as follows:

Body in $ mainly black, only the narrow hind margins of sternites obscurely

whitish or pallid, and last sternite infused with yellowish or reddish-yellow to

a variable extent; body in 2 with the following parts pale yellow to ochreous

yellow: almost entire head (except the dark part on occiput surrounding the

neck, the black ocellar region, an obscure dark central streak on frons of

variable width and intensity to frontal depression, a dark spot on each side

of antennae, the dark antennal joints 2 and 3, and the dark proboscis), broadish

sides of thorax above, continuous anteriorly with a narrowish submedial

dorsal line or streak on each side which often isolates a roundish dark humeral

spot and another more transverse dark spot just above wing-bases, base of

thorax to a variable extent, entire scutellum, greater part of pleurae (except

the dark or black lower halves of sterno- and hypopleurae, the dark parts

surrounding base of halteres, and usually some small obscure dark spots below

wing-bases, and sometimes a darkish streak or infusion across lower part or

half of mesopleuron), almost entire abdomen (except the dark or black bases

of tergites, especially centrally and discally above), and the coxae; narrowish

hind margins of tergites in 2 tending to be more pallid or yellowish whitish,

and the basal parts more orange or yellowish red; legs in 3 black, but with

rather conspicuous yellowish or reddish-yellow knees, with the entire femora,

tibiae and bases of tarsi yellowish in one colour form (described by me as var.

xerophila), and with only apices of femora, entire middle and hind tibiae, upper

surfaces of front tibiae and bases of tarsi yellowish in another colour form

(described as fallax); legs in 2 entirely yellowish, only apices of tibiae and tarsi

(or apical parts of tarsi) darkened.

262 ANNALS OF THE SOUTH AFRICAN MUSEUM

Vestiture with the hairs and scaling relatively short and very sparse in

typical 9, but distinctly longer and denser, especially on abdomen of some 99

(described as var. xerophila), the hairs in 9 distinctly more yellowish, sericeous

yellowish to pale golden on body above; vestiture on body and legs in ¢ much

denser and longer than in 9, entirely whitish or sericeous white, apparently

denser in some 3 with yellowish or partly yellowish legs (those described as

var. xerophila and fallax); the 3 more often with a greyish white tomentum or

pruinescence which appears as broadish greyish white transverse band across

hind margin on sides of tergites.

Head not tumidly prominent anteriorly; antennal joint | short, trans-

verse, not longer than joint 2; joint 3 broadened at or near middle to a variable

extent, subspindle-shaped, without any longish hairs above, its apical processes

about equally long and prominent, the lower one a little broader at base, both

together forming a more or less symmetrical bifid process; proboscis about

1,4-2,6 mm long, stouter in 9.

Wings greyish hyaline, without a milky whitish tint; veins brownish to

dark brown or even blackish-brown; middle cross vein at about, or a little

beyond, middle of discoidal cell; knobs of halteres usually entirely pallid or

pale yellowish above, but in some 3 sometimes with a tendency to be slightly

darkened on outer margin above.

Legs with the basal joint of hind tarsi usually longer or much longer than

half length of hind tibiae.

Length of body: about 3-5 mm

Length of wing: about 3,5-5 mm

Distribution

Koup Karoo, Tankwa Karoo, Little Karoo, Great Karoo, and Namaqua-

land.

The synonymic names may be retained as varietal names for the various

colour forms as indicated in brackets in the description above.

From the material before me it is evident that the typical pulla (syn.

flavigenualis) occurs mostly in the Little Karoo and Great Karoo. The other

colour form, pulla var. fallax, in which the ¢ has yellowish tibiae and the 9

has denser hairs on the abdomen, appears to occur in the Koup Karoo and

adjacent parts of the Great Karoo, extending into Namaqualand in the west

where it is replaced by the variety xerophila, with entirely yellowish legs in

the 3. Still another, more southern, form of pulla, from the region between the

Little Karoo and south Cape, may be added here:

Phthiria pulla var. anomalocera n.

The ¢ of this form differs from the ¢ of pulla Bezzi s.str. in being slightly

smaller; antennal joint 1 distinctly longer, nearly twice length of small trans-

verse joint 2; proboscis relatively shorter, only about 1,5 mm long; abdomen

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE 263

without distinct, or with less conspicuous, bands of greyish tomentum across

hind margins of tergites on sides; last sternite entirely dark or black; and with

the knobs of the halteres entirely very dark or black above.

From a ¢ in the South African Museum.

Distribution

South Cape: Uniondale District (Museum expedition, October 1952).

Phthiria varipes sp. nov.

This species obviously belongs to the South West African crocogramma

and cognata section (Hesse 1938: 833, 835) in which the 99 have very dark or

black legs, less yellow on pleural parts, relatively narrower yellow hind margins

across tergites, often rather broadish wings, and a third antennal joint in which

the upper apical process is shorter, less developed and apparently farther back

than lower one.

It is nearer crocogramma which type of species it represents in South

Africa. Though having the yellow on its body disposed in the same way, it

differs from the latter in the following respects:

Body distinctly smaller; entire occiput behind eyes in at least its upper

half black, not with yellow eye margins; base of thorax entirely black like rest

of thorax above; anterior spiracular part below humeral angle without any,

or scarcely any, yellow; yellow hind margins of tergites distinctly narrower.

Head with the lower apical process of antennal joint 3, though longer, or

projecting more, than upper one, not markedly stouter as in crocogramma;

proboscis distinctly much more slender, much shorter, only about 2 mm long.

Wings more vitreous hyaline, without the faint, but distinct, subopacity

of crocogramma.

Legs with the front coxae more yellowish-brown in front in apical part,

not entirely black; femora not entirely or uniformly dark, but with more pale

yellowish-brown on inner and upper faces in more than their basal halves,

and the knees not so broadly yellowish.

From a 2 holotype in the South African Museum.

Length of body: about 3 mm

Length of wing: about 4 mm

Distribution

Moordenaars Karoo, north-west of Laingsburg (Museum staff, March

1937). .

Phthiria pubescens Bezzi

Phthiria pubescens Bezzi, 1921: 98. Bezzi, 1922: 77-78. Hesse, 1938: 839. (2 only.)

In my revision of the South African species of Phthiria I refered a 3

specimen from ‘Weenen in Natal (Thomasset, March-April 1924)’ to this

species, at the same time ignoring an undescribed ¢ specimen referred to the

264 ANNALS OF THE SOUTH AFRICAN MUSEUM

same species by Bezzi and which was supposed to be in the Hungarian Museum,

but which I could not identify from Bezzi’s references and unsatisfactory key

(Bezzi 1922: 77). This g in any case no longer exists for the dipterous collec-

tions in that museum were destroyed by fire during the revolution of 1957.

Since the publication of my revision another 3 and two 99 from the same

locality and obtained by the same collector as the above-mentioned ¢ have

come to my notice. The differently coloured 99, which obviously belong to

the same species as the 3, prove that my interpretation at the time was wrong

and that the 3 specimen described by me as the J of pubescens is not the 3

of that species. It, together with the other ¢ and the 99, are described below

as a new species.

The ¢ of pubescens Bezzi s.str. is therefore still unknown and this species

is still represented only by the unique 9 specimen, labelled by Bezzi, in the

South African Museum.

Phthiria tinctipennis sp. nov.

Phthiria pubescens (3 only) Hesse (nec Bezzi), 1938: 839, fig. 252 (n. syn.).

As mentioned under the preceding species, a ¢ specimen from Weenen in

Natal was, in the absence of the true 9, wrongly described by me as the g

pubescens Bezzi. It, together with another ¢ and 2 99 from the same locality

and collected at more or less the same time by the same collector, as well as

a slightly aberrant or varietal ¢ from the Transvaal, constitute a distinct and

separate species which is characterized and differs from other known South

African species as follows:

Body in 3, including legs, mainly black, the legs sometimes however

slightly more brownish or even yellowish brown, especially the front and middle

ones; the following parts in 9 very pale yellowish or ivory yellowish: sides

of frons, genae, head below narrowly on sides along eye margin to about or

nearly halfway up behind eyes in occipital part, sides of thorax above (except

for a gap just above wing-bases), entire discal part of scutellum, propleural

callosity above front coxae, a broadish pleural fascia along nearly upper half

of sternopleuron across lower hinder part of pteropleuron up to base of halteres,

metapleural part at same level, almost entire tergites 2 and 3, very broad hind

margins of rest of tergites, and the very broad hind margins of sternites; buccal

rim in dark or blackish; legs in 9 almost entirely yellowish, only the tarsi,

or at least their apical halves, darkened.

Vestiture with the hairs and scaling mainly whitish, longer and denser in

dg, more sericeous yellowish on abdomen above in 9.

Head with the front part not tumidly prominent or swollen; antennal

joint 1 very short, transverse, about as long as, or only very slightly longer

than, 2; joint 3 not distinctly spindle-shaped, only a little narrowed apically

and basally, with distinct bristly hairs above, its lower apical process quite

prominent, even a little more prominent than upper one, the two together

however forming a distinct bifid process; proboscis about 2-2,4 mm long.

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE 265

Wings distinctly, though faintly, dusky, tinted yellowish brownish sub-

apically to even brownish throughout, distinctly more so than in any other

known South African species; veins yellowish-brown or reddish-brown; knobs

of halteres darkened above, sometimes more so in Q.

Hypopygium of 3 as that figured in figure 252 (Hesse 1938: 840) erroneously

referred as belonging to pubescens Bezzi.

From 3 g¢ and 2 99 (3 holotype in British Museum (Natural History),

2 allotype in South African Museum).

Length of body: about 3,5 mm

Length of wing: about 44,5 mm

Distribution

Natal: Weenen (Thomasset, March-April 1924) (¢ holotype, 3 paratype,

and 9 paratype); Weenen (Thomasset, November 1928—March 1929) (9 allo-

type). Transvaal: Potchefstroom (Zumpt, 26 December 1952) (3 paratype).

From crocogramma Hesse, which it very closely resembles, it may at once

be distinguished by the distinctly more dusky wings, relatively shorter discoidal

cell, darkened halteral knobs, yellowish legs in 9 and not distinctly yellowish

knees in 3, much shorter proboscis, more symmetrical bifid apex of antennal

joint 3 in which the upper apical process is not so markedly subapical in position.

The 2 which also superficially resembles /aeta Bezzi, may at once be

distinguished from the latter by the distinctly dusky wings, the distinctly

narrower and less transverse scutellum and which is entirely yellow on disc,

the narrower and more humped body, less extensive yellow on pleurae, etc.

Phthiria nitidigena sp. nov.

Fig. 1B

Body in ¢ entirely black; legs sometimes more dark sienna brownish to

blackish-brown; integument of body, especially above and especially frons,

genae, head below, thorax above, scutellum, and abdomen above, markedly or

brilliantly shining, without any greyish white pruinescence.

Vestiture, as in most other ¢¢ of Phthiria, longish, but not very dense;

hairs on ocellar tubercle, frons, and genae black or dark; short ones on abdomen

above and longer ones on genital structures also dark; longer and sparser ones

on abdomen above and on scutellum pale, slightly straw-coloured yellowish;

rest of hairs on body and legs, especially denser ones on head below, sides of

tergite 1 and on venter, more whitish; fine hairs or hair-like scaling on tibiae

dark, and fine longish hairs on apical parts of femora also appearing dark in

certain lights.

Head (cf Fig. 1B) with the front part rather protruding spout-like, the

frons and genae broad, convex, tumid, much like those of Janigera, but to a

lesser extent; antennal joint 1 short, transverse, not longer than 2; joint 3

more parallel-sided than in most other species, equally broad throughout, not

266 ANNALS OF THE SOUTH AFRICAN MUSEUM

humped or spindle-shaped, with some distinct fine hairs above, ending apically

in a slightly longer lower process and a shorter upper process which is slightly

farther back; palps apically slightly spear-blade-shaped; proboscis slender,

about 2,4-2,7 mm long.

Wings very feebly greyish hyaline, without a very distinct subopacity,

iridescent; veins pale yellowish brownish; middle cross vein a little beyond

middle of discoidal cell; knobs of halteres entirely whitish.

Legs with the basal joint of hind tarsi distinctly much more than half

length of hind tibiae.

From 3 $4 in the South African Museum.

Length of body: about 4 mm

Length of wing: about 4,25-4,5 mm

Distribution

South-western Cape: between Leipoldtville and Elands Bay (Museum

expedition, November 1948).

The markedly shining frons, genae, scutellum and upper parts of the body,

spout-like anterior part of the head, dark hairs on frons and genae, and the

parallel-sided third antennal joints easily distinguish this species from most of

the known South African species.

Phthiria lanigera Bezzi

Phthiria lanigera Bezzi, 1921: 97. Hesse, 1938: 840, fig. 253.

This characteristic Cape species, which appears to be a montane species,

is subject to variation and a distinct variety of it, described below, has since

been taken together with the more typical form.

Phthiria lanigera var. melampogon n.

This variety differs from the typical form of Janigera in the following

respects:

Vestiture with the hairs on frons, genae, and antennal joints | and 2 above

in ¢ distinctly very much denser and longer, those on frons, antennae above,

and upper part of genae black (those on frons and entire genae in typical ¢

distinctly less dense and entirely white); hairs on upper part of genae in also,

very much denser, black, not white, and extreme upper part of genae on sides

of antennae, though also black and shining, not entirely smooth as in /anigera,

but with some black hairs; hairs and woolly scaling on body above in varietal

@ paler yellowish sericeous or pale brassy yellowish, not so golden or deep

golden yellowish.

Wings distinctly and more conspicuously milky whitish; veins also distinctly

paler, more yellowish brown or yellowish, not brown or blackish brown;

knobs of halteres in 9 ivory whitish or yellowish above.

The varietal form also has antennal joint 3 relatively thinner or more

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE 267

slender basally, and the knees are either not, or scarcely, or at least less con-

spicuously, yellowish.

From 5 33 and 9 99, including the ¢ holotype and 9 allotype, in the

South African Museum.

Length of body: about 5,12-6,4 mm

Length of wing: about 5,4-5,52 mm

Distribution

South-western Cape: Wit River Valley in Bainskloof near Wellington

(Museum expedition, December 1949) (¢ holotype, 2 allotype, and paratypes);

Wit River, Bainskloof (Barnard and Lawrence, November 1922); Wit River,

Bainskloof (Barnard, November 1937) (¢ and @ paratypes).

Phthiria nigribarba Hesse

Phthiria nigribarba Hesse, 1938: 843, fig. 255.

Of this species, which I described from only 2 33 in 1938, the South

African Museum has since obtained not only a number of other 3, but also

a series of the as yet undescribed 9. Supplementary notes on the species and

a description of the 2 can now be added as follows:

Body entirely black in both sexes, appearing greyish white, especially

markedly so in 9, due to a conspicuous greyish white pruinescence; hind

margins of sternites sometimes, even in dd, very obscurely and narrowly

pallid; basal part, or half, of palps yellowish in 2 to yellowish brownish or

sienna brownish in J; legs variable in colour, varying from entirely yellowish

in some 99 to yellowish tibiae and tarsi, and yellowish brown, brown, to sienna

brownish femora in other 99, and from entirely dark reddish or sienna brownish

femora with yellowish knees in some ¢<¢ to paler yellowish brown in others.

Vestiture with the hairs rather dense and long in both sexes, denser and

shorter on body in 9, on the whole much like those of Janigera; hairs in upper

part of occiput, on ocellar tubercle, frons, and genae much denser and longer

in 3 and black, entirely sericeous white in 9 like rest of hair on body and legs

in both ¢ and 9.

Head with the anterior part in $ tumidly prominent and swollen as in

3 of lanigera; this part in less swollen, but with the upper part of genae’

and the frons nevertheless broader and more convex than in other species,

excepting those in the Janigera section; antennal joint 1 short, subequal to,

or scarcely longer than, 2; joint 3 spindle-shaped or humped, without distinct

hairs above, its upper apical process distinctly longer than lower one, slightly

curved, but relatively less stout than in J/anigera; proboscis about 2-2,6 mm

long, stouter and relatively shorter in 2 than in d.

Wings conspicuously milky whitish in both sexes, with very pale yellowish

or pallid, to almost white, veins; middle cross vein much beyond middle of

discoidal cell; veins enclosing anal cell markedly straight; anal cell with its

268 ANNALS OF THE SOUTH AFRICAN MUSEUM

apical stalk relatively long in both sexes, usually longer than apical cross vein

of discoidal cell; knobs of halteres entirely pallid or very pale in both sexes.

Length of body: about 3,5—5 mm

Length of wing: about 3,5—5 mm

Distribution

South-western Cape: Malmesbury (Brauns, October 1926) (type material);

Klipheuwel near Cape Town (Zinn, October 1950) (gg and 929 and @ allotype).

Phthiria ovalicornis sp. nov.

Fig. 1E

A species very near nigribarba and also belonging to the Janigera section.

It agrees with and differs from nigribarba in the following respects:

Body also entirely black in both sexes, also covered with a very conspicuous

greyish white pruinescence in 9; palps dark; legs much darker, even in 9,

blackish brown to black, sometimes more sienna brownish in 9, only the knee-

articulations sometimes reddish brown.

Vestiture very similar to that of nigribarba, but the hairs in @ distinctly

denser than in 99 of latter; dark hairs on ocellar tubercle, frons and genae in

¢ distinctly less dense and shorter, and in 2 much sparser and shorter in this

species, but also entirely sericeous white, like rest of hair on body and legs,

in both sexes; hairs in occiput in ¢ also whitish, not dark as in ¢ of nigribarba.

Head in ¢ in front, especially genal parts, very similar to that of nigribarba

and Janigera, but somewhat less tumidly prominent, less evident in 2 where

the genae are much shorter than in nigribarba, though the frons is broader,

more convex than in 9° of most species of Phthiria; antennal joint 1 (cf. Fig. 1E)

also short, not visibly longer than 2, sometimes tending to be even shorter;

joint 3 (cf. Fig. 1E) relatively shorter, characteristically oval or spindle-shaped,

broadest at about, or just before, middle, slightly hollowed or depressed on

inner side from just before middle and with some fine hairs above, more distinct

in 3g, the upper apical process of joint relatively shorter than in nigribarba,

but nevertheless stouter and slightly longer than lower one; palps relatively

shorter, less projecting; proboscis 1,8—2,4 mm long.

Wings also conspicuously milky whitish, with yellowish veins which are

however sometimes less pallid or whitish; second submarginal cell markedly

acute basally, more so than in any other of the known South African species;

middle cross vein much beyond middle of discoidal cell; knobs of halteres

entirely pallid or whitish, but in some ¢¢ tending to be darker above.

Legs with the basal joint of hind tarsi distinctly very much longer than

half length of hind tibiae, much more so than in nigribarba.

From 12 $3 and 12 99 (¢ holotype, 2 allotype and ¢ and @ paratypes)

in the South African Museum.

Length of body: about 3,5-4,5 mm

Length of wing: about 44,5 mm

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE 269

Distribution

North-western Karoo: Augusfontein near Calvinia (Museum expedition,

September 1947).

Phthiria brunnescens sp. nov.

Fig. 1F

Another species in the Janigera section characterized as follows:

Body in ¢ entirely black; legs very dark blackish brown or dark sienna

brownish, only the knee-articulations paler, more yellowish brown.

Vestiture with the hairs dense in g, but rather short, differing from 33

of all other known South African species in having not only the hairs on head

and occiput entirely black, but also dark ones intermixed on thorax above and

on scutellum, short dark hairs on abdomen above, and blackish brown or dark

ones on coxae and legs; the paler hairs on thorax above, to a certain extent on

pleurae, and on sides of abdomen beyond tergite 1 gleaming more yellowish to

pale yellowish brownish; with even the more whitish ones on mesopleuron, sides

of tergite 1, and on venter gleaming more sericeous yellowish in certain lights.

Head with the frons and genae slightly tumidly prominent or inflated,

placing the species at once into the Janigera and nigribarba section; antennal

joint 1 short, subequal to, or only very slightly longer than, joint 2; joint 3

(cf. Fig. 1F) humped as in the /anigera section, but slightly beyond the middle,

with distinct and conspicuous hairs above, ending apically in a slightly curved

process, slightly longer and stouter than lower process; proboscis about 2

mm long; palps rather long, longer than antennae.

Wings greyish hyaline, with a slight subopacity, but not conspicuously

milky whitish as in the Janigera section; veins yellowish brown; middle cross

vein at about, or only a little beyond, middle of discoidal cell; knobs of halteres

darkened above.

Legs with the basal joint of hind tarsi distinctly much more than half

length of hind tibiae.

From a 3 specimen in the South African Museum.

Length of body: about 4 mm

Length of wing: about 4 mm

Distribution

South-western Cape: Citrusdal District (Museum expedition, November

1948).

Phthiria namaquensis sp. nov.

Fig. 1G

A species which is also referable to the /anigera section on account of the

structure of its third antennal joint and its milky whitish wings, but in which

the head in front is not markedly inflated or tumidly prominent. It is charac-

terized as follows:

270 ANNALS OF THE SOUTH AFRICAN MUSEUM

Body and legs black in 3 and also mainly so in 9; the following parts in

2 pallid or yellowish: a short transverse streak on each side in frontal depression,

more or less lower half of genae, continued as a narrow streak along margin

of eyes on head below and halfway up behind eyes in occipital part, the small

boss-like prominence in middle of pleurae between sterno- and hypopleurae

and pteropleuron and sometimes to a certain extent even adjacent parts of

sterno- and metapleurae, and very narrow hind margins of sternites; knees in

both sexes pallid or yellowish; integument of head and body in both sexes

densely covered with greyish white bloom, on thorax above in longitudinal

streaks and on abdomen above, especially in 3, as greyish white rings across

hind margins of tergites.

Vestiture with the hairs fairly dense, more so and longer in 3, but relatively

less so than in other species of this section, entirely sericeous whitish, but in

¢ with some hairs in occiput above behind eyes, those on ocellar tubercle and

frons dark and, in 9, also with some hairs on frons in front dark.

Head with the anterior part in ¢ not so obviously tumid as in nigribarba

or even brunnescens, but with the frons and upper part of genae, even in Q,

slightly broader or more convex than normally so; antennal joint | very short,

transverse, subequal to joint 2; joint 3 (cf. Fig. 1G) more or less subparallel-

sided, not distinctly humped, without distinct hairs above, its upper apical

process slightly, but distinctly, longer and stouter than the conspicuous lower

one; proboscis about 1,6—2,2 mm long, not very much stouter in 9 than in d.

Wings with a distinct milky whitish tint; veins brownish or dark brown;

middle cross vein a little beyond middle of discoidal cell; apical stalk of anal

cell, especially in 9, tending to be shorter than in most other species; knobs

of halteres, even in 9, darkened above to a variable extent.

From 4 gd and 4 99, including the types, in the South African Museum.

Length of body: about 3-4 mm

Length of wing: about 3,5-4 mm

Distribution

Namaqualand: Papendorp (Museum expedition, October 1950) (¢ holo-

type and 9 allotype); Wallekraal (Museum expedition, October 1950) (3 ¢

and 3 2 paratypes).

Phthiria conocephala sp. nov.

Fig. 1D

A species also belonging to the /anigera section as far as the shape of the

third antennal joint, the milky whitish tinted wings and rather dense vestiture

are concerned, but the front part of head is very similar to that of pilirostris

It is characterized as follows:

Body and legs in 2 mainly black; a narrowish streak on each side of frons,

more than lower half of genae, entire head below, excepting only a central

dark streak, pale yellow, this pale yellow also continued as a narrow streak

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE aga

behind eyes, halfway up occiput and then continued as hind margin of eye

itself, as a propleural spot above front coxae, a spot next to it just behind base

of anterior coxae, a streak along middle of pleurae from posterior upper part

of sternopleuron, lower part of pteropleuron, across upper part of hypopleuron,

metapleural part in front of and just below halteres, and to a lesser extent

also upper anterior angle of pteropleuron below wing-base; small area below

wing-base ferruginous.

Vestiture with the hairs and scaling relatively dense and long, especially

on genae, head below, on abdomen, and on femora, predominantly sericeous

or silvery whitish, but those on front part of frons and upper parts of genae

dark or black.

Head (cf. Fig. 1D) front markedly conically produced and narrowed

apically, more so than in 99 of /anigera and nigribarba section, more like that

of 9 pilirostris; buccal cavity sloping backwards; antennal joint | a little, but

distinctly, longer than 2, about 1} times length of 2; joint 3 not very distinctly

humped, only slightly broadened beyond middle, without distinct hairs above,

its upper apical process stouter and longer than the lower feeble prominence,

slightly curved down; proboscis about 2,2 mm long.

Wings distinctly tinted milky whitish; veins brownish or yellowish brown;

middle cross vein much beyond middle of discoidal cell; apical stalk of anal

cell rather long, very slightly longer than apical vein of discoidal cell; squamae

conspicuously pallid or yellowish white; knobs of halteres with a small dark

spot basally above.

Legs with the basal joint of hind tarsi only about half length of hind tibiae.

From a single 2 specimen in the South African Museum.

Length of body: about 4,4 mm

Length of wing: about 4,2 mm

Distribution

Coastal part of western Cape: Graafwater (Museum expedition, October

1947).

REVISED DESCRIPTIVE KEY TO THE KNOWN SOUTH AFRICAN SPECIES OF Phthiria

1 (a) Proboscis without any hairs or dense hairs on labral part; apical part of head beyond

anterior level of eyes to antennal insertions distinctly less conically prominent,

produced, or narrowed, or pointed apically, appearing less tilted upwards; buccal

cavity not straightly and steeply sloping backwards; abdomen without or with

comparatively much narrower yellowish hind margins to tergites and, if more

broadly yellowish, proboscis without hairs above ; 2

(6) Proboscis with distinct, conspicuous, fairly dense, and longish hairs ¢ on basal half

above on labral part; apical part of head beyond anterior level of eyes to antennal

insertions more markedly conically prominent, produced and narrowed or more

pointed apically, appearing more tilted upwards; buccal cavity distinctly more

straightly and steeply sloping obliquely backwards; abdomen with relatively broad

and conspicuous yellowish hind margins to both tergites and sternites

3 pilirostris Hesse

212

2° @)

(5)

4 (a)

(b)

(5)

6 (a)

(d)

7 (a)

ANNALS OF THE SOUTH AFRICAN MUSEUM

Antennal joint 3 more spindle-shaped, or tending to be more or less equally broadened

above and below at broadest part, appearing less distinctly humped, ending apically

in an upper and lower spine-like process, with either both tending to be equally

developed, together forming a more symmetrical bifid process, or the lower one is

distinctly stronger, more prominent; head in front of eyes less markedly tumid or

inflated in appearance, the frons being narrower and upper parts of genae also

relatively narrower, less bulging or inflated; wings, even if appearing slightly hyaline,

tinted cinereous or even dusky, without, or with only a very feeble, subopaquely

milky whitish tint; hairs on head and body on the whole less dense, usually shorter

and often sparse, those on legs also shorter, less dense and often poorly developed

or almost absent a

Antennal joint 3 tending to be more distinctly humped above, sometimes con-

spicuously so, ending in a distinctly stronger, longer, and sometimes curved down,

upper spine-like process which is always stronger than the feebler or much reduced

lower one, the two together not symmetrically bifid; head in front usually distinctly

more markedly tumid or inflated, the frons relatively broader, more convex, and

upper genal parts also broader, more bulging and inflated and, if not, other characters

conform; wings usually clear hyaline, rarely without a very distinct, or even con-

spicuous, subopaquely milky whitish tint; hairs on head and body, also legs, on

the whole much denser, more shaggy and often longer .. ay, =f le

Last sternite mainly yellowish or more conspicuously yellowish; legs entirely or

predominently pale oer or at least tibiae and bases of tarsi mainly pale

yellowish

Last sternite entirely dark or ede, or not distinetly veltoeish legs entirely black

or dark, or more yellowish brown or brownish than pale yellowish, only knees

sometimes narrowly yellowish or pallid .. ad ; si a zis a

Entire femora, tibiae, and bases of tarsi yellowish. . gd un Bezzi var. xerophila Hesse

Only apices or apical parts of femora, entire middle and hind tibiae, upper surfaces

of front tibiae, and bases of tarsi yellowish .. 6 pulla Bezzi var. fallax Hesse

Body relatively broader and thorax distinctly less convex above, less humped;

scutellum markedly transverse, distinctly very much broader than long, at least

twice as broad as long; basal joint of front and hind tarsi, especially of latter, only

about, or only a little more than, half length of hind tibiae; knobs of halteres

darkened above in apical half Bi os 6 laeta Bezzi (nec Hesse 1938: 826)

Body distinctly narrower, the thorax distinctly more convex, more humped; scutellum

distinctly less transverse, very much narrower, more conically narrowed apically,

more convex and, though broader than long, usually considerably less than twice

as broad as long; basal joint of front and hind tarsi, especially of the latter, much,

or very much, more than half length of tibiae; knobs of halteres more extensively,

or entirely, pale above and, if dark above, other characters conform .. A 6

Anterior part of head with the frons and genal parts narrower and shorter, less

spout-like; antennal joint 3 usually broadened at about, or a little beyond, middle,

more spindle-shaped or humped; apical joint or part of palps only slightly broadened,

not spindle-shaped or spear-blade-shaped; frons and genal parts not entirely or

conspicuously shining, usually duller, or in part covered with whitish or greyish

tomentum, and body also distinctly less shining or predominently duller due to

greyish white tomentum; hairs on frons and genae, and fine ones on abdomen above,

mainly pale or whitish ie ae 7

Anterior part of head with the frons and genal parts broader and longer, more

protruding spout-like; antennal joint 3 more parallel-sided or equally broad; apical

joint or part of palps distinctly more spear-blade-shaped; frons and genal parts,

and even head below, more conspicuously brilliantly shining and body above,

including abdomen, distinctly more shining, without greyish tomentum; hairs on

frons and genae, and fine ones on abdomen above, dark or black. . ¢ nitidigena sp. nov.

Wings clearer, more vitreous hyaline, without a very distinct subopacity and, if SO,

this is only very faint and scarcely perceptible, milky whitish; middle cross vein

tending to be nearer middle of discoidal cell; latter relatively shorter, only a little

(5)

(a)

(5)

(a)

(5)

(a)

(6)

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE 273

more than twice as long as broad; head in front and body above much duller, less

shining; antennal joint 3 without distinct hairs above, usually with the apical pro-

cesses more equally lik ai knees narrowly, or to a variable extent, pallid or

yellowish ae 8

Wings less clear, with a more distinct yellowish whitish, cinereous 5 yellowish or

greyish brownish subopacity, sometimes even uniformly dusky; middle cross vein

distinctly more, or even much, beyond middle of discoidal cell; latter relatively

much longer, much more than twice, to nearly or even about 3 times, as long as

broad; head in front, especially upper genal parts, and body above on the whole

tending to be more shiny; antennal joint 3 usually with some distinct, or even con-

spicuous, hairs above, usually with the lower apical process distinctly more

developed; knees not always yellowish and, if so, other characters do not differ.. 9

Antennal joint 1 short, transverse, slightly shorter than, subequal to, or scarcely

longer than joint 2; proboscis longer, usually more than 1,5 mm; knobs of halteres

entirely pale or pallid above, or only darkened to a very limited extent; sides of

tergites with broad bands of greyish tomentum across apical halves; last sternite

tending to be reddish or yellowish to a variable extent .. aE 3 pulla Bezzi

(syn. flavigenualis Hesse)

Antennal joint 1 distinctly longer, elongate, nearly or quite twice length of small

transverse joint 2; proboscis shorter, only about 1,5 mm long; knobs of halteres

entirely dark or blackish above; sides of abdomen or tergites without distinct, or

at least conspicuous, transverse apical bands of greyish tomentum; last sternite

entirely dark .. oh i i 3 pulla var. anomalocera n.

Wings more elongate, clearer ereyish hyaline, with a slight whitish or yellowish

whitish subopacity; veins paler, more yellowish; middle cross vein more beyond

middle of discoidal cell which itself is relatively longer; proboscis longer, about

3—3,5 mm long; knees distinctly contrastingly yellowish; buccal rim and genae not

separated by a very distinct groove ‘ 6 crocogramma Hesse

Wings relatively shorter, distinctly slightly infuscated, dusky or tinted yellowish

brownish to even brownish throughout; veins darker, more reddish brown; middle

cross vein only a little beyond middle of discoidal cell which itself is relatively

shorter; proboscis shorter, relatively stouter, less than 3 mm long; knees not pallid

or yellowish, but front and middle legs sometimes yellowish brownish; buccal rim

and genae separated by a more distinct groove .. 4 6 tinctipennis sp. NOV.

(syn. 3 pubescens Hesse nec Bezzi)

Anterior part of head, comprising frons and genae, distinctly much more prominent,

more conspicuously inflated or tumid, the frons and genae being also distinctly

very much broader; antennal joint 3 markedly broadened, distinctly more humped,

its upper apical spine distinctly more developed, much longer and stouter than

lower process and usually more curved downwards; hairs on frons, genae and on

body as a whole distinctly denser, often more woolly; whitish or greyish pruinescence

on body less developed; knees, excluding the articulations, not pallid or yellowish;

last sternite longer, more elongate and scoop-like as BR ew

Anterior part of head distinctly less prominent, less conspicuously tumid or inflated,

the frons and genae being distinctly much narrower; antennal joint 3 subparallel- _

sided, or at least not so conspicuously broadened or humped, its upper apical spine

only slightly, though distinctly, stronger than the relatively well-developed lower

one; hairs on frons, genae, and body slightly less dense; whitish or greyish pruine-

scence on body on the whole more conspicuously developed; knees, including

articulations, more distinctly pallid or yellowish; last sternite distinctly shorter

3 namaquensis sp. Nov.

Wings distinctly and conspicuously subopaquely milky whitish; middle cross vein

very much beyond middle of discoidal cell; hairs on body and legs, excepting only

on head in front in some forms, entirely or predominently very pale or whitish;

antennal joint 3 usually humped at middle, or only a little beyond middle .. 12

Wings greyish hyaline, without any, or with only a feeble scarcely perceptible,

whitish subopacity; middle cross vein at about, or only slightly beyond, middle of

discoidal cell; hairs on head and occiput, intermixed ones on thorax, on scutellum,

274

12 (a)

13 (a)

(0)

14 (@)

(0)

15 (@)

(0)

ANNALS OF THE SOUTH AFRICAN MUSEUM

disc of abdomen above, coxae, and legs very dark or black, those on thorax above

and to a certain extent on pleurae, and on sides in apical half of abdomen gleaming

more yellowish or yellowish brownish; antennal joint 3 tending to be humped

Nearer apex =. : 3 brunnescens sp. nov.

Antennae longer, stouter, with joint® 1 distinctly much longer, at least 14 to 2 times

length of 2, with longish and dense hairs on both 1 and 2; joint 3 characteristically

and prominently arched or humped at about, or just beyond, middle, with con-

spicuous hairs above, ending apically in a strong, finger-like, curved down, upper

process, dwarfing the slight lower apical prominence; wings rather more pointed

apically, the upper cubital branch ending apically at wing-point; veins dark brown

to blackish brown; knobs of halteres more extensively darkened or black above;

hairs on head, body, and legs distinctly and markedly much denser, longer, and

more woolly or shaggy : ers

Antennae proportionally shorter, with joint 1 very much shorter, scarcely, or only

a very little longer than, or subequal to, as long as, or even a little shorter than

joint 2, with very much shorter, or only very short and less dense, hairs on 1 and 2;

joint 3, though also humped or broadened, less strongly developed, without any, or

with much feebler and shorter, hairs above, ending apically in a less stout and

shorter process, less markedly curved down, forming a more bifid process with an

also relatively prominent lower process; wings more rounded apically; veins paler,

more yellowish or pallid and, if dark, other characters conform; knobs of halteres

either entirely yellowish or pallid above, or with less extensive darkening above;

hairs on head, body, and legs distinctly shorter, less dense, less woolly or shaggy.. 14

Hairs on frons and genae entirely white and distinctly less dense; antennal joint 3

tending to be thicker basally; veins in wings darker; knees more distinctly, even

Hairs on frons, antennae above, and upper halves of genae black and distinctly

denser; antennal joint 3 tending to be thinner basally; veins in wings tending to be

paler, more yellowish or with more yellowish; knees not perceptibly yellowish

6 lanigera var. melampogon n.

Second marginal cell in wings not acuminate or acute at base; anterior part of head

distinctly more bulging, inflated, or tumidly prominent, the frons and genae being

longer (or broader), length of genae at broadest part on sides of head quite, or

nearly, half length of eye at same level; antennae relatively longer, joint 3 distinctly

and relatively longer, more spindle-shaped, its upper apical process longer and

stouter relative to lower one to a variable degree; palps proportionally longer, more

projecting; hairs on head in front and on body distinctly denser and longer;

scutellum on the whole more acute or pointed apically; basal joint of hind tarsi

about, or only a little more than, half length of hind tibiae .. sik ee

Second submarginal cell distinctly more acute or acuminate at base; anterior part

of head distinctly less prominently inflated or tumid, the frons and genae being

shorter (narrower), the length of genae at broadest part on sides of head very much

less than half, only about, or nearly a little more than, a third length of eye at same

level; antennae relatively and proportionally shorter, joint 3 distinctly more oval

or slightly elongate-oval, its upper apical process scarcely, or only a very little, longer

than lower one, the two together forming a more bifid process; palps proportionally

much shorter, less projecting; hairs on head, body, and legs distinctly shorter, less

dense; scutellum more rounded apically; basal joint of hind tarsi distinctly much

more than half length of hind tibiae a bs 3 ovalicornis sp. NOV.

Frons and genae with dark blackish brown or black hairs; palps and legs slightly

paler, more pale yellowish brown to sienna brownish; proboscis longer, about

2-2,5 mm; veins in wings pallid or yellowish and those between anal and axillary

cells and anal and fourth posterior cells markedly straight; apical stalk of anal cell

on the whole longer, subequal to, or as long as, apical cross vein of discoidal cell;

antennal joint 3 shorter, broader, more spindle-shaped, its upper apical process on

the whole stouter, blunter, scarcely or not much curved and the lower apical part

more prominent 3 nigribarba Hesse

Frons with dark, or some dark, hairs, but genae entirely white-haired; palps and

tt (@)

(0)

2 3@)

(6)

3 (a)

(5)

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE 275

legs much darker, darker brownish or black; proboscis shorter, only about 1,5 mm

long; veins in wings darker and those between anal and axillary and anal and fourth

posterior cells not entirely or so markedly straight; stalk of anal cell shorter than

apical cross vein of discoidal cell; antennal joint 3 more elongate, slightly more

than twice as long as joints 1 and 2 combined, not so obviously spindle-shaped, its

upper apical process slightly longer, more slender, more acute apically, and more

curved downwards to the scarcely prominent lower apical angle ¢ simmondsii Hesse

Proboscis without any hairs or dense hairs on labral part; apical part of head beyond

anterior level of eyes to antennal insertions less conically produced, prominent,

narrowed, or pointed apically, appearing less tilted upwards, the buccal cavity not

straightly and steeply sloping backwards and, if rarely so, proboscis without hairs

above; body not predominently, or almost entirely, yellowish above and below

and, if greater part of body is not black, the thorax above and basal halves of tergites

at least black or more extensively black we 3 2

Proboscis with distinct, conspicuous, fairly dense and longish, yellowish or golden:

gleaming hairs on at least basal half of labral part above; apical part of head,

beyond eyes to antennal insertions, distinctly more markedly conically prominent,

produced, narrowed, and more pointed apically, appearing more tilted upwards,

the buccal cavity distinctly more straightly, steeply sloping obliquely backwards;

body predominantly, almost entirely yellow or yellowish above and below, even

thorax above with only three reddish brown or brownish fasciae, and only extreme

bases of tergites discally above and an infusion on hypopleural part dark

° pilirostris Hesse

Antennal joint 3 more spindle-shaped, usually more equally broadened above and

below at broadest part, appearing less distinctly humped, ending apically in an

upper and lower spine-like process either more or less equally prominent and

together forming a more or less bifid process, or with a tendency for lower one to

be stronger; head in front less tumidly prominent, the frons distinctly less broadly

transversely convex and upper parts of genae not so continuously tumid or convex

with frontal convexity; wings, even if with some degree of subopacity, not distinctly

tinted subopaquely milky whitish; vestiture on body and legs distinctly less dense,

much sparser, or shorter, often almost wanting; body usually with extensive yellow,

rarely mainly black .. a 3

Antennal joint 3 tending to be more distinctly humped above, sometimes con-

spicuously so, ending in a distinctly longer or stronger, sometimes curved down,

upper process and a more reduced, sometimes scarcely detectable, lower prominence,

rarely with both more or less equally developed and, if so, other characters listed

here do not differ; head in front slightly or distinctly more tumidly prominent or

protuberant, the frons distinctly more convex, more transversely so, relatively

broader, and upper parts of genae also relatively more convex or tumid and con-

tinuously so with frons and, if not apparent, other characters conform; wings rarely

not distinctly, often conspicuously, tinted subopaquely milky whitish; vestiture

on body and legs usually distinctly more conspicuous, denser, and usually longer; ~

body mainly black or dark, without any, or with very little, yellow present .. 13

Body with distinctly much or more yellow, the head, at least sides of thorax above,

scutellum, hind margins of tergites, pleurae, venter, and sometimes legs con-

spicuously yellow or orange yellow to a variable, greater or lesser, extent; genae,

head below, scutellum, and rest of body above not, or distinctly less, shiny, not

tending to be brilliant; wings usually more one and, if distinctly tinted greyish

or dusky, body with much yellow

Body entirely black, without any yellow markings, the legs entirely very dark blackish

brown or black, and even knees not markedly paler; genae, head below, scutellum,

and to a certain extent integument of rest of body above distinctly more brilliantly

shining and smooth; wings distinctly, though faintly, tinged slightly subopaquely

greyish yellowish or cinereous yellowish, even with a faint brownish tint in certain

lights .. cp 3 #4 ri ay os “i .. Q pubescens Bezzi

276

4 (a)

(d)

(6)

6 (a)

(d)

TG)

(b)

8 (a)

ANNALS OF THE SOUTH AFRICAN MUSEUM

Legs, including coxae, entirely or predominantly pale yellowish or at least mainly,

or more extensively yellowish; almost entire prosternal part, pleural parts below

wings, and hind margins of sternites usually entirely, predominantly, or at least

more extensively yellowish and, if not, legs are yellowish; upper apical process of

antennal joint 3 either nearer apex or almost as long as or as strongly developed as

lower one, the two together forming a more symmetrical bifid process and, if not,

legs at least mainly yellowish ; : 5

Legs, including coxae, entirely very dark, ‘blackish brown, or black, or with much

black, only the knees sometimes pallid or yellowish; prosternal part, pleural parts

below wings mainly dark or black, and hind margins of sternites, if not black,

usually more obscurely, or only narrowly, yellowish; upper apical process of antennal

joint 3 either farther back from apex or smaller, shorter, and less developed than

lower one, the two together usually forming a more asymmetrical bifid process.. 10

Body relatively broader and thorax distinctly less humped; head and frons relatively

much broader; scutellum markedly or more obviously transverse, distinctly very

much broader than long, quite, or more often, a little more than twice as broad as

long; basal joint of hind tarsi only about, or only a little more than, half length of

hind tibiae; front femora above and outer hind faces of tibiae more distinctly

darkened aa : 6

Body relatively narrower, ine thorax diswncily more convex above, more humped:

head and frons narrower; scutellum less transverse, relatively or distinctly much

narrower, more conically narrowed posteriorly and, though broader than long,

usually considerably less than twice as broad as long; basal joint of hind tarsi dis-

tinctly very much more than half length of hind tibiae; front femora and outer

faces of tibiae not, or scarcely, darker and, if much darkened, other characters

do not differ .. xe t ae 8

Broadish hind border, or aliiost hinder half, and somietinries narrow ee of scutellum,

black; yellow hind margins of tergites distinctly broader, occupying almost, or

about, the apical halves of tergites on sides; yellow on sides of thorax above relatively

narrower; lateral extension of black frontal fascia on frons anteriorly more constant

and broader, usually reaching eye; hairs on tergites distinctly less dense, relatively

shorter, and those discally more whitish; fine hairs on frons also less dense, more

whitish; tarsi tending to be paler, or more yellowish, basally or in basal part

2 laeta Bezzi

Entire scutellum discally yellow; yellow hind margins of tergites distinctly narrower,

occupying less than apical halves of tergites on sides; yellow on sides of thorax

above usually broader; black medial frontal fascia either not extending laterally

on frons in front, or the extension is narrower or spot-like; hairs on tergites distinctly

denser, relatively longer, and distinctly sericeous yellowish to pale golden discally;

fine hairs on frons denser, more golden; tarsi tending to be entirely dark or black .. 7

Antennal joint 3 ending apically in two almost equally strong upper and lower

processes, the upper one scarcely much smaller than lower one and tending to be

nearer apex, the two together forming a distinctly more symmetrical bifid process;

upper surface of front femora and sometimes a subapical infusion on inner or upper

apical part of the others and outer faces of all the tibiae more conspicuously or

intensely darkened; knobs of halteres with a dark spot or darkened on outer margin

or sides above . @ flaviscuta sp. nov. (and forms of it)

Antennal joint 3 ending apically i in 1 two “unequally developed processes of which

the upper one is much weaker and thinner than the much stouter lower one and also

farther from the apex, the two together forming a distinctly more asymmetrical

bifid process; upper surface of front femora and outer surfaces of all the tibiae not,

or only feebly and less conspicuously, darkened; knobs of halteres not distinctly

darkened in part or with a spot above... = .. 9 hesperia sp. nov.

Wings vitreous hyaline, without any, or abarckive any, detectable subopacity; black

on head more reduced, that on occiput less extensive, that on head above more or

less narrowly confined to ocellar tubercle, or as a narrow line towards middle, and

as a spot anteriorly on each side of antennae, without a broadish central band on

head below, and buccal rims and buccal cavity also pale; yellow on pleural parts

(6)

9 (a)

(0)

10 (a)

(0)

Ae x(a)

(b)

2) .@)

(d)

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE pars |

distinctly much more extensive, greater part of pleurae above coxae being entirely

or predominantly yellow; yellow on sides of thorax broader, more extensive, and

with a tendency for two yellowish submedial lines above as well, and base of thorax

above yellowish or orange yellowish to a variable extent; scutellum entirely yellowish;

transverse black basal bands across tergites almost confined to middle above;

antennae with joint 1 more transverse, about as long as, or subequal to, joint 2,

usually yellowish or at least yellowish above, with joint 3 more ic ae te or

humped, without any distinct hairs above = g

Wings distinctly dusky, tinted subopaquely cinereous yellowish to brownish: black

on head distinctly more developed, almost the entire occiput and a broad central

fascia from it across ocellar tubercle to antennae black, without a spot on each side

of antennae, but with the buccal rims and buccal cavity dark, and a central broad

fascia on head below also black; yellow on pleural parts distinctly more reduced,

present only as a propleural spot, a central band across upper part of sternopleuron,

spots behind pteropleuron and on metapleural part; yellow on sides of thorax much

narrower, the entire discal part black, and base of thorax above black; scutellum

yellowish only on disc, its broad declivous sides black; transverse black basal bands

across tergites extending down sides as well; antennae with joint 1 entirely black,

slightly more elongate, tending to be slightly longer than 2, with joint 3 scarcely

or not broader across middle, not aay humped, and with distinct and con-

spicuous hairs above ne v4 ae 2 tinctipennis sp. nov.

Hairs on body, and especially on “Aapdlanaea: distinctly much sparser, less con-

spicuous, shorter, and sometimes almost absent. . 2 pulla Bezzi var. xerophila Hesse

Hairs on body, and Saag! on abdomen, distinctly much denser, longer, more

conspicuous .. ah .. @ pulla Bezzi var. fallax Hesse

Yellow or orange vellow transverse ands across hind margins of tergites distinctly

very much broader, occupying nearly, or even about, apical half of tergites; yellow

on sides of thorax relatively broader, the fascia across sternopleuron occupying

almost entire upper part of it, and scutellum with less black on declivous sides;

wings relatively broader and longer, without, or with scarcely detectable or feebler,

subopacity; proboscis longer, not shorter than 2mm .. See Valet

Yellow transverse bands across hind margins of tergites and sternites distinctly

very much narrower, those of tergites occupying considerably less than apical

halves; yellow on sides of thorax relatively narrower, the fascia across sternopleuron

occupying only hinder upper half, and scutellum with more extensive black on

declivous sides; wings relatively shorter and narrower, with a more conspicuous

subopaquely milky whitish tint; proboscis shorter, only about 1,8-2 mm

2 cognata Hesse

Proboscis distinctly stouter, longer, about 2,8-3,5 mm; occiput with some yellow

behind eye-margins, either on vertex or right around eyes; anterior spiracle with

more yellow around it; yellow hind margins of tergites broader, not much less than

apical half of tergites; legs on the whole darker, with broader yellowish knees;

wings with a distinct, though feeble, subopacity . . 12

Proboscis more slender, shorter, only about 2 mm n long: entire eeciput behind eyes

in at least upper half black; anterior spiracle without any, or with scarcely any,

yellow; yellow hind margins of tergites narrower; legs with more yellowish brown

or yellowish on femora, especially on inner upper surfaces on at least basal halves,

and front coxae pale in front, and knees not, or less broadly, yellowish; wings more

vitreous hyaline, without any detectable subopacity se 2 varipes Sp. NOV.

Thorax entirely black above; yellow on sides of thorax in front slightly narrower,

and scutellum with more black on declivous sides; yellow hind margins of tergites

_ Narrower, occuyping only about apical halves of tergites; frons with a consistent

broad central black band extending to antennae; wings clearer hyaline

typical 2 crocogramma Hesse

Base of thorax yellowish or yellowish-spotted to a variable extent; yellow on sides

of thorax relatively broader, and scutellum without, or with much less, dark on

declivous sides; yellow hind margins of tergites broader, occupying more than

apical halves of tergites; black fascia on frons much reduced, either wanting or

278

13 (a)

14 (a)

(0)

15 (@)

(d)

16 (a)

(0)

ANNALS OF THE SOUTH AFRICAN MUSEUM

thinning out anteriorly; wings with a slight, but distinctly more evident, subo-

pacity”... ab a se i ay A 2 var. of crocogramma Hesse

Antennae with joint 1 distinctly much longer, at least 14 times to nearly, or about,

twice length of 2, with joint 3 usually distinctly more humped, ending in a con-

spicuous, slightly curved, upper process which dwarfs the scarcely detectable lower

one; head in front distinctly more conically prominent or produced; head below,

propleural callosity above front coxae, and a longitudinal fascia along middle of

pleurae, ivory yellowish or yellow; hairs on head, body, and He distinctly longer,

markedly denser, and more woolly or shaggy .. ; » tary EE

Antennae with joint 1 much shorter, subequal to, as long as, Or - scarcely longer than,

2, with joint 3 usually not so conspicuously humped, sometimes more spindle-

shaped, or subparallel-sided, or even oval, ending apically in a relatively less

developed upper process which usually forms a more evident bifid process with the

lower one which itself is only a little shorter; head in front distinctly less prominent,

not conically produced; head below, or at least greater part of it below, and pleurae,

either entirely black or not with yellowish on all the above-mentioned sites; hairs

on head, body, and legs distinctly shorter, relatively less dense, less woolly or

shaggy .. =a ae ie 2 seh Aye a de of 22 9 ee

Head in front broader, less conically prominent; antennal joint 3 more con-

spicuously humped, with conspicuous hairs above, ending apically in a longer,

stouter, finger-like process; frons entirely black; head below without a central dark

fascia; scutellum more broadly rounded; hairs and scaling on body and legs dis-

tinctly much denser and longer, those above brassy or golden yellow; slightly larger

and more bulky forms, usually more than 4,5 mm long Ae “4 “aS

Head (cf. Fig. 1D) in front narrower, distinctly more conically prominent or pro-

duced; antennal joint 3 less conspicuously humped, without any conspicuous hairs

above, ending in a shorter, less stout, upper process; frons with a yellow stripe on

each side along eye-margins; head below with a central dark fascia; scutelum

distinctly more conically pointed or narrowed apically; hairs and scaling on body

and legs less dense, shorter, and those above distinctly silvery white; slightly less

bulky forms, only about 4,4 mm long, with a wing-length of about 4,2 mm

2 conocephala sp. nov.

Upper part of genae smooth, without hairs, or with a few pale ones; wings distinctly

less conspicuously milky whitish, with darker, darker brownish, or blackish brown

veins; knobs of halteres eae with a variable dark spot above; knees more distinctly

yellowish ae 2 lanigera Bezzi

Upper part of genae ah a | dense. tuft of black ‘hairs: wings distinctly and more

conspicuously milky whitish, the veins distinctly paler, more yellowish brown or

yellowish; knobs of halteres entirely ivory whitish or yellowish above; knees not,

or scarcely perceptibly, yellowish .. 2 .. @ lanigera var. melampogon 0.

Genae and head below entirely black; pleural parts entirely black; antennal joint

3 more spindle-shaped or oval; wings with the veins very pale, pallid or yellowish,

the middle cross vein usually much beyond middle of discoidal cell; apical stalk of

anal cell longer, at least as long as, or even longer than, apical cross vein of discoidal

cell; knobs of halteres entirely very pale or pallid above; basal joint of hind tarsi

usually about, or scarcely more than, half length of hind tibiae .. bi. ee

More than lower half of genae and continuous as a narrow streak round margin

of eyes to about halfway up behind eyes pallid; small, middle, boss-like prominence

in middle of pleurae between sterno- and hypopleurae and pteropleuron, and some-

times even to a certain extent on adjacent parts of sterno- and metapleuron, also

pallid; antennal joint 3 tending to be more subparallel-sided, less broadened at

middle; wings with the veins distinctly darker, more brownish to blackish brown,

the middle cross vein only a little beyond middle of discoidal cell; apical stalk of

anal cell shorter, markedly short, much shorter than apical cross vein of discoidal

cell; knobs of halteres slightly darkened above or with a variable dark infusion;

basal joint of hind tarsi distinctly much more than half length of hind tibiae

2 namaquensis sp. NOV.

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE 279

17 (a) Base of second submarginal cell more normal or at least not markedly acute; antennal

joint 3 relatively longer, more spindle-shaped, its upper apical process distinctly

stouter and longer than lower one, the apex less bifid in appearance; greater part

of palps luteous or yellowish; legs on the whole paler, with more yellowish or

yellowish brown, the apical parts of femora and the tibiae, and greater part of tarsi

pale yellowish or luteous; hairs relatively less dense, but slightly longer

2 nigribarba Hesse

(6) Base of second submarginal cell distinctly much more markedly acute; antennal

joint 3 (cf. Fig. 1E) relatively shorter, more oval, its upper apical process not much,

or scarcely, longer than lower one, the apex thus more symmetrically bifid; palps

dark; legs, including tibiae, distinctly much darker, mainly blackish brown or black,

only the knee-articulations being reddish brownish, and bases of tarsi tending to be

dark sienna or reddish brownish; hairs relatively shorter, but distinctly much

Genser... ne Ae cit si oe Sy, ie 2 ovalicornis sp. nov.

Subfamily Usiinae Becker

This subfamily, originally based on the typical Palaearctic genus Usia

Latreille, was supposed by me (Hesse 1938: 712) not to be represented in

southern Africa after the transference of the genera Corsomyza Wied. and

Hyperusia Bezzi, which Bezzi (1924: 103-109) placed in the Usiinae, to the

Bombyliinae (Hesse 1938: 712-713).

Recently Hull (1973:212-211) however emended and redefined this sub-

family, restricting it to four genera of which two, Usia Latreille and Dagestania

Paramonov, are Palaearctic and Oriental in their distribution and two, Apolysis

Loew and Oligodranes Loew, occur not only in the Palaearctic and Nearctic

Regions but also in the Ethiopian Region including the South African subregion.

As a result of Hull’s taxonomic assignment of genera in this subfamily

the latter two genera which I placed in the subfamily Phthiriinae in my revision

(Hesse 1938: 822) are now to be considered as important South African repre-

sentatives of the emended subfamily Usiinae.

Genus Apolysis Loew

Apolysis: Hesse, 1938: 848. Hull, 1973: 217.

The discovery of still other South African species of this genus does not |

appear to necessitate much change in the diagnostic characters recorded by

Loew (1860) and Engel (1933: 127) and supplemented by me in 1938, and more

fully detailed by Hull in 1973. Worthy of record however are that in the case

of the South African forms the wings appear to be more frequently distinctly

tinted milky whitish, that the abdomen of both sexes in some species is often

entirely or predominantly pallid or yellowish, that the legs of certain species

are relatively short and stoutish, and that the hind femora of still others are

slightly curved.

Descriptions of new species and comments on already described ones

are as follows:

280 ANNALS OF THE SOUTH AFRICAN MUSEUM

Apolysis monticola sp. nov.

Fig. 2D

In the absence of the 3g, these 99 are here described as a new species even

though they appear to be very near the § xanthogaster Hesse (1938: 853, fig.

259). They are characterized as follows:

Body and legs mainly black; integument dull, with very faint greyish white

bloom on head, thorax above, and scutellum; hind margins of tergites narrowly

pallid or yellowish, those of sternites more broadly yellowish, sometimes very

much so or even with greater part of venter, apart from the broad pallid hind

margins, reddish yellow, and with at least the sides (articulating part between

tergites and sternites) conspicuously yellowish; legs sometimes, or at least

the tibiae, more castaneous brown.

Vestiture with the hairs, even on head and thorax, very sparse, entirely

pale or whitish, those on abdomen short and a little longer posteriorly; those

on legs short, not conspicuous.

Head with antennal joint | distinctly longer than 2, about 14-2 times as

long; joint 3 distinctly slightly produced and rounded apically beyond the

spine which is dorsal and subapical in position; proboscis stoutish, about

0,6-1 mm long; palps longish, distinctly projecting, at least as long as antennal

joints 2 and 3 combined, and usually about as long as antennae.

Wings (cf. Fig. 2D) greyish hyaline, with a very faint greyish white sub-

opacity in certain lights; veins brown to dark brown; first basal cell a little

longer than second, and part of third vein between it and apical cell sub-equal

to, as long as, or even slightly longer than, cell itself; knobs of halteres entirely

whitish.

From 6 99, including the holotype, in the South African Museum.

Length of body: about 2,2-3,12 mm

Length of wing: about 2,3-3,4 mm

Distribution

Natal: Cathedral Peak in Natal Drakensberg, alt. 1 950 m (B. Stuckenberg,

March 1955).

The darker legs, the relatively longer wings, slightly longer first basal cell,

relatively longer second submarginal cell, much darker veins, and shorter

proboscis distinguish this species from the ¢ xanthogaster which also comes

from the Drakensberg on the Orange Free State side.

Apolysis thamnophila sp. nov.

A species very near the preceding species and also cingulata Hesse (1938:

853).

From monticola it differs in having the integument of body above distinctly

more shining, distinctly broader yellowish hind margins to the tergites, dark

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE 281

hairs on thorax above posteriorly and on scutellum, shorter and sparser hairs

on abdomen, a first basal cell which is as long as second, and very much shorter

than distance between first basal cell and base of second submarginal cell.

From @ cingulata Hesse it differs in being relatively smaller, 2,2 mm as

against 3,5 mm and with a wing-length of only about 2,4 mm as against 4 mm,

in having a darker buccal rim, narrower ivory yellowish hind margins to

tergites, entirely dark legs, including knees, more whitish hairs on occiput,

darker hairs on thorax posteriorly and on scutellum, entirely yellowish white

halteral knobs, shorter proboscis, only about 1 mm long as against 2 mm in

cingulata.

From a single 2 specimen in the South African Museum.

Distribution

Little Karoo: Rust en Vrede near Oudtshoorn (Museum expedition,

October 1951).

Apolysis stuckenbergi sp. nov.

Fig. 2A

This species which belongs to the humilis section is named after Dr B.

Stuckenberg of the Natal Museum who is a great South African dipterist and

who has collected many interesting species of Bombyliidae in Natal. The species

is characterized as follows:

Body black; hind margins of tergites and sternites narrowly and obscurely

pallid or whitish in 9; legs entirely very dark blackish brown to black; integu-

ment of body dull, with dull greyish white bloom, absent on thorax above in

6, but in 2 separated by two submedial dark lines and a broken lateral one.

Vestiture with the hairs on body and legs sparse, longer in ¢ than in 9,

entirely whitish.

Head with the eyes in ¢ in contact in front of ocellar tubercle, separated

in 9, on vertex nearly, or quite, 3 times width of tubercle; transverse frontal

depression in 9 at about middle and the middle longitudinal depression from the

ocellar tubercle ending in it; antennal joint 1 distinctly longer than 2, at least

14 times as long as 2; joint 3 (cf. Fig. 2A) elongate, with a rather distinct

prominence in front of apical spine, the latter dorsal and subapical in position, _

with the apex of joint beyond spine slightly produced and rounded; proboscis

(cf. Fig. 2A) about I-1,2 mm long, slightly stouter in 2; palps long, projecting

conspicuously, nearly or quite as long as antennae.

Wings greyish hyaline; veins dark blackish brown; first basal cell a little,

but distinctly, longer than second, shorter than distance between it and second

submarginal cell; second posterior cell widely divergent apically; knobs of

halteres darkened or black above in 3, usually entirely whitish in 2 or sometimes

with a small darkish spot above.

Legs slender; first basal joint of hind tarsi a little less than half length of

hind tibiae.

282 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 2. Left side views of heads and antennae, and right wings of some new species of Apolysis.

(All to the same scale.)

A. 9° stuckenbergi; B. capicola; CC. 9 brachycera; D. 2 monticola; EE. 2 seminitens;

F. 2 semiflava; G. ¢ hirtella.

From 5 39 and 5 99 (¢ holotype in the Natal Museum, 9 allotype in

the South African Museum).

Length of body: about 1,8-2,2 mm

Length of wing: about 2,12—2,6 mm

Distribution

Natal: Karkloof (B. Stuckenberg, | April 1956).

From humilis Loew it differs mainly in being smaller, with entirely whitish

hair on occiput, thorax above, and scutellum, and slightly less cinereous-

tinted wings.

Apolysis oreophila sp. nov.

Some other 99 from the Drakensberg in Natal resemble 99 of the preceding

species stuckenbergi so closely that they may almost be considered as only

representing a variety of the latter. In the absence of the ¢ and the slight

differences present in these 99 they are here considered as a distinct species.

They differ from stuckenbergi in being distinctly larger, about 2,4-3 mm,

with a greater wing-length of about 3-3,2 mm (as against 2,12—2,6 mm in the

latter); in having the subopacity of the wings in certain lights more distinctly,

though faintly, whitish and not cinereous, and with the veins more yellowish

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE 283

brown than blackish brown; apical stalk of anal cell apparently shorter, sub-

equal to, or even shorter (not a little longer) than apical cross vein of second

basal cell; hairs on body and legs distinctly longer, relatively more dense, and

the fine hairs on hind tibiae longer, more conspicuous; and in having slightly

broader and more constantly pallid hind margins to the abdominal segments,

especially the sternites.

From 6 9° (holotype in the Natal Museum and paratypes in the Natal

and South African Museums).

Distribution

Natal: Cathedral Peak area in the Natal Drakensberg, alt. 1950 m

(B. Stuckenberg, 23 March 1955).

Apolysis capicola sp. nov.

Fig. 2B

This southern Cape species is referable to the xanthogaster and monticola

section, but may also be compared with the stuckenbergi and oreophila section.

It agrees and differs from these species mentioned in the following respects:

Body, including legs, mainly black; integument entirely dull, thus differing

from cingulata and thamnophila; hind margins of tergites scarcely, or only

very narrowly, pallid in J, those of sternites in g narrowly pallid, the tergites

in 2 sometimes as in 3, but sometimes, especially those posteriorly, narrowly

and distinctly pallid or yellowish, broader so than in g, and hind margins of

sternites in some 99 relatively broadly yellowish, more like those of some 99

of monticola and thus distinctly broader so than in stuckenbergi and oreophila,

Vestiture on body distinctly not very dense, entirely sericeous whitish.

even on thorax above.

Head with the eyes in ¢ in contact above for about 2,42,8 times length of

ocellar tubercle; interocular space on vertex in 2, at narrowest part, distinctly

slightly broader than length of antennal joint 3 (sub-equal to length of joint

3 in 9 monticola); antennae (cf. Fig. 2B) with joint 1 short, subequal in length

to 2 or scarcely longer than 2 (distinctly or much longer than 2 in monticola,

stuckenbergi and oreophila); joint 3 with the dorsal prominence in front of

subapical excavation not, or distinctly less angularly, prominent than in —

stuckenbergi and oreophila; palps relatively stoutish, subequal in length to,

or scarcely longer than, antennae (more slender and longer in monticola and

the latter two species); proboscis (cf. Fig. 2B) stouter and appearing shorter

than in stuckenbergi and oreophila.

Wings differing from those of all the species mentioned in being distinctly,

though faintly, tinted subopaquely milky whitish; veins yellowish brown to

brown; first basal cell scarcely longer than second; second posterior cell slightly

divergent apically; knobs of halteres darkened or black above in d, entirely

whitish in 9, but in some 99 also slightly darkened above.

284 ANNALS OF THE SOUTH AFRICAN MUSEUM

Legs with the basal joint of hind tarsi only a very little less than half length

of hind tibia.

From 6 g¢ and 25 99, including the 3 holotype and 9 allotype, in the

South African Museum.

Length of body: about 1,84—-2,84 mm

Length of wing: about 1,8-3,04 mm

Distribution

South-western Cape: Pearly Beach in Bredasdorp District (South African

Museum expedition, December 1958) (holo- and allotypes); Wit River Valley,

Bainskloof near Wellington (November 1937); Leipoldtville (South African

Museum expedition, November 1956).

Apolysis fumalis Hesse

Apolysis fumalis Hesse, 1938: 855.

Since my description of the 2 of this species in 1938 the ¢ has also been

taken and the following supplementary notes on the species have to be added:

Body of 3 entirely black; integument of thorax above in ¢ without the

brownish grey bloom of 9; legs dark chestnut brown.

Vestiture with the hairs on occiput in both sexes, those on genae and head

below in 3, and those on rest of body in 3, including those on coxae and greater

part of legs dark blackish brown to black, those on thorax above, scutellum

and abdomen above in 2 sometimes also more brownish or brownish yellow

than yellow.

Head in 3 with the eyes, in front of ocellar tubercle, in contact for a

distance nearly 3 times length of the tubercle; antennal joint 1 in both sexes

quite twice as long as joint 2; proboscis about 1,4-1,8 (or 2) mm long; palps

long, conspicuous, and in ¢ and some 99, as long as antennae.

Wings distinctly dusky or smoky brownish, darker in ¢ than in Q; first

basal cell longer than second, longer than part of third vein between it and

second submarginal cell; second posterior cell widely divergent apically; knobs

of halteres in $ entirely dark castaneous brownish.

Legs with the basal joint of hind tarsi much more than half length of hind

tibia.

The original @ holotype is in the British Museum (Natural History),

the 3 allotype is in the South African Museum.

Length of body: about 3-3,5 mm

Length of wing: about 3,8-4,5 mm

Distribution

Natal: Karkloof (B. Stuckenberg, 1 April 1956) (¢ allotype and the

other 99).

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE 285

Apolysis minuscula sp. nov.

A minute aberrant species which is characterized as follows:

Body mainly black, the integument smooth, polished and shining; hind

margins of tergites (broadened on extreme sides), the slightly broader hind

margins of sternites, and the broad articulating part on each side below between

tergites and sternites in 9 ivory yellow or yellow: hind margins of tergites in

dg only very narrowly or obscurely yellowish, evident only on sides posteriorly;

antennae, proboscis and legs yellowish brown to dark castaneous brownish,

though antennal joints 1 and 2 sometimes paler.

Vestiture with the hairs on body and legs, and even head, fine, very short

and sparse, inconspicuous, almost imperceptible or wanting, even in J, and,

where detectable, apparently dark.

Head with the eyes in 3 broadly separated above, a little more than half

width of the interocular space in 9; frons medially foveately depressed in

basal half in front of anterior ocellus, somewhat tumidly convex and rounded

anteriorly, more evident in 9, the head thus more rounded anteriorly above;

antennal joint | a little longer than 2; joint 3 more elongate ovate in J, usually

more ovate in 9, at least half as broad as long; proboscis, relative to body,

markedly long, about 0,68-1,5 mm; palps shorter than antennae, not very

conspicuously visible and not projecting much.

Wings vitreous hyaline, with a scarcely detectable milky whitish tint or

subopacity in certain lights; veins or parts of veins in anterior half of wings

yellowish, their distal parts, or veins in posterior half, paler, more whitish;

first basal cell as long as second, much shorter than part of third vein between

it and second submarginal cell; second posterior cell widely divergent apically;

apical stalk of anal cell markedly long, relatively longer than in in other known

South African species of this genus, at least twice as long as base of third

posterior cell.

Legs with the basal joint of hind tarsi much shorter than half length of

hind tibia, sometimes even only a third of its length.

1 g and 6 99, including the 9 holotype and ¢ allotype, in the South African

Museum.

Length of body: about 1,2-1,7 mm

Length of wing: about 1,6-1,9 mm

Distribution

Bushmanland: Between Springbok and Pella (Museum staff, October

1939) (¢ holotype and 3 2 paratypes); Aggenys (Museum staff, October 1939)

($ allotype and a 2 paratype). North-western Cape: Kenhardt area (Museum

staff, October 1939) (1 @ paratype).

Frequents flowers of Mahernia grandiflora and M.nana, and is easily

recognized by its small size, cyrtosiine appearance, smooth and shining

integument, and almost entire absence of vestiture.

286 ANNALS OF THE SOUTH AFRICAN MUSEUM

Apolysis lindneri Hesse

Apolysis lindneri Hesse, 1962: 1, fig.

This is a very small species which Dr E. Lindner of the ‘Staatliches Museum

fiir Naturkunde in Stuttgart’ caught on the yellow flowers of the composite

Chrysanthemoides monilifera when he visited Cape Town in 1958 and which

I described in 1962.

It differs from all the other known South African species of Apolysis

except the preceding minuscula, in its small size. It is characterized as follows:

Body, including antennae and proboscis, entirely dark, dark blackish

brown; eyes reddish brown; hind margins of sternites slightly narrowly pallid

or pale.

Vestiture in form of fine, not very dense, greyish bloom on head, thorax,

abdomen, and pleurae, and shortish, somewhat sparse hairs, slightly longer

on sides of thorax anteriorly, on scutellum posteriorly, and on abdomen; these

hairs gleaming pale on sides of head behind eyes, darker on occiput, pale on

sides of thorax and on scutellum, darker on disc of thorax, more pale on

abdomen, pale, shortish and sparse on legs, slightly longer and denser along

outer hinder surfaces of hind tibiae.

Wings rather narrowish, slightly longer than body, greyish hyaline, but

distinctly faintly milky whitish in certain lights; veins brownish, even fine hind

margin and microtrichial fringe dark; first basal cell very slightly longer than

second, distinctly shorter than part of vein between it and apical cell; latter

cell slightly shorter than this same part; second posterior cell divergent apically ;

apical stalk of anal cell rather long, nearly twice length of base of third posterior

cell; knobs of halteres whitish.

Head with the frons parallel-sided, slightly broader than width of eye

(front view), slightly transversely depressed across middle, the apical part only

slightly convex; antennal joint 1 subequal to 2, the two together slightly

shorter than joint 3; the latter about 2,5 times as long as broad; proboscis

rather stoutish, only about 0,68 mm long; palps short, not perceptibly

projecting.

Legs rather slender and long, the tarsi elongate, with the basal segment of

hind ones quite, or nearly, half length of hind tibiae.

From the single 2 specimen in the Museum in Stuttgart.

Length of body: about 1,2 mm (excluding proboscis)

Length of wing: about 1,6 mm

Distribution

Western Cape Province: Cape Town (E. Lindner, 31 October 1958).

From minuscula, the other minute species from north-western Cape, it

may at once be distinguished by the duller head and body, covered with fine

greyish bloom, the longer and denser hairs on thorax and abdomen and legs,

absence of distinct and conspicuous yellowish hind margins to tergites and

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE 287

sternites, a first basal cell in wings which is slightly longer than the second,

slightly longer, more slender legs, with relatively longer basal joint tarsi, longer

basal joint of hind tarsi, and slightly shorter and stouter proboscis.

Apolysis thornei Hesse

Apolysis thornei Hesse, 1938: 857, fig. 261.

Of this species the South African Museum has since obtained a very long

series of both sexes from flowers of Mahernia brandiflora and M. nana in Bush-

manland, north-western Cape and the Koup Karoo.

From this series it is quite evident that the species is variable and that

certain statements made by me in 1938 have to be modified somewhat.

In the first place specimens (more typical form) from Bushmanland,

north-western Cape, and Nieuveld Karoo have the hind margins of the tergites

in the 99 only narrowly yellowish, more broadly yellowish only at apex of

abdomen; the wings slightly more apparently milky whitish; costal cell relatively

broad at middle; and knobs of halteres in ¢¢ darkened above.

Specimens from the Koup Karoo have the hind margins of the tergites

distinctiy or very much broader yellowish.

Some 99 from Bushmanland are entirely black, without any yellowish

or pallid hind margins, and with the integument of the entire body above

shining, and with the wings distinctly less milky whitish, more vitreous hyaline.

Some $4 from Namaqualand have not only the wings less milky whitish,

but the knobs of halteres entirely pallid.

Some ¢¢ from the Moordenaars Karoo near Laingsburg have not only

pallid halteral knobs, but much yellowish on abdomen and even more yellowish

legs.

Apolysis semiflava sp. nov.

Fig. 2F

This species which belongs to the thornei section and which, like thornei

and some other species, frequents the flowers of a species of Mahernia, is easily

recognized by the following characters:

Body with the head, thorax (except slightly yellowish humeral angle and

postalar margin in 9), and scutellum black; abdomen in @ entirely very pale ~

yellowish, the hind margins of segments sometimes even more whitish; abdomen

in $ with the hind margins of tergites narrowly pallid or yellowish to a variable

extent, broader on sides and those of sternites even broader so to a variable

extent, the entire venter sometimes tending to be yellowish and large scoop-like

last sternite (or tergite) entirely yellowish; integument of thorax and scutellum

above in $ more or less shining, duller in 2 and covered with greyish white

bloom; frons and upper parts of genae in ¢ with silvery tomentum; abdomen

in $ also markedly and densely covered with greyish white bloom; legs with the

femora yellowish brown, castaneous to dark brown, or almost black, the tibiae

288 ANNALS OF THE SOUTH AFRICAN MUSEUM

and tarsi, and usually the knees, in 3 paler or even more yellowish to a variable

extent, in 9 the tibiae and at least basal halves of tarsi, and also apices of femora

more consistently yellow.

Vestiture with the hairs on body and legs whitish, not very dense, but

distinctly much denser and longer in 3, those on abdomen in 9 long only

posteriorly.

Head with the eyes in 3 in actual contact above for some distance, about

4-22 times as long as ocellar tubercle; frons in 2 with the transverse depression

relatively shallow, less evident than medial longitudinal one in front of tubercle;

antennal joint | very short, as long as, or scarcely, or only a very little longer

than 2; joint 3 elongate-oval, about twice as long as broad, broadest slightly

beyond middle, its apical spine nearly terminal, the apex of joint scarcely pro-

duced beyond spine; palps short, inconspicuous, confined to buccal cavity;

proboscis about 1,12—1,48 mm long, usually a little stouter in 9.

Wings (cf. Fig. 2F) rather conspicuously tinted subopaquely milky whitish;

veins whitish, only those in costal part slightly more yellowish or pallid; first

basal cell distinctly much longer than second and much longer than part of

third vein between it and second submarginal cell; second posterior cell sub-

parallel-sided, scarcely, or not, diverging apically; apical stalk of anal cell

relatively long; knobs of halteres entirely whitish in both sexes.

Legs comparatively short, the basal joint of hind tarsi distinctly much

less than half length of hind tibia; claws feeble, short, mainly pale.

From 8 gd and 41 99, including the 2 holotype and J allotype, in the

South African Museum.

Length of body: about 2,08-3 mm

Length of wing: about 2,4-3 mm

Distribution

Little Karoo: Oudtshoorn District (Museum expedition, October 1952)

(2 holotype and ¢ allotype); Uniondale District (Museum expedition, October

1952). Karoo: between Vondeling and Willowmore (Museum expedition,

October 1952). Koup Karoo: Lammerfontein in the Moordenaars Karoo in

the Laingsburg Division (Museum expedition, October 1952).

Its entirely yellow abdomen in 9, broader yellowish hind margins of

abdominal segments in 3, the yellowish tibiae, milky whitish wings with

whitish veins, and shorter proboscis distinguish it at once from thornei which

species it appears to replace in the Little Karoo.

Apolysis lactearia sp. nov.

This species, represented by only the female sex among the new accessions,

can only be confused with 9° of thornei. The differences are however of such

a nature that they cannot be considered as merely varietal. From both the

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE 289

typical and varietal forms of the 2 thornei the 22 of this new species differ as

follows:

Body, including legs, almost entirely black; narrowish hind margins of

tergites and only slightly broader ones of sternites pallid; narrow hind border

of metapleurae also yellowish; articulating part below between tergites and

sternites broadly yellowish; knees not tending to be slightly yellowish as in

some 99 of thornei; integument of body, including head, thorax above and

scutellum, distinctly duller, mainly covered with greyish white bloom.

Vestiture with the hairs on body and legs distinctly very much denser and

longer, more conspicuously so on abdomen, but also entirely whitish as in

thornei.

Head with the frontal depression apparently less deep, especially that

medial part of it in front of ocellar tubercle; antennal joint 3 relatively shorter,

usually less than 24 times as long as broad, its apex beyond spine distinctly

very much less produced than in thornei, the spine more terminal and itself

slightly stouter; proboscis also slender, relatively long, 2,4-2,8 mm long; palps

as in thornei section short, inconspicuous, not projecting much.

Wings distinctly more conspicuously milky whitish; veins also pale

yellowish; first basal cell also much longer than second; costal cell apparently

slightly narrower, the costal vein not so markedly curved outwards as in some

2° of thornei; second posterior cell subparallel-sided and even sightly narrowed

apically as in the thornei section; knobs of halteres entirely whitish.

From 3 99, including the holotype, in the South African Museum.

Length of body: about 2,6-3 mm

Length of wing: about 2,8-3 mm

Distribution

North-western Karoo: Augusfontein near Calvinia (Museum expedition,

September 1947).

Apolysis namaensis sp. nov.

A 2 specimen from Namaqualand also belongs to this section and is very

near Jactearia from which it differs in the following respects:

Frons scarcely, or less, impressed in front of ocellar tubercle, less shining

along the middle, being mainly dull; proboscis relatively shorter, only about

2 mm long and distinctly stouter; antennal joint 3 with the humped part nearer,

or just a little beyond, middle (in /actearia nearer apex), the spine however

very similar and also almost terminal in position; legs distinctly stouter and

shorter, the femora especially are stouter, and legs with relatively longer and

much denser hairs; and wings milky whitish as in /actearia, but with the parts

of costal and first veins between apex of false vein much darker, dark brown

or blackish brown, not yellowish.

Length of body: about 3 mm

Length of wing: about 3 mm

290 ANNALS OF THE SOUTH AFRICAN MUSEUM

Distribution

Namaqualand: Wallekraal on road to Hondeklipbaai (Museum expedition,

October 1950).

Apolysis brachycera sp. nov.

Fig. 2C

Still another 2 specimen from the western coastal region is referable to

the thornei and Jactearia section, differing from both /Jactearia and namaensis

in the following respects:

Wings, though also milky whitish, with white and not yellowish veins;

antennal joint 3 (cf. Fig. 2C) distinctly very much shorter or rather more ovate,

scarcely more than 14 times as long as broad and its apical spine (even more

terminal in position than in Jactearia and namaensis) distinctly longer and more

slender; legs distinctly much stouter and shorter, the femora being even stouter

than in namaensis and basal joint of hind tarsi relatively shorter, distinctly

much less than half length of hind tibia whereas in Jactearia and namaensis

it is about, or very little less than, half length of tibia; hairs on body, especially

on abdomen, relatively shorter.

Length of body: about 3 mm

Length of wing: about 3 mm

Length of proboscis: about 1,88 mm

Distribution

West coastal region between Leipoldtville and Elands Bay (Museum

expedition, October 1947).

Apolysis seminitens sp. nov.

Fig. 2E

This striking new species, which was also caught in the flowers of a species

of Mahernia and which resembles the species semiflava superficially, however

belongs to the maherniaphila section in which the eyes in the §¢ are broadly

separated, the hairs on the body and legs are shorter and denser, and the legs are

on the whole shorter and stouter. It is characterized as follows:

Body with the head, thorax and scutellum black, the thorax and scutellum

above brilliantly shining; humeral angles and postalar calli reddish brown,

especially in 9; antennae and proboscis also sometimes reddish brown to

blackish brown; abdomen and also hind margin of metapleural part in both

sexes entirely very pale yellowish or pallid, the hind margins of both tergites

and sternites sometimes even more broadly whitish; legs with the coxae and

anterior and middle femora to beyond middle and to a variable and much

lesser extent hind femora, or only outer or outer lower part of hind ones

darkened yellowish brownish to dark brown or blackish brown, with the apical

parts of the femora or sometimes most of the hind ones, the tibiae, and at least

basal halves of tarsi very pale yellowish.

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE 291

Vestiture with the fine tomentum or bloom on anterior part of frons

on each side of antennae and on upper half of genae silvery greyish; hair

on body and legs entirely whitish, relatively poorly developed, very short

and sparse or almost absent on thorax above, that on head below, humeral

angles, and across hind margin of scutellum longer, more conspicuous; that

on abdomen however distinctly denser than on rest of body, much denser in

6 than in 9; hairs on legs much sparser and shorter than in most other species,

being very short and scarcely detectable on tibiae.

Head with the eyes in g broadly separated above, a little narrower than

in 9, about as wide as broad ocellar tubercle, in 2 a little wider than tubercle;

frons more deeply transversely depressed across middle in 2 than in 3; antennal

joint 1 markedly short, transverse or bead-like, as long as, or scarcely as long

as, the equally short and transverse bead-like second joint; joint 3 (cf. Fig. 2E)

slightly elongate-oval to ovate, about 1? to twice as long as broad, broadest

slightly beyond middle, its spine subterminal, quite half as long as broadest

part of joint; proboscis about 1-1,4 mm long, slightly stouter in 2; palps

short, not conspicuous.

Wings conspicuously subopaquely milky whitish; veins pallid or whitish;

first basal cell very much, or markedly, longer than second, and considerably

more than twice as long as part of third vein between it and second submarginal

cell; second posterior cell subparallel-sided, not diverging apically; knobs of

halteres entirely whitish in both sexes.

Legs comparatively short and stoutish, the femora rather stout and hind

ones markedly curved outwards; basal joint of hind tarsi comparatively short,

very much less than half, usually nearly or about a third, length of hind

tibiae; claws markedly long and slender, more than half length of last tarsal

joint.

From 10 3¢ and 10 99, including the ¢ holotype and @ allotype, in the

South African Museum.

Length of body: about 1,48-3,2 mm

Length of wing: about 1,88-3,08 mm

Distribution

Southern Namaqualand: Knersvlakte (Museum expedition, October 1950) _

(types). West coastal region: Papendorp at mouth of Olifants River (Museum

expedition, October 1950).

From the similarly-coloured semiflava it may at once be distinguished by

the separated eyes in the 3, the more shining and polished thorax in both sexes,

the relatively longer first basal cell, almost hairless tibiae, and the more strongly

developed claws.

From maherniaphila Hesse (1938: 859) it may at once be distinguished

by the entirely yellow abdomen, yellow tibiae, relatively shorter and stouter

legs, shorter proboscis, relatively longer first basal cell, and denser hairs on

abdomen.

292 ANNALS OF THE SOUTH AFRICAN MUSEUM

Apolysis hirtella sp. nov.

Fig. 2G

Still another bicolorous new species, with the eyes separated in the 3 and

belonging to the maherniaphila section, is to be added here. It is characterized

as follows:

Body with the head, thorax and scutellum black, dull, but with greyish

white bloom sometimes evident on sides of antennal insertions and body

above; ocelli reddish or reddish brown; humeral angles and postalar calli

also reddish brownish or piceous; abdomen in ¢ variable in colour, either

entirely or predominantly yellowish or with tergites | and 2 mainly dark or

with only hind margins of segments broadly yellowish or yellowish red, and

sometimes with the scoop-like last sternite (or tergite) mainly dark; abdomen in

2 more constantly entirely yellowish or pallid, but sometimes with the bases

of tergites, especially in basal half, or tergites 1-4, darkened to a variable

extent as in 3; antennae, proboscis and legs dark reddish brown, blackish

brown to black, the femora more consistently dark, and tibiae usually more

reddish brown, with the knees however yellowish.

Vestiture with the hairs on body and legs conspicuous, comparatively

dense and long in comparison with many other species, denser and longer in

3, especially on abdomen where they are characteristically erect and recurved

forwards, entirely or predominantly silvery whitish, though those on abdomen

in 2 sometimes with a slight sericeous yellowish tint.

Head with the eyes in 3 broadly separated above, on vertex even slightly

wider than distance between outer margins of posterior ocelli, but slightly

narrower than in 9; transverse depression on frons slightly deeper and more

evident in 9; antennal joint 1 (cf. Fig. 2G) short, but distinctly longer than

the very short second joint, sometimes nearly or about twice as long; joint 3

slightly elongate, elongate—oval to oval, about 14 to 2 times as long as broad,

at broadest part usually a little beyond middle, its apex rotundately rounded,

projecting a little beyond dorsal spine which is more subapical and markedly

short and feeble; proboscis about 1,44—2,2 mm long; palps small, inconspicuous.

Wings, relative to body, long, distinctly tinted milky whitish; veins mainly

pallid or pale yellowish, more whitish in hinder and basal half, the costal vein

beyond apex of false vein to apex of wing and even round posteriorly to second

posterior cell and sometimes also apical parts of posterior veins in this region

darkened or markedly dark; costal cell markedly broad and outwardly curved

at level of cross vein; first basal cell longer than second and distinctly longer

than part of third vein between it and second submarginal cell; second posterior

cell subparallel-sided; knobs of halteres entirely pallid or whitish, but some-

times with an indication of a dark spot above in some ¢<.

Legs rather short and stoutish, the femora rather thick; basal joint

of hind tarsi short, only about a third length of hind tibiae; claws short but

strong.

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE 293

From 6 g¢ and 6 99, including the 3 holotype and 9 allotype, in the

South African Museum.

Length of body: about 1,8-2,72 mm

Length of wing: about 2,56-3,08 mm

Distribution

Koup Karoo (labelled Gouph): Laingsburg District (Museum staff,

February 1938) (types); Lammerfontein in the Moordenaars Karoo near

Laingsburg (Museum expedition, October 1952).

Easily recognized by its rather dense vestiture, erect and recurved hairs on

abdomen of 3, mainly yellowish abdomen, and stoutish legs, in which respects

it also differs from maherniaphila.

From semiflava and seminitens, which also have a mainly or entirely

yellowish abdomen, it differs in being distinctly more hairy, and from the former

species in the separated eyes in the 3, and from the latter in being more hairy,

in not having a smooth and shining front part of the body, entirely dark or

black legs, etc.

REVISED KEY TO THE KNOWN SOUTH AFRICAN SPECIES OF Apolysis

1 (a) First basal (cf. Fig. 2D) cell in wings as long as, or scarcely, or only a little longer

than second, and as long as, or subequal to, or sometimes even shorter than, part

of third vein between it and apical cell (if longer first basal cell only slightly longer

than second); second posterior cell widely divergent apically; wings rarely with a

very distinct milky whitish tint (if so first basal cell not long); palps usually elongate,

conspicuously projecting (if short other characters do not differ) 2

(b) First basal cell (cf. Fig. 2F) distinctly much longer than second and distinctly much

longer than part of third vein between it and apical cell; second posterior cell sub-

parallel-sided, sometimes even slightly narrowed apically; wings usually with a more

distinct, sometimes conspicuous, milky whitish tint; oe usually much shorter,

or very short and insignificant si 5 : a Sifie

2 (a) Palps longer or elongate, conspicuously yicible. Sr OSLIIE ae beyond buccal

cavity; antennal joint 3 distinctly longer, more elongate, much more than 2,5 times

as long as broad and usually equally long or broad in both sexes; wings more

elongate, relatively narrower, cinereous hyaline, cinereous, or even distinctly dusky

or infuscated, the veins distinctly darker; larger forms, usually much more than

2or2,5mmlong .. 3

(6) Palps much shorter or very ‘short, insignificant not easily visible, either hidden in

buccal cavity or not conspicuously projecting; antennal joint 3 distinctly relatively

shorter, more oval or ovate, or not much more than 2 or 2,5 times as long as broad

and either shorter or broader and more distinctly ovate in 92; wings relatively shorter

and broader, clear hyaline or with a faint or distinct milky whitish tint, the veins paler,

more yellowish; smaller or minute forms, usually less than 2 or 2,5 mm long.. 10

3 (a) Hairs on head, occiput and genae, on thorax and scutellum above either entirely

pale or whitish like rest of hairs on body and legs, or with fewer dark ones and then

not on all the parts mentioned; wings clearer or only faintly greyish hyaline and with

a more milky whitish subopacity in certain lights; apical part of antennal joint

3 beyond spine-like process slightly, but distinctly, more produced or apical margin

above, just beyond spine, distinctly more indented or incised . 4

(6) Hairs on head, occiput and sometimes genae and on thorax above ‘and scutellum

distinctly not white, darker or black, or with more numerous dark ones; wings

darker, distinctly more cinereous, dusky or infuscated, the subopacity more ‘greyish

294

(a)

(6)

(a)

(0)

(a)

(5)

(a)

(6)

ANNALS OF THE SOUTH AFRICAN MUSEUM

than whitish; apical part of antennal joint 3 beyond spine usually less produced,

more rotundately rounded, the i margin above, just beyond spine, usually

less incised .. * oe a 11

Abdomen with the hind margins ii Pipes aad stceaiies etaaden more con-

spicuously, yellowish or pallid, or the venter itself may be mainly yellowish; antennal

joint 3 with the prominence on dorsal margin, just before subapical excavation,

less sharply or angularly prominent, the joint thus less humped in appearance 5

Abdomen with the hind margins of the tergites and sternites only very narrowly or

obscurely pallid or yellowish, greater part of venter not mainly yellowish; antennal

joint 3 with the prominence, just before apical excavation, distinctly more sharply

or angularly prominent, giving joint a more humped appearance re Lee

Body above, especially abdomen, dull, not shining; hind margins of tergites less

broadly pallid or yellowish; sternites in some 29, and sometimes in some ¢¢ also,

more broadly yellowish or even entire venter yellowish; hairs on body more whitish

or entirely sericeous whitish; proboscis on the whole or ae shorter, stouter,

usually only about 0,6-1 mm long . 6

Body above, especially abdomen above, distinctly more , shining: hind margins of

tergites more broadly, more conspicuously, ivory yellowish or yellowish; greater

discal part of venter, even in 92, not conspicuously yellowish; hairs on occiput,

thorax above or only basally, and on scutellum, and sometimes on abdomen above

to a certain extent, distinctly more yellowish or even darker, not whitish; proboscis

relatively longer, usually about 1-2 mm .. ; as 8

Legs distinctly paler, more reddish brown, the tibiae and tarsi even 1 Baler: entire

venter, even in 4, distinctly yellowish; hairs on body distinctly denser, more shaggy;

wings relatively shorter, the veins paler, more yellowish brown; second submarginal

cell relatively short, shorter than part of third vein between it and first basal cell;

proboscis about 1,33 mmlong .. : 36 xanthogaster Hesse

Legs on the whole darker or entirely black: entire venter in known ¢¢ not yellowish ;

hairs on body distinctly less dense, sparser; wings more elongate, the veins darker

brown to dark brown; second submarginal cell normally long, subequal in length

to, or even slightly longer than, part of third vein between it and first basal cell;

proboscis relatively shorter, about 0,6-1 mm long wt ; iy 7

Wings more greyish hyaline; hind margins of sternites in 2 more bneaig yellowish,

even entire venter yellowish; antennal joint 1 distinctly longer than joint 2; palps

more slender, relatively longer, distinctly longer than antennal joint 3; interocular

space in 9, at narrowest part, on vertex relatively narrower, subequal to length of

antennal joint 3; knobs of halteres in 2 entirely whitish .. .. 2 monticola sp. nov.

Wings more distinctly tinted subopaquely milky whitish; hind margins of sternites

apparently less broadly yellowish, the entire venter not yellowish; antennal joint

1 markedly short, subequal in length to, or very little longer than, 2; palps shorter,

stouter, relatively shorter, subequal in length to, or scarcely longer than, antennal

joint 3; interocular space on vertex in @ relatively broader, distinctly broader than

length of antennal joint 3; knobs of halteres either entirely whitish or slightly

darkened above in 9, consistently darkened above in ¢ .... do 2 capicola sp. nov.

(more especially some 29)

Rim of buccal cavity ivory yellowish; hind margins of tergites more broadly ivory

yellowish; knees slightly yellowish and tibiae and tarsi tending to be less dark than

femora; hairs on occiput and thorax above more straw-coloured yellowish; knobs of

halteres slightly darkened above towards base; proboscis slightly longer, about

2 mm long; antennal joint 1 slightly longer, longer than 2; slightly larger form,

about 3,5 mm long, with a wing-length of about4mm.. .... Q cingulata Hesse

Rim of buccal cavity dark or dark reddish brown; hind margins of tergites only

very narrowly ivory yellowish; legs, including knees, entirely dark or black; hairs

on occiput and thorax anteriorly whitish, those on base of thorax above and on

scutellum dark or blackish; knobs of halteres entirely pale yellowish white above;

proboscis shorter, only about 1 mm long; antennal joint 1 shorter, scarcely longer

than 2; smaller form, about 2,2 mm long, with a wing-length of about 2,4 mm

Q thamnophila sp. nov.

(a)

(5)

(a)

(6)

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE 295

Antennal joint 1 much longer, at least twice length of 2; prominence in front of

subapical excavation on joint 3 above more angularly prominent; proboscis distinctly

more slender, thinner; palps more slender, longer than antennal joint 3; wings

more greyish hyaline, not or scarcely tinted faintly milky whitish, the veins darker

brownish to blackish brown; hind margins of tergites and sternites in both sexes

narrowly, or scarcely, pallid; knobs of halteres in 29 usually entirely white... 10

Antennal joint 1 markedly short, about as long as, or only a very little longer than,

2; prominence on dorsum of joint 3 less angularly prominent; proboscis distinctly

stouter, appearing shorter; palps relatively shorter and stouter, shorter, or at least

not much longer, than antennal joint 3; wings distinctly tinted more subopaquely

whitish, the veins more yellowish brown; hind margins of tergites and sternites in

some 2° relatively more broadly yellowish; knobs of halteres in some 29 darkened

above as in dd ie ate A a 32 capicola sp. nov.

Smaller species, about 1,8-2 mm one: with a wing-length of about 2,12-2,4 mm;

wings faintly more greyish or cinereous, more so in 4, the veins darker, dark brownish

to blackish brown; apical stalk of anal cell relatively longer, usually a little longer

than apical cross vein of second basal cell; narrow pallid hind margins of tergites

and sternites, even in 9, tending to be less constant, more obscure; hairs on body

and legs shorter, sparser, and fine ones on hind tibiae relatively shorter, less con-

spicuous . & 2 stuckenbergi sp. nov.

Slightly larger. species, about 2423) mm long, with a wing: -length of about 3-3,2

mm; wings less greyish, with a more distinct whitish subopacity, the veins slightly

paler, more yellowish brown; apical stalk of anal cell relatively shorter, subequal

to, or even shorter than, apical cross vein of second basal cell; narrow pallid hind

margins of tergites and sternites, in 9 at least, more constantly present, especially

the more broadish ones on sternites; hairs on body and legs longer, slightly darker,

and fine hairs on hind tibiae relatively longer, more conspicuous

2 oreophila sp. nov.

Wings clearer, less dusky or infuscated, only greyish hyaline or tinted slightly cine-

reous, the veins more reddish brown; part of third vein between first basal cell and

apical cell tending to be longer, subequal to, or only a little shorter than, latter

cell; knobs of halteres in ¢ darkened above only; hind margins of tergites and sternites

in both sexes less conspicuously, or broadly, yellowish, only narrowly or obscurely

pallid; hairs on body less extensively dark or black, those on head below, on abdomen,

and on legs, even in 3, paler or more whitish; hump before apical excavation on

antennal joint 3 less prominent Ses ; .. & 2 humilis Loew

Wings distinctly dusky, more infuscated, or smoky brownish, even more so in 3d,

the veins darker, very dark brown, blackish brown, or even black; part of third

vein between first basal cell and apical cell tending to be markedly shorter than

apical cell; knobs of halteres in ¢ usually darkened above and below; hind margins

of tergites and sternites, in 2 especially, more conspicuously yellowish and even

more broadly so on venter; hairs on body more extensively dark or black, those on

head below, on abdomen and legs in ¢ tending to be black, in 2 more yellowish,

yellowish brown to brown (if paler other characters do not differ); hump before

apical excavation on antennal joint 3 more prominent or even knob-like

3 2 fumalis Hesse

Slightly larger, more bulky species, about 1,5-2,5 mm long, with a wing-length of

about 1,5-2,5 mm; body duller, with more conspicuous, denser, greyish white

bloom, in @ especially, with two rows of velvety black quadrangular patches on

abdomen above (those on tergites 2 and 3 the largest), and with black patches on

thorax; proboscis much shorter, stouter, only about 0,4-0,6 mm long; wings more

conspicuously milky whitish, their first basal cell distinctly longer than part of

third vein between it and apical cell and latter also considerably longer than this

part; apical stalk of anal cell shorter, less than twice width of base of third posterior

cell; hairs on body and legs much denser, longer, markedly so in 9; eyes in gd in

actual contact for some distance in front of ocellar tubercle. . 5 9 brevirostris Hesse

Very much smaller, or minute, less bulky species, only about 1,2-1,7 mm long,

with a wing-length of about 1,5-1,9 mm; body either less dull and with much feebler

296

13 (a)

(b)

14 (a)

(5)

15 (a)

(6)

ANNALS OF THE SOUTH AFRICAN MUSEUM

greyish bloom, or smooth and shining, without any bloom, without any distinct,

conspicuous, black, velvety patches on abdomen; proboscis, relative to body, much

longer, more slender, about 0,68-1,5 mm; wings tinted more faintly milky whitish,

the first basal cell distinctly shorter than, or only subequal in length to, part of vein

between it and apical cell, and latter sub-equal in length to, or even shorter than,

same part; apical stalk of anal cell longer, quite or at least twice length of base of

third posterior cell; hairs on body and legs either much sparser and shorter, or

very sparse and short; eyes in known ¢¢ broadly aT ae only a little narrower

than in 99° ee S48 : ee _ 13

Head and body smooth, sity sess: waiehiant any seven tomentum; hind

margins of tergites and sternites, especially in 2, more broadly and more con-

spicuously yellowish; hairs on head, body and legs minute or absent in 9, very

sparse and short on thorax and abdomen in dg; legs shorter, the tarsi shorter and

basal joint of hind ones much shorter than half length of hind tibia; veins in hinder

part of wings paler yellowish or becoming paler distally; hind margin and micro-

trichial hairs in wings very pale or whitish; first basal cell as long as second; anterior

part of frons distinctly more convex or subtumid; proboscis in @ slightly less stout,

reaching the length of 1,9 mm ae ; 3 2 minuscula sp. nov.

Head and body more dulled toa variable extent by fine greyish tomentum; hind

margins of tergites and sternites, even in 9, only narrowly, scarcely, or obscurely,

pallid or yellowish; hairs on head and body, especially abdomen and legs, even in

9, distinctly denser and longer, those on hind tibiae distinctly longer, more evident;

legs relatively longer, the tarsi longer, and basal joint of hind ones quite or nearly

half length of hind tibia; veins in wings darker; hind margin of wings and micro-

trichial fringe dark; first basal cell slightly longer than second; frons anteriorly less

convex; proboscis, even in 2, rather stoutish and short, only about 0,68 mm long

2 lindneri Hesse

Eyes in $< in actual contact above for some distance, about 24 to nearly 3 times

length of ocellar tubercle, and with the upper facets distinctly much coarser than

lower ones; distance between posterior ocelli in 22 subequal to, as long as, or scarcely

perceptibly longer than, distance between them and inner margin of eyes; vestiture

on body and legs usually longer, even if sparse, and that on abdomen, especially in 29,

sometimes tending to be less dense; legs on the whole more slender, longer, the basal

joint of hind tarsi usually relatively longer, at least half length of hind tibiae .. 15

Eyes in 3g widely separated above by a broad space which is only a little narrower

than that of 2?, and with the upper facets not coarser than lower ones; distance

between posterior ocelli in 92 distinctly or perceptibly longer than distance between

them and inner margin of eyes; vestiture on body and legs on the whole shorter,

usually denser, and that on abdomen, especially in 29, usually markedly denser;

legs on the whole shorter, stouter, the basal joint of hind tarsi usually shorter, much

shorter than half length of hind tibiae .. 2A J. a

Legs either entirely dark or black or, if tibiae sae tarsi are isda they are not pallid

or pale yellowish; yellow on abdomen distinctly less developed, the hind margins

of tergites and sternites in known 3 either not yellowish or pallid or only narrowly,

obscurely and less conspicuously so, and in 99 abdomen is not entirely or predo-

minantly yellowish, at most with only broadish pallid or yellowish hind margins;

wings slightly less conspicuously milky whitish and, if so, veins tend to be more

yellowish than whitish or pallid and, if whitish, nn are dark; proboscis relatively

longer, usually more than 1,5 mm “2 a

Legs on the whole much paler, the femora more . yellowish brownish or brownish

and greater part of tibiae and tarsi, excluding only the brownish apical parts of

latter and sometimes slightly brownish apices of tibiae, entirely or predominantly

pale yellowish; yellow on abdomen more developed, the entire or greater part of

abdomen in @ very pale yellowish or pallid, and in ¢ with broader pallid or yellowish

hind margins, sometimes very broad on sides, and with last elongated sternite

entirely or predominantly yellowish or at least with much yellowish; wings more

conspicuously subopaquely milky whitish, the veins very pallid or whitish; pro-

boscis relatively short, only about 1,12-1,48 mm long .. 3 2 semiflava sp. Nov.

(a)

(6)

(a)

(5)

(a)

(5)

(a)

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE 297

Integument of body above, especially thorax and scutellum, and in 2 also entire

or greater part of frons, more shining, less dull, not densely covered with greyish

white bloom; antennal joint 3 more elongate, relatively longer, about 2,5-3 times

as long as broad, its apex beyond subapical spine distinctly, though slightly, more

produced, more rounded, the spine thus distinctly more subapical in position;

frontal depression in 2° relatively deeper and medial part in front of anterior ocellus

also distinctly deeper; wings either not distinctly milky whitish or tinted less so;

hairs on body and legs, especially on abdomen, and more so in 99, distinctly shorter

and sparser .. ea

Integument of body above and i in 9 also greater part of frons distinctly duller, very

densely covered with greyish white bloom; antennal joint 3 less elongate, shorter,

not more than, usually less than, 2 times as long as broad, its apex beyond spine

distinctly not produced, the spine appearing more terminal; frontal depression

in 2° less deep, especially the medial part in front of anterior ocellus; wings more

conspicuously subopaquely milky whitish; hairs on body and legs, especially on

abdomen, even in 29, distinctly longer, more shaggy and relatively denser .. 19

Wings with a distinct, though faint, milky whitish tint; costal cell relatively broad

at middle, its anterior margin more curved outwards near cross vein; knobs of

halteres in gd usually either entirely dark above or darkened above to a variable

extent; abdomen in 9° with the hind margins of tergites and sternites yellowish to

a variable extent, more so apically; anterior part or half of frons, genae, and abdomen

above in 2° distinctly duller tok ; 3 & thornei Hesse (and forms of it)

Wings more vitreous hyaline, without any perceptible milky whitish tint; costal

cell appearing narrower, its anterior margin not markedly curved outwards; knobs

of halteres, in known 34, entirely whitish or pallid; abdomen in known 22 entirely

black or with scarcely any indication of yellowish hind margins; entire frons and

body above in 99 shining .. a distinct 2 form of thornei Hesse and some 34 (18)

Base of wings up to cross vein in costal cell more subopaquely whitish

6 form of thornei Hesse

Base of wings up to cross vein distinctly more opaquely yellowish

3 form of thornei Hesse

Veins in wings distinctly more yellowish; antennal joint 3 distinctly more elongate,

at least twice as long as broad, its apical spine relatively stouter and shorter; legs

relatively longer, the basal joint of hind tarsi longer, only a little less than, or nearly,

half length of hind tibiae; hairs on abdomen relatively longer . . a Sei 20

Veins in wings white or whitish; antennal joint 3 distinctly shorter, more ovate,

scarcely more than 1,5 times as long as broad, its apical spine distinctly more slender

and longer; legs relatively stouter arid shorter, the basal joint of hind tarsi shorter,

aad much less than half ree of hind tibiae; hairs on abdomen relatively

shorter . ls a 2 brachycera sp. nov.

Frons more aicrinetly, and. more Gesaee Geprescd in front of ocellar tubercle,

also more shining along middle; proboscis slightly longer, more slender, about

2,48-2,9 mm long; humped part of antennal joint 3 nearer apex; legs distinctly

more slender, longer, the femora more slender, with relatively shorter hairs; part of

costal vein and first main vein beyond apex of false vein yellow or more yellowish, ~

the costal vein only slightly darkened apically .. ‘ 2 lactearia sp. nov.

Frons scarcely or less impressed in front of tubercle, less shining along middle,

mainly dull; proboscis distinctly shorter and stouter, only about 2 mm long; humped

part of antennal joint 3 nearer, or just a little beyond, middle; legs distinctly shorter

and stouter, the femora distinctly stouter, with relatively longer hairs; part of costal

vein and first vein, beyond apex of false vein, much darker, dark brown or blackish

brown .. ; 2 namaensis sp. Nov.

Abdomen not predeniiiantde: or Satis vellowielt or pallid in both sexes, only hind

margins of tergites and sternites being yellowish to a variable extent, much less so,

or only narrowly so, in 3; legs distinctly longer, more slender, the femora relatively

less stout, and basal joint of hind tarsi relatively longer, about, or a little less than,

or even a little more than, half length of hind tibiae; proboscis on the whole longer,

reaching up to 2 mm; hairs on abdomen, even in g,sparser 4 2 maherniaphila Hesse

298 ANNALS OF THE SOUTH AFRICAN MUSEUM

(b) Abdomen predominantly or entirely pallid or yellowish, or with broader yellowish

hind margins, in $4, and entirely so in 99; legs distinctly shorter, stouter, the

femora relatively thicker and stouter, and basal joint of hind tarsi shorter, usually

very much less than half length of hind tibiae; proboscis on the whole shorter, less

than 2 mm; hairs on abdomen, even in 99, denser, sometimes markedly dense and

conspicuous .. F ; aia i = ee

22) i(@) Integument of head abate: eer and : cellabas brilliantly annie! smooth and

polished in appearance; hairs on body and legs, especially on thorax and scutellum,

less developed, sparser, those on abdomen not so markedly recurved forwards;

abdomen in both sexes entirely pallid or very pale yellowish; legs with more yellowish,

sometimes mainly yellowish or with at least apical parts or halves of femora, entire

tibiae, and basal halves of tarsi pale yellowish, and the hind femora tending to be

more curved; antennal joint 1 very short, not or scarcely as long as second; antennal

joint 3 more narrowed basally, its spine much longer and more apically situated;

costal cell less markedly broad, its anterior vein not so convexly arched; first basal

cell relatively longer .. 3 2 seminitens sp. nov.

(6) Integument of head above, thorax, ‘and scutellum dull, densely covered with greyish

white bloom; hairs on body and legs, even on thorax above, distinctly markedly

denser, those on abdomen above, especially in ¢, markedly upright or recurved

forwards; abdomen in 3 sometimes with the bases of tergites darkened to a variable

extent and in 2 sometimes with only hind margins of sternites yellowish to a variable

extent; legs mainly or entirely dark, dark castaneous brown to black, only the knees

yellowish, and with the hind femora less markedly curved; antennal joint 1 a little

or distinctly longer than 2; antennal joint 3 on the whole broader, less narrowed

basally, its apical spine much feebler, more dorsal or subapical in position; costal

cell broader, more markedly dilated, its anterior vein convexly arched; first basal

cell relatively shorter oy a ue Re AF 3 9 hirtella sp. nov.

Genus Oligodranes Loew

Oligodranes: Hesse, 1938: 861. Hull, 1973: 219.

As is stated under the subfamily Usiinae in this paper the genus Oligodranes

is the other genus, formerly placed in the subfamily Phthiriinae, which Hull,

in his recent monograph of the genera of the Bombyliidae, transferred to the

subfamily Usiinae.

In view of the discovery of four additional South African species of this

genus, which I take to be the South African counterpart of the Palaearctic

form and of which I gave a supplementary description in 1938 based solely

on the South African representatives, the following supplementary notes on

the genus should now be added:

The South African forms seem to show distinct colour differentiation

in the two sexes. All the known 3<¢ are consistent in being predominantly

or entirely black; the yellow colouring on body and legs being much reduced

or almost absent. The 99 on the other hand always show more extensive yellow

colouring or a striking pattern of yellow or pallid markings on the head, thorax,

pleurae, and abdomen.

As far as cephalic characters are concerned it is to be noted that the eyes

in $3 are not always in contact above; these in some species are widely separated

as in the case of the 99. In this respect they agree with ¢¢ of some species in

the genus Apolysis. In the latter genus however the last tergite (or sternite)

in the $¢ is scoop-like and the genitalia are prominent, thus rendering

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE 299

differentiation between the sexes easy. In Oligodranes the genitalia of the $3

are very small, insignificant, usually hidden or not easily seen. Confusion of

the sexes is therefore not excluded if the eyes of the ¢ are broadly separated

as in the 9. Such a mistake was made by me in 1938 in the case of the supposed

2 of namaensis Hesse (see below).

As in the case of species of Apolysis most of the South African repre-

sentatives of Oligodranes frequent flowers and in the latter case mostly those of

species of Mesembryanthemum.

Oligodranes namaensis Hesse

Oligodranes namaensis Hesse, 1938: 864, fig. 263b.

Since my description of this species in 1938 the Museum has acquired

a long series of both sexes of this species. As they were caught together fre-

quenting the flowers of species of Mesembryanthemum and also show certain

characters in common there can be no doubt that the sexes belong to the same

species.

This discovery necessitates the correction of an error in identification

made by me in 1938. A comparison of the real 2 of namaensis with the supposed

2 allotype of namaensis has not only shown that the latter is not the Q2 of

namaensis, but that it is a g with separated eyes belonging to another species

which is described below as a new species. Only the characters given for the

dS specimen of namaensis are therefore valid in my description of 1938. A

supplementary redescription of this species, which also includes the real

and differently-coloured 9, is given here to render its recognition more easy:

Body and legs in ¢ mainly black, with the frons in front to a variable

extent, palps, humeral angles and anterior spiracular area to a variable extent,

an infusion just below base of wings, postalar calli to a variable extent, meta-

pleural part just below halteres, ligamentous connection between squama and

scutellum, very narrow hind margins of tergites and broader ones of sternites

pallid or yellowish; knees of § usually also yellowish.

Body in & also partly black, but with the following parts very pale yellow:

sides of frons in middle, anterior part of frons, entire genae, palps, greater part

of head below, occiput broadly behind eyes (except broadish, black, central,

postvertical stripe and middle part of occiput), triangular humeral spot, a

broad spot just below it, notopleural part, postalar calli broadly, broadish

base of thorax (or two spots), entire scutellum, propleural spot, anterior spira-

cular part, prosternal part, area just below wing-bases, a longitudinal fascia

along middle of pleurae, greater part of metapleural region, fairly broadish

hind margins of tergites, broader hind margins of sternites or sometimes almost

entire venter, entire coxae (excepting only a brownish anterior basal spot on

hind ones), and the femora (excepting only the brownish anterior upper sur-

faces, or spots, in apical halves, especially of hind ones).

Vestiture with the hairs sparse, longer in 3, shorter and denser on abdomen

300 ANNALS OF THE SOUTH AFRICAN MUSEUM

in 9, entirely pale or whitish, though with slight sericeous yellowish gleams on

abdomen in 9.

Head with the eyes above in ¢ in actual contact for a distance at least

twice length of ocellar tubercle; interocular space on vertex in 9 quite twice

width of ocellar tubercle; transverse depression across middle of frons in 2

rather deep and conspicuous; antennal joint | about 14-2 times length of

joint 2; joint 3 about 12-2 times combined length of joints 1 and 2; proboscis

1,12-1,4 (average 1,2) mm long; palps slender, subequal in length to, or about

as long as, antennal joint 3.

Wings hyaline, iridescent, with a very feeble milky whitish tint in certain

lights, more detectable at base; veins yellowish brown to brown, more

yellowish at base; middle cross vein at about between basal third and basal

fourth of discoidal cell; first basal cell usually a little shorter than distance of

third vein between it and base of second submarginal cell and latter cell, along

lower vein, also distinctly shorter than this part of third vein; knobs of halteres

entirely pallid or whitish in both sexes.

Hypopygium of ¢ as shown in figure 263b (Hesse 1938: 864).

From 12 ¢¢ (including original 3 holotype) and 34 99.

Length of body: about 1,88—2,28 mm

Length of wing: about 2,2-2,6 mm

Distribution

Namaqualand: Kamieskroon (Museum staff, September 1930) (¢ holo-

type). Bushmanland: Aggenys between Springbok and Pella (Museum staff,

October 1939) (34, 2 allotype and 9 paratypes). West coastal region: Citrusdal

District (Museum expedition, November 1948) (33).

Oligodranes triseriatellus sp. nov.

Fig. 3A, C

This slightly variable species, which was also caught on the flowers of a

species of Mesembryanthemum, is very near namaensis. From the latter it

however differs in the following respects:

Body in 3 with the hind margins of tergites and sternites distinctly much

more broadly and more conspicuously yellow, sometimes extensively so; body

in 2 with the distribution of black and yellow variable, differing from 2 of

namaensis in having the entire occiput behind ocellar tubercle and entire frons

(excepting sometimes for a much narrower median dark spot or line about

middle) yellow, sides and base of thorax above distinctly much more broadly

and extensively yellow, the part on each side above wing-base also continuously

yellow, isolating a rather conspicuous, round, black spot or peninsula on

each side, in having the pleurae more extensively yellow (sometimes with only

a large black spot in upper half of mesopleuron), with the yellow on abdomen

above much more extensive than in namaensis, the hind margins very broadly

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE 301

Fig. 3. Left side views of heads and left hind legs of some new species of Oligodranes.

(All to the same scale.)

A. ¢ triseriatellus; B. 3 anomalus; C. ¢ triseriatellus; D. 3 anomalus; E. 2 puberulus.

so, reducing or more often breaking up the black basal parts of tergites into

three longitudinal rows of black spots or patches which become smaller

posteriorly and which, in some specimens, are even absent in posterior half of

abdomen or very much reduced in size, almost the entire abdomen being

yellowish; legs in 2 often also with more yellowish on the tibiae or on middle

of tibeae, especially hind ones, and with the brownish infusions on upper or

anterior parts of femora in apical halves usually also more conspicuous.

Head (cf. Fig. 3A) with antennal joint 1 tending to be partly yellowish in

$ and entirely yellowish in 2; proboscis on the whole comparatively longer,

1,2-1,8 mm; palps also shorter, usually even less conspicuous, usually distinctly

shorter than antennal joint 3.

Wings with the veins on the whole paler, more yellowish; distance between

middle cross vein and base of second submarginal cell relatively shorter,

especially in 3, usually only a little, or scarcely, longer than lower vein of

cell itself whereas in namaensis this distance is more constantly markedly longer;

middle cross vein with a tendency to be farther away, at least at, or slightly

more than, basal third of discoidal cell.

302 ANNALS OF THE SOUTH AFRICAN MUSEUM

Legs with the left hind one of 3 shown in Figure 3C.

From 4 g¢ and 15 99, including the types, in the South African Museum.

Length of body: about 1,72-3,28 mm

Length of wing: about 2,12-3,68 mm

Distribution

Bushmanland: Pofadder (Museum staff, October 1939) (types and para-

types); Aggenys between Springbok and Pella (Museum staff, October 1939)

(paratypes). North-western Cape: Putsonderwater (Museum staff, October

1939) (paratypes). Great Karoo: Richmond District (Museum staff, November

1939) (Q paratype).

As is evident from the description this species appears to be variable not

only in size, but also in the extent of the yellowish or black colouring on the

head, abdomen and legs in 99. Some 99 have the three rows of black spots on

the abdomen less isolated or larger, tending to coalesce into black basal bands

across the tergites; in others the spots are small and the tibiae yellowish to

a variable extent, sometimes almost entirely yellowish.

The 2 paratype from Richmond has not only a tendency for the three

series of black basal spots on tergites 1-3 to coalesce but also to have a dark

medial postvertical stripe on the occiput as in 2 of namaensis.

Oligodranes puberulus sp. nov.

Fig. 3E

Another new species caught on flowers of a species of Mesembryanthemum

is characterized as follows:

Body and legs mainly black in both sexes, but in 3 with the frons in front

and upper half of genae, palps, humeral angle, margins of anterior spiracle,

notopleural ridge, area just below wing-bases, postalar calli, ligamentous

connection between squama and scutellum, metapleural part just below halteres,

and narrow hind margins of tergites and sternites pale yellowish or pallid, the

hind margins of tergites being more whitish; the following parts in 2 are very

pale yellowish or pallid; front part or half of frons, genae, palps, head below

anteriorly and continuous broadly to halfway up on each side behind eyes,

humeral tubercle and anterior spiracular area, notopleural ridge, more extensive

area below wing-basis, postalar calli, ligamentous connection, hind border of

scutellum, hind margins of tergites (slightly broader than in 3) and broader

hind margins of sternites, prosternal part to a variable extent, propleural part

just above front coxae, a longitudinal band along middle of pleurae, and greater

part of metapleural region in front of and below halteres; knees in 3 yellowish

red and in 9 more yellowish as well as apices of coxae, especially front ones;

integument of body dull, with greyish bloom, espeically in ¢.

Vestiture with the hairs longer and much denser than in namaensis, longer

in ¢ than in Q, entirely silvery whitish.

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE 303

Head (cf. Fig. 3E) with the eyes in ¢ in contact above for a distance about

24 times length of ocellar tubercle, the line of contact rather deeply impressed

and facets in upper half of eyes rather coarse; interocular space on vertex in

2 about twice width of ocellar tubercle; transverse frontal depression in 9

deep and distinct; antennal joint 1 about 1}-2 times length of small transverse

joint 2; joint 3 at least twice length of joints 1 and 2 combined, subparallel-

sided, its dorsal subapical spine short and rather stoutish; palps appearing

rather long, conspicuously visible, almost, or about, as long as antennal join

3; proboscis about 1,4-1,68 mm long.

Wings vitrous hyaline, iridescent; veins yellowish; middle cross vein at

about, or a little more than, a third of discoidal cell; part of third vein between

middle cross vein and second submarginal cell distinctly shorter than length

of latter cell; knobs of halteres entirely whitish, pallid, or yellowish in both sexes.

From 2 g¢ and 2 99, including the type specimens, in the South African

Museum.

Length of body: about 2,6-3,08 mm

Length of wing: about 2,88-3,4 mm

Distribution

Bushmanland: Aggenys between Springbok and Pella (Museum staff,

October 1939) (types); Pofadder (Museum staff, October 1939).

From both namaensis and triseriatellus this species may at once be dis-

tinguished by the much denser and longer vestiture, the relatively more con-

spicuous and longer palps, the comparatively longer second submarginal cell

in relation to part of third vein between it and middle cross vein, the position

of the latter cross vein, and in @ the less extensive yellow markings on head

and body.

Oligodranes flavifemoris sp. nov.

This species resembles puberulus very closely, but differs from it in the

following respects:

Body with the integument of the frontal triangle, sides of face and upper

parts of genae in 3 entirely dark, not yellowish, but covered with pale or greyish

white bloom; head below in @ slightly more extensively yellowish; hind margins

of tergites in 2 distinctly more broadly pallid or yellowish, sometimes occupying

nearly half of tergites posteriorly; hind margins of sternites in both sexes

distinctly very much more broadly yellowish, the entire venter in some 39

sometimes tending to be yellowish; legs with the coxae and femora in 2 mainly

and conspicuously yellow, not black as in 2 puberulus, only the upper or anterior

surfaces in apical halves of at least front and hind femora darkened or brownish

to a variable extent as in 99 of namaensis.

Vestiture as dense as in puberulus.

Head with antennal joint 1 apparently a little longer, especially in 8,

quite 24 times as long as small transverse joint 2; proboscis about 1,2-1,66

mm long.

304 ANNALS OF THE SOUTH AFRICAN MUSEUM

Wings, even though faintly tinted milky whitish, distinctly more so than

in puberulus; veins slightly darker, more brownish; knobs of halteres in 3

darkened above.

From | g and 8 99, including the types, in the South African Museum.

Length of body: about 2,32-3,64 mm

Length of wing: about 2,6-3,6 mm

Distribution

Southern coastal region: Pearly Beach in the Bredasdorp District (Museum

staff, December 1958) (types). Koup Karoo: Laingsburg District (Museum

staff, February 1938). West coastal region (Olifants River Valley): Bulhoek

between Clanwilliam and Klawer (Museum expedition, October 1950).

Oligodranes anomalus sp. nov.

Fig. 3B, D

Oligodranes namaensis (described as 2) Hesse, 1938: 864, fig. 263a (n. syn.).

As has been stated in my supplementary redescription of namaensis in

this paper, the discovery of the real ° of the latter species eliminates the specimen

described by me in 1938 as the 9 of namaensis. This specimen moreover is not

a 2 but a J in which the eyes are broadly separated above as in 99 of this genus.

It belongs to a newly discovered section of Oligodranes in which the ¢¢ are

not holoptic but dichoptic as in 99 where the eyes are not in contact above

but widely separated. This representative of the section by itself is characterized

as follows:

Body and legs mainly black, but with the following parts pale yellowish

or pallid: extreme anterior part (around antennal insertions) of frons, genae,

buccal cavity, palps, humeral angles and anterior spiracular area, notopleural

fold, area just below wing-bases, postalar calli and area below it, ligamentous

connection between squama and scutellum, part of metapleural region below

halteres, hind margin of metapleural part, narrow hind margins of tergites

and sternites, and hind margin of last sternite (or tergite); coxae and legs more

deep dark reddish brown than black, the apices of coxae, especially front ones,

and the knees slightly paler, more yellowish reddish; integument dull.

Vestiture with the hairs on body and legs sparse, slightly denser on abdomen,

entirely pale or whitish.

Head (cf. Fig. 3B) with the eyes broadly separated above, about 1§ times

width of ocellar tubercle; frons almost parallel-sided, slightly depressed centrally

at about middle, not deeply and transversely as in 99 of Oligodranes; antennal

joint | (cf. fig. 263a in Hesse 1938: 864 and Fig. 3B) very short, only a very

little longer than joint 2; joint 3 about twice as long as joints | and 2 combined,

about 22 as long as broad, its subterminal spine short, but rather stoutish;

palps about as long as antennal joint 3, but not easily seen at base of proboscis ;

proboscis about 1,08 mm long.

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE 305

Wings hyaline, iridescent, with a very faint whitish subopacity in certain

lights; veins yellowish brown; middle cross vein at a little more than basal

third (6: 15) of discoidal cell; part of third vein between cross vein and second

submarginal cell shorter than latter cell; knobs of halteres entirely whitish.

Legs with the left hind one shown in Figure 3D.

From a single 3 specimen originally erroneously described as the @ allo-

type of namaensis Hesse.

Length of body: about 1,88 mm

Length of wing: about 2,4 mm

Distribution

Namaqualand: Kamieskroon (Museum staff, September 1930).

Oligodranes elegans Hesse

Oligodranes elegans Hesse, 1938: 865.

The original description of this species was based on two specimens from

Aus in South West Africa, in the British Museum (Natural History). These

two specimens were taken to be 99, but in view of the fact that the allotype of

namaensis Hesse was mistaken by me for a 9 on account of its broadly separated

eyes and that a subsequent re-examination of this specimen, and the discovery

of the real 9 of namaensis, proved it to be a 3 belonging to a previously

unsuspected section of Oligodranes in which the 33 have separated eyes as in

a similar section of the genus Apolysis, it is more than probable that I confused

the sexes and that the two 9° of elegans may prove to be two 3.3. This suspicion

is supported by the fact that the ¢¢ of all the known South African species of

Oligodranes are mainly or predominantly black and that the known 99 have

more extensive yellow colouring on the body or are even predominantly

yellowish.

On the other hand the fact that the elegans specimens, according to my

original description, have a more distinct medial frontal depression and also

an anterior frontal depression however indicates a @ sex.

REVISED DESCRIPTIVE KEY TO ALL THE KNOWN SOUTH AFRICAN SPECIES OF

Oligodranes

1 (a) Eyes in actual contact above for a relatively long distance, their upper facets distinctly

much coarser than lower ones; frontal triangle or frons anteriorly usually more

extensively pallid or yellowish or appearing yellowish in certain lights; legs (cf

Fig. 3C) usually relatively longer, more slender, the first posterior tarsal joint usually

longer, distinctly more than half length of the tibia me 2

(6) Eyes broadly separated above, nearly or about 14 to about 2 times width of ocellar

tubercle, their upper facets not coarser than lower ones; frons in front not or less

extensively pallid and, if yellowish at all, then only narrowly so along upper margin

of buccal cavity; legs (cf. Fig. 3D) on the whole shorter, stouter, the first posterior

tarsal joint sometimes scarcely, or only a little more than, half length of the tibia 5

306

2 (a)

(5)

3 (a)

(5)

5 (a)

(6)

(d)

ANNALS OF THE SOUTH AFRICAN MUSEUM

Hairs on body and legs shorter, sparser, less dense, more inconspicuous; knees

distinctly much paler, more distinctly yellowish; line of contact between eyes not,

or less, deeply impressed; palps shorter, not conspicuous, apparently much shorter

than antennal joint 3; hind margins of tergites tending to be Tae more broadly

pallid or yellowish .. 3 od 3

Hairs on body and legs distinctly ‘longer, ‘denser, much more ‘conspicuous; knees

darker, or at least not conspicuously pale; line of contact between eyes distinctly

more deeply impressed; palps distinctly more conspicuous, apparently longer,

almost, or about, as long as antennal ge 3; hind margins of i tending to be

more narrowly pallid or yellowish . oie a ah 4

Hind margins of tergites and sternites, especially ean not or scarcely, or only

very narrowly, pallid or yellowish; veins in wings darker, more brownish; distance

between middle cross vein and base of second submarginal cell distinctly, or much,

longer than latter cell; middle cross vein usually at about, or less than, basal third of

discoidal cell; palps slightly longer, subequal to, or isa a little shorter than, antennal

joint 3. . & namaensis Hesse

Hind margins oF tergites and sternites Gicaaciy more broadly, or more conspicuously,

yellowish, sometimes even extensively so; veins in wings paler, more yellowish;

distance between middle cross vein and base of second submarginal cell only a little

longer, sometimes subequal to, even slightly shorter, than cell itself; middle cross

vein usually at a little more than basal third of discoidal cell; palps distinctly much

shorter, much shorter than antennal joint 3 .. 6 triseriatellus sp. nov.

Integument of frontal triangle and upper part of genae, sides of face and genae

pallid or yellowish; knobs of halteres entirely whitish or pale yellowish; middle

cross vein in wings slightly farther away from base of discoidal cell, at about between

basal third and fourth; apical stalk of anal cell relatively shorter, much shorter than

apical cross vein of discoidal cell; veins in wings paler, more yellowish

3 puberulus sp. nov.

Integument of frontal triangle, sides of face, and upper part of genae black, but with

greyish white or pale tomentum; knobs of halteres darkened above; middle cross

vein in wings slightly nearer base of discoidal cell, at only about basal fourth; apical

stalk of anal cell relatively longer, only a little shorter than apical cross vein of dis-

coidal cell; veins in wings darker, more brownish Ay 3 flavifemoris sp. nov.

Humeral angles, upper pleural parts, postalar calli, and narrow hind margins of

tergites and sternites distinctly pallid or yellowish; antennal joint 1 shorter, scarcely,

or only a little, longer than joint 2; palps shorter than antennal joint 3; proboscis

slightly shorter, a little less than 1,5 mm; wing-veins slightly darker

3 anomalus sp. nov. (= 2 namaensis Hesse)

Humeral angles, pleurae, postalar calli, and hind margins of tergites entirely black

like rest of body, only hind margins of sternites obscurely, very narrowly pallid;

antennal joint 1 longer, quite 14 times length of joint 2; palps longer, quite as long

as, or even slightly longer than, antennal joint 3; proboscis slightly longer, quite,

or a little more than, 1,5 mm; wing-veins paler, more yellowish

elegans Hesse (if original specimens be 3d)

Body with much or more extensive yellow coloration, the greater part of frons,

sides of occiput behind eyes, or even greater part of occiput, broadish sides of

thorax in front of wings, postalar calli, even base of thorax, greater part of pleurae,

entire scutellum, broad hind margins of tergites and sternites, and greater part of

femora more extensively or conspicuously yellow; distance between middle cross

vein and base of second submarginal cell in wings subequal to, or longer than, cell

itself; hairs on body and legs distinctly shorter and sparser_.. ; :3 2

Body mainly black or with distinctly less extensive yellow coloration, the greater

part of frons, upper half of or entire occiput, greater part of thorax, even on sides,

greater part of pleurae, entire or at least basal half of scutellum, and in some forms

the legs, black, and only humeral angles, upper part of pleurae just below wings,

SOUTH AFRICAN SPECIES OF PHTHIRIINAE AND USIINAE 307

narrower postalar calli, narrower hind margins of tergites and sternites, and in

some forms the femora, being yellow; distance between middle cross vein and base

of second submarginal cell usually shorter, or much shorter, than cell itself; hairs

on body and legs distinctly longer and denser .. uy 4 ve ne 3

2 (a) Occiput behind ocellar tubercle with a central dark band; base of frons or more

often the middle and sides of frons usually dark or black; sides and base of thorax

above distinctly less broadly or extensively yellow, the part on each side just above

wing-base black and not continuous; yellow hind margins of tergites, though

broadened on sides, distinctly much narrower, the black not tending to be broken

up to form three longitudinal rows of patches or spots... .. 9 namaensis Hesse

(6) Entire occiput behind ocellar tubercle and eyes and entire frons (excepting sometimes

a narrow median spot or line at about middle) yellow; sides and base of thorax

above distinctly more broadly and extensively yellow, the part on each side above

wing-base continuously yellow, but above it with a rather conspicuous round black

spot confluent with black discal part; yellow on abdomen much more extensive,

the black basal parts of tergites tending to be reduced and broken up by the yellow

to form three longitudinal rows of black spots or patches, becoming smaller

posteriorly... i 7 ue .. 9° triseriatellus sp. nov. (and forms of it)

3 (a) Frons anteriorly, lower half of sides of head behind eyes, humeral tubercle, postalar

calli, hind part of scutellum, propleural part, middle and hind part of pleurae, and

hind margins of tergites and sternites pallid or yellowish; interocular space on

vertex at least 2} times width of ocellar tubercle; veins in wings darker, yellowish

brown or brownish; knobs of halteres entirely very pale yellowish white to ivory

whitish; hairs on body slightly denser and longer iy 4

(6) Head and body, especially thorax, scutellum, pleurae, and abdomen above, entirely

black, only narrowish or obscure hind margins of sternites pallid; interocular space

on vertex narrower, only about 2 times width of tubercle; veins in wings more

yellowish; knobs of halteres more yellowish or even tinged slightly brownish above;

hairs on body sparser and shorter .. elegans Hesse (if original specimens be 29)

4 (a) Legs entirely dark or blackish brown, only apices of coxae and extreme apices of

femora yellowish; head below mainly dark or black, especially hinder half; yellowish

hind margins of tergites distinctly narrower, very much narrower than half length of

tergites; antennal joint 1 appearing shorter, usually less than twice length of joint 2;

wings scarcely or not faintly tinted milky whitish at Q puberulus sp. nov.

(5) Legs with the greater part of, or entire, coxae and femora yellow, only upper surfaces

of front and hind femora basally and apically, or only apically, darkened to a

variable extent; head below mainly or entirely yellowish; yellowish hind margins

of tergites distinctly broader towards apex, almost or only a little narrower than

half length of tergites; antennal joint 1 slightly longer, at least twice length of joint 2;

wings distinctly, though faintly, tinted more milky whitish

SUMMARY

In this paper 14 species of the genus Phthiria (subfamily Phthiriinae) are

dealt with and of these 9 are described as new. A new variety of an already

described species is added. Supplementary redescriptions of some species,

previously inadequately described or described from one sex only, are given,

and the synonymy of some others, described by me in 1938, is established.

In the two genera Apolysis and Oligodranes, which Hull transferred to

the subfamily Usiinae in 1973, 12 new species of the former and 4 of the latter

are described. As in the case of Apolysis it has been found that the J¢ of some

South African species of the genus Oligodranes are dichoptic like the 99.

308 ANNALS OF THE SOUTH AFRICAN MUSEUM

To supplement the descriptions three figures portraying heads, antennae,

wings and legs of some species are given.

Revised and descriptive keys to all the known South African species of

the three genera dealt with are also given.

ACKNOWLEDGEMENTS

In the preparation of this revision my thanks are due to Dr B. Stucken-

berg of the Natal Museum for submitting certain specimens of Phthiria from

Natal for identification; to the late Mr C. Thorn, a Technical Assistant of

the South African Museum, who accompanied me on various collecting trips

to Namaqualand and the North-western Cape and who collected many species

of Bombyliidae frequenting flowers; and lastly to Mr H. Zinn, the other

Technical Assistant, who made many collecting trips with me to the Karoo and

adjacent parts and whose contributions to the bombyliid collections in the

Museum are second to none.

REFERENCES

Bezzi, M. 1921. On the bombyliid fauna of South Africa (Diptera) as represented in the

South African Museum. Ann. S. Afr. Mus. 18: 1-180.

Bezzi, M. 1922. Enumeratio Bombyliidarum (Dipt.) quas ex Africa meridionali Dr. H. Brauns

misit. Broteria (Zool.) 20: 64-86.

Bezzi, M. 1924. The Bombyliidae of the Ethiopian Region: 109-115. London: British Museum

(Natural History).

ENGEL, E. O. 1933. Bombyliidae. In: LINDNER, E. Die Fliegen der palaearktischen Region.

Lief. 69. Stuttgart: Schweizerbart.

Hesse, A. J. 1938. A revision of the Bombyliidae (Diptera) of southern Africa. Ann. S. Afr.

Mus. 34: 1-1053.

Hesse, A. J. 1962. Apolysis lindneri sp. nov., eine neue Bombyliide aus Siidafrika (Dipt.).

Stuttg. Beitr. Naturk. 80: 1-2.

HuLt, F. M. 1973. Bee flies of the world: the genera of the family Bombyliidae. Bu//. U.S.

natn. Mus. 286: i—xii, 1-687.

Loew, H. 1860. Die Dipteren-Fauna Siid-Afrika’s. Erste Abteilung. Berlin: Bossellmann.

(Abh. naturw. Ver. Sachsen Thiir. Halle 2, 1861: 57-402.) (Bidrag till kannendomen om

Afrikas Dipteren. Ofvers. K. Vetensk Akad. Forh. Stockh. 17, 1860 (1861): 81-97.)

INSTRUCTIONS TO AUTHORS

Based on

CONFERENCE OF BIOLOGICAL EDITORS, COMMITTEE ON FORM AND STYLE. 1960.

Style manual for biological journals. Washington: American Institute of Biological Sciences.

MANUSCRIPT

To be typewritten, double spaced, with good margins arranged in the following order:

(1) Heading, consisting of informative but brief title, name(s) of author(s), address(es) of

author(s), number of illustrations (figures, enumerated maps and tables) in the article.

(2) Contents. (3) The main text, divided into principal divisions with major headings; sub-

headings to be used sparingly and enumeration of headings to be avoided. (4) Summary.

(5) Acknowledgements. (6) References, as below.

Figure captions and tables to be on separate sheets.

ILLUSTRATIONS

To be reducible to 12 cm x 18 cm (19 cm including caption). A metric scale to appear

with all photographs.

All illustrations to be termed figures (plates are not printed; half-tones will appear in their

proper place in the text), with arabic numbering; items of composite figures to be designated

by capital letters (A, B, C etc.).

REFERENCES

Harvard system (name and year) to be used: author’s name and year of publication given

in text; full references at the end of the article, arranged alphabetically by names, chronologi-

cally within each name, with suffixes a, b, etc. to the year for more than one paper by the same

author in that year.

For books give title in italics, edition, volume number, place of publication, publisher.

For journal articles give title of article, title of journal in italics (abbreviated according to the

World list of scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses,

volume number, part number (only if independently paged) in parentheses, pagination.

Examples (note capitalization and punctuation)

BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FIscHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris

88: 100-140.

FISCHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des

littorines. Archs Zool. exp. gén. 74: 627-634.

Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee

region of Ceylon. Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the

Indian Ocean. Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. |

Zoologische und anthropologische Ergebnisse einer Forschungsreise im westlichen und

zentralen Siid-Afrika. 4: 269-270. Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16:

269-270.

ZOOLOGICAL NOMENCLATURE

To be governed by the rulings of the latest International code of zoological nomenclature

issued by the International Trust for Zoological Nomenclature (particularly articles 22 and

51). The Harvard system of reference to be used in the synonymy lists, with the full

references incorporated in the list at the end of the article, and not given in contracted form

in the synonymy list.

Example

Scalaria coronata Lamarck, 1816: pl. 451, figs 5 a, b; Liste: 11. Turton, 1932: 80.

A. J. Hesse

ADDITIONS TO THE SOUTH AFRICAN

SPECIES OF PHTHIRITINAE AND USIINAE

(DIPTERA: BOMBYLIIDAE)

WITH KEYS TO ALL THE.KNOWN SPieae

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