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

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FiscHer, P. H., Duvat, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archives de zoologie

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Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. Annals and

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Koun, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bulletin of

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THIELE, J. 1910. Mollusca. B. Polyplacophora, Gastropoda marina, Bivalvia. In: ScHuLTzE, L. Zoologische und anthro-

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(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 104 Band

February 1996 Februarie

Part 10 #£Deel

B\0O” S

Gms

THE GENUS BRANCHIPODOPSIS

(CRUSTACEA, BRANCHIOPODA, ANOSTRACA)

IN SOUTHERN AFRICA.

MORPHOLOGY, DISTRIBUTION,

RELATIONSHIPS AND

THE DESCRIPTION OF FIVE NEW SPECIES

M. L. HAMER

C. C. APPLETON

Cape Town Kaapstad

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D4271

THE GENUS BRANCHIPODOPSIS

(CRUSTACEA, BRANCHIOPODA, ANOSTRACA)

IN SOUTHERN AFRICA.

MORPHOLOGY, DISTRIBUTION, RELATIONSHIPS AND

THE DESCRIPTION OF FIVE NEW SPECIES

M. L. HAMER* & C. C. APPLETON

Department of Zoology & Entomology, University of Natal, Private Bag XOI,

Scottsville, 3209, South Africa

(With 41 figures and 1 table)

[MS accepted 27 January 1995]

ABSTRACT

The southern African species of Branchipodopsis are reviewed. Eleven previously known

species are redescribed and five new species (B. barnardi, B. dayae, B. drakensbergensis,

B. hutchinsoni and B. underbergensis) are described. The male second antennae (clypeus),

cercopods, last abdominal segments and penes, as well as extensions of the genital segments

of the females of certain species, are illustrated as are species distributions. The 16 species

have been divided into groups based on a tentative cladistic analysis and a key, using the

morphology of the clypeus, is presented. Intraspecific variation in two widespread species,

B. tridens and B. wolfi, is illustrated and discussed. The characters used in the taxonomy of

the genus Branchipodopsis, aspects of species diversity, habitats, dispersal and future

Tesearch are commented on.

CONTENTS

PAGE

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13 GGT TOTS 68 cer SABE D SSB OER ACH TcCREe (an CSROET Ha BOGn OC ANE an SHAE AO nN HEMT et Akt Ri tent ores 315

Key to the southern African species of the genus Branchipodopsis ................+0+++ 365

LI SGICT OTe cedar citer at Be cB ae caaR EHO smelt tap ant airtitrit actin ena nl ARTA gene ae 366

PAE MME WICE SCIEN Sic e dectee acck eee ee nena enero soos e oes season ee seu eR TERE ae eee ennen 373

ETE TOSS a coat CRE RRR ORO REPEC CEN E Ee OSE ROROR ERASE ORTGEr CRT Ere CM SAE MmmninSr sd Sena an BAA an 374

/s UIE PICIT ELS (CRY ZE Te Cer Meted He ABR a ors ci Ren ans mali Angee, Sabir ARAL EA ne ERE a rien 375

INTRODUCTION

The anostracan family Branchipodidae is represented in Africa, Asia,

Australia and Europe and is characterized by having the basal joints of the

second antennae of the male fused to form a so-called ‘clypeus’. In addition,

according to Linder (1941), the genital segments of the male are negligibly

swollen ventrally and there are no distinct seminal vesicles. The penes are close

together, the basal parts are rigid and possess proximal outgrowths and usually

distal ones as well. The retractable parts of the penes are armed with

longitudinal rows of spines. The female ovisac is short and broad.

* present address: Natal Museum, Private Bag 9070, Pietermaritzburg, 3200

311

Ann. S. Afr. Mus. 104 (10), 1996: 311-377, 41 figs, 1 table.

312 ANNALS OF THE SOUTH AFRICAN MUSEUM

Initially, the family Branchipodidae Baird, 1852, included most of the

known anostracan genera. Daday (1910) restricted the family to four genera,

which he placed in two subfamilies: the Parartemiinae (which included the

genus Parartemia), and the Branchipodinae (with three genera—Branchipodop-

sis, Tanymastix and Branchipus). This subfamilial distinction was based on the

number of abdominal segments (eight in Parartemiinae and nine in Branchi-

podinae) and whether the cercopods articulated with the telson (Branchipodinae)

or not (Parartemiinae). The monotypic genus Metabranchipus was later

described by Masi (1925) and included in the family Branchipodidae. Barnard

(1929) and Linder (1941) disputed Daday’s division. Linder (1941) pointed out

that both groups have eight abdominal segments and a telson, although the

anterior boundary is almost obscure in Parartemia, and that the cercopods of

this genus are as articulated as in the other genera. In addition, Linder (1941)

placed little taxonomic value on the number of abdominal segments.

Linder (1941) provided a detailed synopsis of the family and stated that the

degree to which the basal joints of the second antennae are joined varies from

almost complete in Branchipus to the form where only the proximal halves are

joined, as in those specimens of Branchipodopsis hodgsoni examined by him.

He suggested that, in all of the Anostraca, it is difficult to decide the degree to

which the basal joints must be fused to be called a clypeus and this presents

some problems in determining the limits of the family Branchipodidae. Sub-

sequent to Linder’s (1941) monograph, few taxonomic contributions, apart from

single species descriptions, have been made to the family. Brtek (1972)

described a new genus, Janymastigites, from North Africa and divided the

Branchipodidae into three subfamilies, the Branchipodinae (Branchipodopsis,

Branchipus and Parartemia), the Tanymastiginae (Tanymastix and Tany-

mastigites), and a third comprising the single genus, Metabranchipus. An in-

depth review of the family and the characters uniting the genera currently

included in it is thus necessary. This is, unfortunately, beyond the scope of the

present study, which focuses on southern African anostracans. Only the genus

Branchipodopsis is represented in the region and it has been neglected, probably

to an even greater extent than the other genera of the Branchipodidae.

Sars (1898) described the first Branchipodopsis species, B. hodgsoni, from

Port Elizabeth, South Africa, and established the genus but he did not provide a

generic diagnosis. Branchipus abiadi Brauer, 1877, a species from Rio de Oro,

Mauritania in North West Africa, was later transferred to the genus Branchipo-

dopsis by Linder (1941). Sars (1901) described Branchipodopsis affinis from

Central Asia. In 1910, Daday described a further three species from southern

Africa. In addition, he provided a diagnosis of the genus. He used the median

ventral process, a small outgrowth from the anteroventral edge of the clypeus,

the presence of two flat outgrowths on either side of it, and two more lamelli-

form and short outgrowths near the apical joints as diagnostic generic charac-

ters. Barnard (1929), however, in his review of the southern African phyllo-

pods, suggested that the median ventral process is not always present; it may be

present in some specimens of a species but not in others. Barnard (1929) pro-

vided illustrations, descriptions and locality data for then known southern Afri-

can species, as well as for seven new species of Branchipodopsis from the

region.

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 313

Linder (1941) provided additional data on Branchipodopsis, including details

of the second maxilla, the structure and setation of the thoracopods, and the

male genital segments and penes. He confirmed Barnard’s (1929) comment on

the median ventral process but suggested that it might be present in all speci-

mens of a species but might have broken off in some. The proximal part of this

structure is very narrow and could break without leaving any trace of its

presence on the clypeus. Linder (1941) considered the lamelliform setiferous

processes at the apical joint to be the most unique characteristic of Branchi-

podopsis, since these processes are not present in any other anostracans.

Subsequent to the work of Linder (1941), the genus Branchipodopsis has

been neglected. A number of articles have been published on the Asian species

B. acanthopenes (Mulhotra & Duda, 1970), B. terpogossiani Smirnov, 1936,

and B. affinis (Roen 1952; Hartland-Rowe 1968; Tiwari 1972; Battish 1983;

Brtek et al. 1984; Vechov 1988), but only two contributions have been made on

the African representatives of the genus. Brehm (1958) illustrated, but did not

name or provide detailed data of, one possible new species and other specimens

that he identified as B. karroensis Barnard, 1929. These specimens, however,

cannot be located and his identification is probably inaccurate. Loffler (1968)

described B. candea, a new species from Mt Elgon in Central Africa, and

provided data on B. wolfi Daday, 1910. Two recent publications (Hamer &

Appleton 1991; Seaman ef al. 1991) provided brief ecological information on

B. wolfi and B. tridens Daday, 1910, respectively.

In southern Africa, sporadic collecting of Anostraca over the last 15 or so

years and a more concentrated effort over the last four years have produced

additional material of the genus Branchipodopsis. Examination of this has

revealed the need to update Barnard’s (1929) review in terms of distribution

data, illustrations and descriptions, as well as to describe five new species. A

key, based on the male clypeus, is provided and a division of the genus into

groups based on cladograms is presented. The results of and problems associ-

ated with a cladistic analysis are outlined. Comments are made on the characters

used for species identification and aspects of intraspecific variation, speciation,

diversity, distribution and habitats are discussed.

MATERIALS AND METHODS

List of museums. The following abbreviations are used to indicate the

museum collections included in this study: AM—Albany Museum, Grahams-

town, South Africa; BMNH—British Museum of Natural History, London,

England; HNHM—Hungarian Natural History Museum, Budapest, Hungary;

USNM—National Museum of Natural History, Smithsonian Institution, Wash-

ington, U.S.A.; NM—Natal Museum, Pietermaritzburg, South Africa; SAM—

South African Museum, Cape Town, South Africa; SMN—State Museum of

Namibia, Windhoek, Namibia; TM—Transvaal Museum, Pretoria, South

Africa; NMZ—National Museum, Bulawayo, Zimbabwe.

Material catalogued TM VLKE was collected on the Vernay-Lang Kalahari

Expedition by the Transvaal Museum. Material catalogued under DB was

collected by Dr T. Rutherford of the University of Lesotho and Dr J. A. Day of

314 ANNALS OF THE SOUTH AFRICAN MUSEUM

the University of Cape Town and was provided by Dr Denton Belk of Our Lady

of the Lake University, Texas, U.S.A., from his personal collection.

Collecting. Hand-held dip nets of varying sizes and with mesh of 0.5-

2.0 mm were used to collect anostracans. Specimens were preserved in 70 per

cent ethanol or 4 per cent formalin. Dried pool sediment was collected from a

number of localities for later rehydration in the laboratory. Hatched anostracans

were cultured until mature enough to be identified.

Illustrations. Drawings were made using a Wild M-5 dissecting microscope

or a Leitz Labor Lux 12 compound microscope and drawing tube. The clypeus

was dissected from those species for which sufficient material was available,

and cleaned and dehydrated in a graded ethanol series. Specimens were then

critical point dried, mounted on stubs and coated with 20 nm of gold for

scanning electron microscope observation using a Hitachi S-570 at an

accelerating voltage of 10 or 12 kV. For some species, eggs were removed

from the egg sac of females, rinsed in distilled water, oven-dried at 50°C for

24 hours, coated and viewed.

Measurements. Specimens of suitable quality were measured using a dis-

secting microscope and graticule. All measurements are presented as total body

length (mean + standard deviation where n > 10, or as a range of lengths where

n < 10), from the front of the head (excluding the clypeus) to the tip of the

cercopods (excluding the setae) for sexually mature specimens.

Fig. 1. Diagram of idealized male second antennae (clypeus) illustrating terminology used in

text and figures. Abbreviations: aj—apical joint; bj—basal joint; bp—basal process;

c—clypeus; il—inner lobe; Ip—lamelliform process; mvp—median ventral process; ol—outer

lobe; pr—projection; tu—tubercle.

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA S15

Terminology. The terminology used to describe the regions of the male

clypeus is from Barnard (1929) and Linder (1941) and is illustrated in Figure 1.

Each half of the clypeus (c) consists of a broad, stout basal joint (bj) and a

slender, inwardly curved apical joint (aj). A lamelliform, setiferous process (Ip)

is present on the basal joint just proximal to the apical joint. The basal process

(bp) is a somewhat flattened process situated anteriorly/anterodorsally on the

basal joint. The basal process may have adjacent, apical lobes; these are termed

the inner (il) and outer lobe (01), which are rounded or conical structures of

varying widths. Tubercles (tu) are smaller, digitiform projections that may be

present apically or dorsally on the basal process. A smaller, acute projection

(pr) may also be present near the apex of the basal process. The median ventral

process (mvp) is a small, usually spinose, rounded or oval process, situated on

the ventral surface in an anterior, median position on the basal joint. This

structure is not present in all species.

Systematic analysis. Cladograms were initially drafted manually using

Hennig augmentation, and were then generated using the ‘mhennig’, ‘bb*’ and

‘ie-’ commands of the program Hennig86. The following characters were used

for the analyses: pair of spines on ventral surface of last abdominal segment;

proportion of the cercopod inner margin with spines; cercopod shape; cercopod

length in relation to body length; size of lamelliform process; curve of apical

joint; distal inflation of posterior margin of apical joint; shape of the apex of the

apical joint; presence of median ventral process; basal process shape; relative

size of lobes of basal process; width of basal process in relation to height;

presence of a dorsal tubercle on basal process; presence of a small projection on

apex of inner lobe; large dorsal spinous projection at base of basal process.

These were coded as binary characters, and it was assumed that the simplest

form represented the primitive state. The genus Parartemia was used as the out-

group in the analysis.

The 16 species were divided into groups using a combination of the clado-

grams generated. Those species that were consistently grouped and that exhibit

distinct similarities in the form of the clypeus were placed in the same group.

This analysis and division must, however, for reasons discussed later, be con-

sidered as tentative.

DESCRIPTIONS

Class BRANCHIOPODA Latreille, 1817

Order ANOSTRACA Sars, 1867

Family Branchipodidae Baird, 1852

Genus Branchipodopsis Sars, 1898

Branchipodopsis Sars, 1898: 26. Daday, 1910: 293. Barnard, 1929: 192. Linder, 1941: 228.

Eubranchinella Daday, 1910: 256.

Mongolobranchipus Dybowski, 1927: 39.

Type species. Branchipodopsis hodgsoni Sars, 1898: 26, pl. 3.

316 ANNALS OF THE SOUTH AFRICAN MUSEUM

Diagnosis

Males with basal joints of second antennae fused to form a clypeus. Each

basal joint broad with a basal process in an anteromedian position and a setifer-

ous lamelliform process distally. Apical joint slender and curved inwards. A

median ventral process may be present (Fig. 1). Basal part of penes with

rounded projection on median margin about halfway along length. Apical evers-

ible part of penes with one longitudinal and a second, less regular row or

scattered arrangement of spines (Fig. 2A-D). Thoracopods with a single pre-

epipodite (Fig. 2E). Egg sac of female short and oval.

Typical morphology of labrum and maxillae is illustrated in Figure 3.

Fig. 2. A. Penes of Branchipodopsis wolfi (DB817). Ai. Ventral view of right penis; arrow

indicates peg-like projection; 1—row of regular spinous processes; 2—irregular row of

spinous processes. Aii. Dorsal view of right penis. B. Ventral view of right penis of

B. wolfi (NMZ/Cr 6); arrow indicates peg-like projection; mp—median rounded projection.

C. Lateral view of right penis of B. wolfi (NMZ/Cr6). D. Ventral view of right penis of

B. natalensis (DB818). E. Right thoracopod 4 of B. tridens (AMLEN 150A).

Scales: A-D = 1 mm; E = 0.5 mm.

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 317

Branchipodopsis barnardi sp. nov.

Figs 3-5

Material

Holotype. SAM-A40839, 1 3 (13.9 mm); collected from rock pool

(1.5 x 2 m), 15 cm deep, Sehlabatebe National Park area, Drakensberg, Natal—

Lesotho border, by M. Hamer and O. Wirminghaus, 23 April 1993.

Paratypes. SAM-A40840, 23 ¢ (14.8 + 1.0 mm), 44 2 (15.5 + 1.5 mm);

same collection data as holotype.

Other material examined. SAM-A40841, 1 3 (14.6 mm), 2 2 (15.0,

15.3 mm); collected from type locality, by O. Wirminghaus and H. Adie,

11 April 1993.

of portion of setae (position indicated by arrow on Fig. 3A) showing variation in shape of

scale-like structure detail of setae on maxilla I. C. Maxilla II. D. Lateral view of labrum.

E. Dorsal view of labrum. Scales: A, E = 0.1 mm; B, C = 0.05 mm; D = 0.05 mm;

C = 0.5 mm.

318 ANNALS OF THE SOUTH AFRICAN MUSEUM

B———

NCD Ral ve

Fig. 4. Branchipodopsis barnardi sp. nov. (SAM-A40840, paratype). A. Clypeus.

B. Dorsal view of cercopods of male. C. Ventral view of posterior region of abdomen;

arrow indicates spine. D. Ventral view of basal part of penes; arrow indicates median

projection. E. Lateral view of genital segments of female showing egg sac and bulbous

processes (indicated by arrow). F. Dorsal view of female antenna. G-I. Basal processes of

three specimens illustrating intrapopulation variation in shape of apical region.

Scales = 0.5 mm.

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 319

Description of male

Clypeus. Basal process long and slender. Apical region with digitiform inner

lobe and short, flat outer lobe (Figs 4A, 5A-B). Reticulate patterning on dorsal

proximal surface of basal process (Fig. 5C). Lamelliform process long, promi-

nent and elliptical in shape with long sparse setae (Fig. 4A). Apical joint

slender, distal third weakly curved inwards and apex blunt (Figs 4A, 5A).

Median ventral process absent.

Cercopods. Moderate length (ratio to body length about 0.2: 1). Proximally

stout, tapering distally and with outer margins weakly convex (Fig. 4B). Short

to medium length setae on outer margins, replaced by three spines distally.

Inner margins with small patch of sparse setae proximally, followed by 12-15

evenly spaced spines (Fig. 4B).

Abdominal spines. Pair of short, stout and blunt spines present on ventral

surface of last abdominal segment (Fig. 4C).

Penes. Basal part short and stout, with proximal region bulbous and with

indistinct rounded projection on inner margin (Fig. 4D).

Description of female

Thoracic segments. Last thoracic segment with pair of obvious, rounded and

bulbous extensions dorsally (Fig. 4E).

Antenna. Apical point on second antenna almost one-half length of antenna

(Fig. 4F).

Egg morphology

Surface of egg with irregularly shaped, five-sided depressions separated by

high rounded ridges (Fig. 5D).

Differential diagnosis

Branchipodopsis barnardi is easily distinguished from all other species of

the genus by the large elliptical lamelliform process, the slender apical joint and

the long, digitiform and distally bilobed shape of the basal process.

Distribution

Branchipodopsis barnardi has, to date, only been collected from rock pools

in the high-altitude (2000 m) Drakensberg region, on the border of Sehlabatebe

National Park in Lesotho and the Bushman’s Nek area of Natal (Fig. 39).

Remarks

In the cladograms presented in Figure 41, B. barnardi did not show a close

relationship with any of the other species. As a result of the simple structure of

the basal process and shape of the apical joint, the position of B. barnardi on

this cladogram suggests a primitive form. Branchipodopsis barnardi does, how-

ever, share certain characters with B. drepane Barnard, 1929. Both species

have slender bilobed basal processes, with a smaller outer lobe, and the apical

joint is smooth and weakly curved. For these reasons, B. barnardi and B. dre-

pane have been included in the same group (see Table 1).

320 ANNALS OF THE SOUTH AFRICAN MUSEUM

The colour of live specimens varied according to the pool from which they

were collected. One population (used for DNA extraction) was a pale orange-

red colour before preservation, whereas the type specimens were a pale aqua

blue. The labral area and a thin stripe along the gut region were red in all speci-

mens. There was also some degree of intrapopulation variation in the shape of

the apex of the basal process as illustrated (Fig. 4G-I).

Etymology

Branchipodopsis barnardi is named for the late Dr K. H. Barnard, of the

SAM, who was the author of the first review of the southern African ‘Phyllo-

poda’ in 1929 and who collected a large number of specimens currently housed

in that institution.

#9 +. ec RTS

Fig. 5. Branchipodopsis barnardi sp. nov. (SAM-A40840, paratype). A. Dorsal view of

clypeus. B. Basal processes; il—inner lobe; arrow indicates position of outer lobe.

C. Apical region of left basal process showing reticulate patterning. D. Egg.

Scales: A-B = 0.5 mm; C-D = 100 um.

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 321

Branchipodopsis browni Barnard, 1924

Figs 6-7

Branchipodopsis browni Barnard, 1924: 217, pl. 26 (fig. 4); 1929: 198, fig. Sa.

Material

Syntypes. SAM-A6705, large number of specimens, 24 ¢ measured (12.0 +

0.7 mm), 26 2 measured (13.3 + 0.6 mm); collected from Kalkfontein South,

Great Namaqualand, Namibia, by J. S. Brown, date unknown.

Other material examined. SAM-A7600, 7 ¢ (13.1-15.6 mm), 4 2 (17.5-

20.0 mm); collected from Williston, Cape Province in 1939, collector unknown.

AM LEN 79B, 6 ¢ (11.8-13.0 mm), 7 2 (13.1-14.8 mm); collected from a

large (20 x 30 m), shallow (20-35 cm) turbid pool, sparsely vegetated with

grass, 10 km south of Carnarvon on the road to Loxton, Karoo, Cape Province,

by M. Hamer, 16 February 1990. AM LEN 164A, 3 ¢ (6.6, 8.0, 8.1 mm), 1 2

(6.9 mm); hatched from dried sediment collected 8 km north-east of Fraserburg,

Karoo, Cape Province, by M. Hamer, 6 January 1993. AM 614/93, 1 ¢

(14.2 mm), 1 2 (16.5 mm); collected from inundated area along road, 8 km

from Richmond, by K. Martens, 6 April 1993.

Redescription of adult male

Clypeus. Basal process conical proximally, apically inflated and bluntly

rounded, with a series of short denticles on apex (Figs 6A, 7A-C). A distal,

anteriorly curved arid pointed projection on inner margin of basal process

(Fig. 7B-C). One or two small, acute projections dorsomedially near apex of

basal process (Fig. 7B-C). Lamelliform process very large (as long as basal

process) and oval (Fig. 6A). Apical joint slender, long, smooth and curved

inwards with apex slightly inflated and blunt (Fig. 6A). Median ventral process

large (approximately one-half length of basal process), oval and spinose

(Figs 7A, 8A).

Cercopods. About one-fifth of body length, margins straight, with long,

plumose setae along outer margins and proximal two-thirds of inner margins.

Distal third of inner margins with 6-8 widely spaced, long spines (Fig. 6B).

Abdominal spines. Two small ventral spines on last abdominal segment

(Fig. 6C).

Penes. Median projection bluntly rounded and prominent. Distal region of

basal part with small but distinct peg-like projection on outer margin (Fig. 6D).

Remarks

Barnard (1924) stated that B. browni closely resembles B. hodgsoni but that

the presence of abdominal spines in the former species separates them. The

arrangement of the basal process and associated projections is, however, quite

distinct in these two species. In the cladogram (see Fig. 41), B. browni was

positioned closest to B. wolfi. These two species have an apical projection on

unlobed or unilobed basal processes and distinctly bent apical joints.

Variation in the size of the dorsomedian projection of the basal process and

the apical denticles is illustrated in Figure 7B and C.

B22 ANNALS OF THE SOUTH AFRICAN MUSEUM

| \ |

Sein ce i RB i

eee A,B

—— aD

Fig. 6. Branchipodopsis browni. A. Dorsal view of clypeus (AMLEN79B). B. Cercopods

of male (AMLEN79B). C. Ventral view of last abdominal segment of male (SAM-A6705,

syntype); arrow indicates spine. D. Ventral view of right penis with apical region extended

(SAM-A6705); arrow indicates peg-like projection. E. Female second antenna (SAM-

A6705). Scales = 0.5 mm.

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 323

Distribution

Branchipodopsis browni has only been reported from the above five

localities in the arid southern Namibian and Karoo regions (Fig. 38).

Fig. 7. Branchipodopsis browni. A. Basal processes and median ventral process of

AMLEN 164A specimen. B. Detail of apex of left basal process; arrow indicates median,

dorsal projection. C. Detail of left basal process of AMLEN79B specimen; arrow indicates

small median, dorsal projection. Scales: A = 150 wm; B = 50 um; C = 100 um.

Branchipodopsis dayae sp. nov.

Figs 8-9

Material

Holotype. SAM-A40842, 1 3 (17.3 mm); collected Eland’s Bay, Cape

Province, by J. Lighton, 23 November 1980.

324 ANNALS OF THE SOUTH AFRICAN MUSEUM

Paratypes. SAM-A40843, 2 3 (15.3, 16.9 mm), 2 2 (12.8, 14.4 mm);

collection data as for holotype.

Other material examined. SAM-A40849, 1 3, in poor condition; hatched in

laboratory from sediment collected from dry depression on roadside,

Koppieskraal Pan, approximately 36 km from Namibian border on R31, Cape

Province, by M. Hamer, 22 December 1992.

Fig. 8. Branchipodopsis dayae sp. nov. (SAM-A40843, paratype). A. Dorsal view of

clypeus; arrow indicates small projection on apical, median margin of basal process.

B. Cercopods of male. C. Ventral view of penes. D. Median view of apical, everted part of

penes; arrow indicates blunt, peg-like projection. Scales = 0.5 mm.

Description of male

Clypeus. Basal process broad and apically bilobed. Outer lobe rounded,

inner more conical, with small acute projection apically on median margin

(Figs 8A, 9A-B). Lamelliform process small and round (Fig. 8A). Apical joint

weakly curved inwards, distal region foot-shaped (Figs 8A, 9A, C). Median

ventral process small, round and with short, blunt spines (Fig. 9D).

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 325

Fig. 9. Branchipodopsis dayae sp. nov. (SAM-A40843, paratype). A. Dorsal view of

clypeus. B. Apical region of left basal process (arrow indicates small projection on inner

lobe). C. Distal region of apical joint. D. Dorsal view of median ventral process.

Scales: A = 0.5 mm; B-C = 100 pm; D = 50 pm.

Cercopods. Moderate length (ratio to body length approximately 0.25: 1),

outer margins weakly convex and with short setae along almost entire length

except distal region where these replaced by 3-5 widely spaced, short spines

(Fig. 8B). Proximal fifth of inner margins with short setae and remainder of

length with 12 short spines (Fig. 8B).

Abdominal spines. A pair of blunt spines present on ventral surface of last

abdominal segment.

Penes. Basal part stout, inner margin with distinct conical projection proxi-

mally and lateral margin with indistinct peg-like projection distally (Fig. 8C-D).

Differential diagnosis

The distal region of the apical joint and the shape of the two lobes of the

basal process of B. dayae are unique to this species. Branchipodopsis drepane

also has characteristic apical joint apices but these are hook-shaped rather than

326 ANNALS OF THE SOUTH AFRICAN MUSEUM

foot-shaped. This species also shares the small, rounded lamelliform processes

with B. dayae but the basal processes of the two species are clearly different.

The basal process morphology of B. dayae resembles that of B. karroensis

Barnard, 1929, and these two species were shown to be closely associated by

the cladistic analyses (Fig. 41).

Distribution

Branchipodopsis dayae has been collected from Eland’s Bay on the Cape

west coast and from the Gordonia region between Namibia and Botswana

(Fig. 40). It is possible that this species also occurs in other waterbodies in the

area between these two localities.

Etymology

Branchipodopsis dayae is named after Dr J. A. Day of the University of

Cape Town, who has contributed to the knowledge of temporary waterbodies in

southern Africa. She also collected numerous Anostraca and kindly made her

collection, including the type specimens of this species, available for study.

Branchipodopsis drakensbergensis sp. nov.

Figs 10-13

Material

Holotype. SAM-A40834, 3 (12.5 mm); collected from small (40 x 50 cm)

rock pool in boulder, 15 cm deep, Loteni Nature Reserve, Drakensberg, Natal,

by M. Hamer, 6 January 1992.

Paratypes. SAM-A40835, 14 ¢ (12.6 + 0.5 mm), 5 2? (11.9-12.8 mm);

same collection data as holotype specimen.

Other material examined. SAM-A40836, 17 ¢ (11.1 + 0.5 mm), 34 2 (10.4

+ 0.5 mm); collected from small pool (30 x 40 cm) in boulder, 10 cm deep,

Loteni Nature Reserve, Drakensberg, Natal, by M. Hamer, 6 January 1992.

AM LEN 158A, 7 ¢ (7.5-10.5 mm), 5 2 (9.0-18.1 mm); collected in rock pool

on top of Prentjiesberg, Farm Montana, Ugie, Cape Province, by R. McC. Pott,

4 April 1991. AMLEN 159A, 12 ¢ (2 with damaged cercopods, remainder

13.1-13.9 mm), 4 2 (12.3-14.0 mm); locality as for AM LEN 158A; collected

by R. McC. Pott, 22 January 1992. SAM-A11593, 4 ¢ (9.4-12.0 mm), 5 ?

(10.0-11.3 mm); collected from Giant’s Castle, Drakensberg, Natal, by Ewer,

1951. AM REA77A, 12 ¢ (8.9 + 0.6 mm), 26 2 (9.1 + 0.8 mm); collected

from temporary vlei at Witkoppies, Benoni, Transvaal, by P. A. Reavell,

31 April 1971.

Description of male

Clypeus. Basal process slender, distally bilobed, with inner lobe twice as

long as outer, both apically narrow and rounded (Figs 10A, 11A-B). Inner lobe

with 1-2 small projections medianly on apex (Fig. 12A-C). Large spinous

projection proximally on dorsal surface of basal process (Figs 10B-C, 11A-C),

one-half to two-thirds length of basal process and with apical denticles or scales

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 327

Fig. 10. Branchipodopsis drakensbergensis sp. nov. A. Dorsal view of clypeus (SAM-

A40835, paratype). B. Lateral view of basal process of Loteni specimens (SAM-A40835).

C. Lateral view of basal process of Ugie specimen (AMLEN 159A); il—inner lobe; ol—outer

lobe; s—spinous projection. D. Cercopods of male (SAM-A11593). E. Ventral view of

basal part of penes (SAM-A40835). F. Lateral view of genital segments of female

(AM LEN 159A) showing egg sac. G. Female antenna (AMLEN 159A). Scales = 0.5 mm.

(Fig. 13A—-C). Median ventral process prominent, ovate and with long spines

distally (Figs 10A, 11B). Lamelliform process ovate, with long setae (Figs 10A,

11C). Apical joint strongly curved inwards with rounded apices (Fig. 10A).

Cercopods. Moderate length (ratio to body length approximately 0.25: 1).

Outer margins strongly convex, with setae along entire length. Patch of setae on

328 ANNALS OF THE SOUTH AFRICAN MUSEUM

proximal fifth of inner margins, followed by 17-19 evenly spaced spines

(Fig. 10D).

Abdominal spines. A pair of prominent spines on last abdominal segment.

Penes. Basal part long, with distinct, round projection about halfway along

inner margin (Fig. 10E).

Egg morphology

Eggs with deeply crumpled appearance created by irregular narrow

depressions separated by high ridges (Fig. 13D). Surface of eggs roughened

with numerous small lumps.

Fig. 11. Branchipodopsis drakensbergensis sp. nov. A. Dorsal view of basal processes of

clypeus, Loteni specimen (SAM-A40835, paratype); outer lobe of left basal process not

visible; arrow indicates outer lobe of right basal process; s—spinous projection. B. Basal

processes of Ugie specimen (AM LEN 159A); arrow indicates median ventral process.

C. Left basal process of Benoni specimens (AM REA77A); Ip—lamelliform process.

Scales = 100 pm.

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 329

Differential diagnosis

Branchipodopsis drakensbergensis resembles B. tridens in the presence of a

dorsal projection on the basal process and the bilobed basal process. However,

the prominent median ventral process, absent or vestigial in B. tridens, and the

distinctly slender basal process as opposed to the broad shape of these structures

in B. tridens, separate the two species. As a result of this similarity, the Giant’s

Castle specimens (SAM-A11595) were previously labelled as B. tridens in the

SAM collection.

Fig. 12. Branchipodopsis drakensbergensis sp. nov. A. Detail of apex of inner lobe of basal

process, Loteni specimen (SAM-A40835, paratype). B. Detail of apex of inner lobe of basal

process, Ugie specimen (AMLEN 159A). C. Detail of apex of inner lobe of basal process,

Benoni specimen (AMREA77A). Scales: A-B = 10 ym; C = 50 um.

330 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 13. Branchipodopsis drakensbergensis sp. nov. A. Basal process, illustrating apex :

spinous process (indicated by arrow), Loteni specimen (SAM-A40835, paratype). B. Detail

of apex of spinous process, Ugie specimen (AMLEN 159A). C. Detail of apex of spinous

process, Benoni specimen (AMREA77A). D. Egg.

Scales: A, D = 100 um; B = 50 um; C = 10 pm.

Distribution

Branchipodopsis drakensbergensis occurs in high-altitude rock pools in the

southern and central Drakensberg. The single Transvaal locality could indicate a

wider distribution for this species as well as a wider habitat range (Fig. 39).

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 331

Remarks

There is some variation in clypeus morphology between populations from

different localities. For example, the specimens from Loteni have a flattened,

spatulate process on the apex of the inner lobe of the basal process as opposed

to a pointed one in the specimens from Benoni and Ugie (Fig. 12A—C) and there

is a slight difference in the shape of the basal process and spinous projection

(Fig. 10B-C). The latter projection has long extensions apically in the Loteni

specimens (Fig. 13A), whereas these are flattened and scale-like in the

specimens from Benoni (Fig. 13C) and somewhat intermediate in those from

Ugie (Fig. 13B). In addition, the cercopods of the Benoni specimens have fewer

spines (9) on the inner margin than the specimens from the Drakensberg. These

differences could indicate two distinct species but, in the light of the large

amount of interpopulation variation in other widespread species of the genus and

because of the overall similarity in clypeus shape, a single species has been

described.

In terms of relationships to other species of the genus, B. drakensbergensis

could be related to B. tridens, since both species have a spinous projection on

the basal process and bilobed basal processes. However, the cladograms

(Fig. 41) do not indicate this relationship and the spinous projection could be a

homoplasious character.

Etymology

Branchipodopsis drakensbergensis is named after the mountain range from

which most of the material was collected.

Branchipodopsis drepane Barnard, 1929

Fig. 14

Branchipodopsis drepane Barnard, 1929: 199, fig. 5e-f.

Material

Holotype. SAM-A7259, 1 6 (12.4 mm), distal part of right apical joint

damaged; collected from Great Fish River, near Gibeon, Namibia, by R. W.

Tucker, date unknown.

Other material examined. BMNH 1932.2.25.41, 1 ¢ (12.3 mm); same

locality and collection data as type specimen.

Redescription of adult male

Clypeus. Basal process long and slender with distinct indentation about half-

way along lateral margin. Distally, basal process with a smaller outer lobe and

a higher and larger, rounded inner lobe (Fig. 14A). Lamelliform process small,

slender and oval (Fig. 14A). Apical joint weakly curved inwards, with

hook-shaped, acute apex (Fig. 14A). Median ventral process obovate, apically

flat and with surface covered by small spines (Fig. 14B).

Cercopods. Ratio to body length approximately 0.26: 1, slender, and with

outer margins only slightly convex. Outer margins and proximal half of inner

332 ANNALS OF THE SOUTH AFRICAN MUSEUM

margins with plumose setae. Distally, inner margins with six large, widely

spaced spines (Fig. 14C).

Abdominal spines. Last abdominal segment with two stout, blunt spines on

ventral surface.

Penes. Proximal region of basal part with obscure bulge on lateral surface,

followed by small rounded projection on median margin (Fig. 14D).

Distribution

Branchipodopsis drepane has only been collected from the type locality in

southern Namibia (Fig. 38).

Fig. 14. Branchipodopsis drepane (BMNH 1932.2.25.41). A. Dorsal view of clypeus;

il—inner lobe; ol—outer lobe. B. Ventral view of basal processes and median ventral

process. C. Cercopods of male. D. Ventral view of basal part of penes; arrows indicate

lateral and median proximal projections.

Scales = 0.5 mm.

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 333

Branchipodopsis hodgsoni Sars, 1898

Figs 15-16

Branchipodopsis hodgsoni Sars, 1898: 26, pl. 3 (figs 1-9). Daday, 1910: 301, fig. 51.

Barnard, 1929: 194, fig. 5k-1.

Branchipodopsis braueri Pesta, 1921: 94.

Material

Syntypes. SAM-A1488, 1 3 (11.6 mm), 1 ? with damaged abdomen and

cercopods; hatched from dried sediment collected from Port Elizabeth, Cape

Province, by J. V. Hodgson, May 1897.

Other material examined. SAM-A6721, 14 6 (14.0 + 1.3 mm), 4 2

(10.5-16.5 mm); collected from Ashton, Cape Province, collector unknown,

27 August 1910. SAM-A13631, 1 ¢ (11.9 mm), 5 2 (8.8-14.4 mm); collected

from freshwater pool, Kenton-on-Sea, Cape Province, by R. A. Jubb, April

Fig. 15. Branchipodopsis hodgsoni. A. Dorsal view of clypeus (HNHMI/A-76); arrows

indicate dorsal and ventral lobes of basal process. B. Basal processes and median ventral

process of AMLEN 149A specimen; 1—dorsal lobe; 2—ventral lobe. C. Lateral view of

basal process (AMLEN 149A). D. Cercopods of male (SAM-A6721). E. Ventral view of

basal part of right penis (AM LEN 149A); arrow indicates peg-like process. F. Female

second antenna (SAM-A6721). Scales = 0.5 mm.

334 ANNALS OF THE SOUTH AFRICAN MUSEUM

1963. AM 107A, 1 ¢ with right half of clypeus damaged (9.2 mm), 1 2

(10.4 mm); same collection data as SAM-A13631. HNHMI/A-76, 1 6

(11.8 mm), 1 ? (8.3 mm); data as for syntypes. AM LEN 149A, 7 ¢

(6.3-12.6 mm); collected from a shallow (5-10 cm), clear stretch of water with

sparse, dead vegetation on side of main road from Bredasdorp to Struisbaai,

Cape Province, by M. Hamer, 17 July 1990.

Redescription of adult male

Clypeus. Basal process with broad, double-lobed base; second lobe pos-

itioned slightly ventral to first (Figs 1SA-C, 16A). Narrow, medianly directed

apex with two small pointed tubercles (Figs 15A, 16A). Lamelliform process

oval and slender (Fig. 15A). Apical joint strongly curved, with distinct bend and

blunt apex (Fig. 15A). Median ventral process large, oval and with blunt spines

(Fig. 16B).

Cercopods. Moderate length (ratio to total body length approximately

0.24: 1). Outer margins convex with short plumose setae along entire length.

Proximal two-fifths of inner margins with plumose setae, followed by

8-10 acute, evenly spaced spines (Fig. 15D).

Abdominal spines. Last abdominal segment without ventral spines.

Penes. Basal part with large rounded projection on median margin and

small, but distinct, peg-like projection distally on lateral margin (Fig. 15E).

Distribution

Branchipodopsis hodgsoni appears to be concentrated in the south-east

coastal region of South Africa, from Kenton-on-Sea in the north to Bredasdorp

in the south. The locality of Ashton in the western Cape represents the furthest

inland record (Fig. 38).

Fig. 16. Branchipodopsis hodgsoni (AMLEN 149A). A. Right basal process; 1—dorsal

lobe; 2—ventral lobe. B. Dorsal view of median ventral process.

Scales: A = 50 pm; B = 25 um.

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 335

Remarks

Branchipodopsis hodgsoni is quite distinct from the other species of the

genus in that the basal process has a bilobed appearance, with one lobe ventral

to the other. This double-lobed arrangement is not always as evident as

illustrated but this is usually a result of the preservation of the specimens. The

two small apical tubercles on the dorsal lobe are also unique to B. hodgsoni.

Branchipodopsis hutchinsoni sp. nov.

Figs 17-18

Material

Holotype. SAM-A40844, 1 3 (12.3 mm); collected from heavily vegetated

roadside ditch (30 x 1 m), 20-30 cm deep, 30 km along dirt road from Hutch-

inson to Richmond, Karoo, Cape Province, by M. Hamer, 16 February 1990.

Paratypes. SAM-A40845, 18 ¢ (12.0 + 0.7 mm), 32 2 (11.4 + 0.8 mm);

collection data as for holotype.

Description of male

Clypeus. Basal process broad, apically bilobed, lobes subequal, rounded-

conical, with inner lobe slightly more conical than outer (Figs 17A, 18A-B).

Inner lobe with small, acute process on median, apical surface (Fig. 18B). A

medianly directed, conical tubercle on dorsal surface at base of inner lobe

(Fig. 18A-C). Lamelliform process obovate, with setulose scalloped margin

(Fig. 17A). Apical joint with distinct inward bend, apically blunt and broad

(Fig. 17A). Median ventral process prominent, ovate and with short spines

(Fig. 17A).

Cercopods. Moderate length (ratio to body length approximately 0.25: 1),

outer margins strongly convex with medium length setae along four-fifths of

length, distal fifth with 3-6 short spines. Inner margins with patch of setae on

proximal quarter, followed by 12-16 strong spines (Fig. 17B).

Abdominal spines. A small blunt spine present medianly at the base of each

cercopod. A pair of rounded processes present ventrally on penultimate segment

(Fig. 17C).

Penes. Basal part slender with small rounded projection proximally on

median margin and a narrow, peg-like projection apically on lateral margin

(Fig. 17D).

Egg morphology

Surface of egg with mildly crumpled appearance created by three- to five-

sided depressions separated by broad ridges with sharp crests (Fig. 18D).

Differential diagnosis

Branchipodopsis hutchinsoni, B. karroensis, B. dayae and B. natalensis

Barnard, 1929, all have broad bilobed basal processes with both lobes of

approximately equal size. The inner lobe of both B. karroensis and B. dayae

also has a small apical projection but neither species has a dorsal tubercle such

336 ANNALS OF THE SOUTH AFRICAN MUSEUM

as that of B. hutchinsoni and B. natalensis. The latter two species can be

separated by the rounder shape of the basal processes, the strongly curved

cercopods and the scalloped lamelliform processes of B. hutchinsoni.

Distribution

Branchipodopsis hutchinsoni has, to date, only been collected from the type

locality in the Karoo (Fig. 39).

Fig. 17. Branchipodopsis hutchinsoni sp. nov. (SAM-A40845, paratype). A. Dorsal view

of clypeus; arrow indicates small projection on apex of inner lobe. B. Cercopods of male.

C. Ventral view of posterior region of abdomen showing rounded projections on penultimate

segment (indicated by large arrows) and blunt spinous processes (indicated by small arrows).

D. Ventral view of genital region of male showing basal parts of penes; arrows indicate

proximal median process and distal lateral process. E. Dorsal view of genital segments of

female showing absence of outgrowths in this region. F. Female second antenna.

Scales = 0.5 mm.

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 337

Fig. 18. Branchipodopsis hutchinsoni sp. nov. (SAM-A40845, paratype). A. Dorsal view

of basal processes of clypeus. B. Detail of right basal process. C. Detail of apex of median

dorsal projection on basal process. D. Egg.

Scales: A-B, D = 100 um; C = 10 um.

Etymology

Branchipodopsis hutchinsoni is named in honour of the limnologist Pro-

fessor Evelyn Hutchinson, in memory of his contributions to the knowledge of

freshwater habitats.

Branchipodopsis kalaharensis Daday, 1910

Figs 19-20

Branchipodopsis kalaharensis Daday, 1910: 296, fig. 49. Barnard, 1929: 194, fig. 5j; 1935:

487.

Material

Syntypes. 2 3 and 2 2 housed in the Senckenberg Museum; collected from

the Kalahari, by D. Schultze, date unknown (Daday 1910).

338 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material examined. SAM-A11594, 2 6, 1 with damaged cercopods (other

11.2 mm), 6 2 (10.0-13.5 mm); collected from Kanke Pan (90 miles west of

Molepole), Botswana, by the Vernay-Lang Kalahari Expedition, 19 March

1930. TM VLKE94, 8 ¢ (10.0-14.0 mm); same collection data as

SAM-A11594. TM VLKE460a, 8 ¢ (10.6-12.8 mm); collected from

Sunnyside, Botswana, by the Vernay-Lang Kalahari Expedition, 20 April 1930.

Redescription of adult male

Clypeus. Basal process broad, apically bilobed with a narrow conical inner

lobe and an outer, less prominent but similarly shaped lobe (Figs 19A, 20A-B).

Lamelliform process narrow and oval (Fig. 19A). Apical joint strongly curved

inwards and with distinct bend (Figs 19A, 20A). Median ventral process absent

or obscure.

Cercopods. Long (ratio to body length approximately 0.3: 1), with proximal

two-thirds of outer margins convex and with plumose setae of moderate length.

Distal third of outer margins with 8-10 widely spaced, spiniform setae. Proxi-

mal quarter of inner margins with plumose setae, followed by approximately

15 regularly spaced, strong spines (Fig. 19B).

Fig. 19. Branchipodopsis kalaharensis (TM VLKE94). A. Dorsal view of clypeus; il—inner

lobe; ol—outer lobe. B. Cercopods of male. C. Ventral view of basal part of left penis;

arrow indicates blunt apical projection. Scales = 0.5 mm.

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 339

Abdominal spines. Last abdominal segment without ventral spines.

Penes. Basal part with large rounded projection on proximal region of

median margin and obscure blunt projection on apex of lateral margin

(Fig. 19C).

Remarks .

Daday (1910) attributed Branchipodopsis kalaharensis to Wolf, who initially

examined the type material. The description by Wolf was, however, never

published (Forr6 & Brtek 1984).

Fig. 20. Branchipodopsis kalaharensis (TM VLKE 94). A. Dorsal view of clypeus.

B. Detail of right basal process. Scales: A = 0.5 mm; B = 100 um.

Daday (1910) suggested that, because of the simple structure of the basal

process of this species, the other species of the genus are derived from it. Of the

five species that had been described at that stage, Daday named the Indian

species B. affinis as the closest relative of B. kalaharensis. In southern Africa,

there are a number of species that have bilobed basal processes but the lobes in

these species are much rounder than in B. kalaharensis. It is, however, quite

possible that the simple bilobed shape of the basal process of B. kalaharensis

does represent the ancestral form for species such as B. dayae, B. hutchinsoni,

B. karroensis and B. natalensis. This is not reflected by the cladistic analyses

(Fig. 41), where B. kalaharensis was linked to B. simplex Barnard, 1924, which

has an unlobed basal process, but has apical joints of similar shape and also

lacks a distinct median ventral process.

Distribution

Branchipodopsis kalaharensis has only been collected from central

Botswana (Fig. 38).

340 ANNALS OF THE SOUTH AFRICAN MUSEUM

Branchipodopsis kaokoensis Barnard, 1929

Fig. 21

Branchipodopsis kaokoensis Barnard, 1929: 200, fig. 5q.

Material

Barnard (1929) stated that the type material, consisting of one male and four

females, was housed in the SAM but this could not be located. The collection

data and catalogue number of four specimens in the BMNH suggest that these

are the types supposedly in the SAM. Barnard (1929) gave the length as 13 mm,

but his measurements did not include cercopods and this could account for the

difference in measurements presented here, or he could have presented an

average length for all five specimens. Details for these specimens are as

follows: BMNH 1932.2.25.42-45, 1 3 with left half of clypeus removed

(21.0 mm), 1 2? (13.5 mm); collected from Choabendus (115 miles north-west of

Outjo), Kaokoveld, Namibia, by K. H. Barnard, date unknown.

Redescription of adult male

Clypeus. Basal process broad, apically trilobed with inner lobe digitiform,

the outer lobe similarly shaped but more acute and the median lobe the largest

and apically flattened. A long, conical, spiniform projection present dorsally at

proximal region of basal process (Fig. 21A). Lamelliform process narrow and

ovate. Apical joint stout, curved inwards and apically deeply bifid. A short,

spiniform projection on dorsal surface approximately midway along apical joint,

followed by a larger, triangular projection ventrolaterally just proximal to apex

(Fig. 21A). Median ventral process absent.

Cercopods. Moderate length (ratio to body length approximately 0.26: 1),

slender and with outer margins slightly convex. Outer margins with moderately

long plumose setae along about half of length, followed by 5-6 small, sharp

spines. Inner margins with plumose setae along proximal quarter of length,

followed by 15 prominent spines (Fig. 21B).

Abdominal spines. Last abdominal segment with two large, broad spines on

ventral surface (Fig. 21C).

Penes. Basal part short and broad without distinct median projections.

Remarks

The penes of the single specimen examined could be shrunken by the

preservative since they look different to those of other species of the genus.

Barnard (1929) remarked on the similarity between B. kaokoensis and B. tri-

dens, but stated that no B. tridens specimens had any hint of projections on the

apical joint of the clypeus. However, the two species were consistently very

closely linked in the cladistic analyses (Fig. 41) and there is little doubt that

B. kaokoensis and B. tridens represent advanced, closely related forms.

Distribution

Branchipodopsis kaokoensis has only been recorded from the type locality in

Namibia (Fig. 38).

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 341

Fig. 21. Branchipodopsis kaokoensis (BMNH 1932.2.25.42-45). A. Dorsal view of

clypeus; arrows indicate projections on apical joint. B. Cercopods of male. C. Ventral view

of last abdominal segment of male. Scales = 0.5 mm.

Branchipodopsis karroensis Barnard, 1929

Fig. 22

Branchipodopsis karroensis Barnard, 1929: 198, fig. Sm-n.

Material

Syntypes. SAM-A5919, 2 3 (9.8 mm; 12.0 mm), 5 ? (9.5-12.3 mm);

collected from Hoogeveld, south-west of Beaufort West, Cape Province, by

S. H. Haughton, date unknown.

Other material examined. BMNH 1932.2.25.36-40, 1 ¢ (10.0 mm), 1 @

(11.3 mm); data as for syntypes.

Redescription of adult male

Clypeus. Basal process broad. Apex with two rounded lobes and a small

acute process on the median margin of inner lobe. Lamelliform process oval.

Apical joint curved inwards with slight inflation just proximal to blunt apex.

Median ventral process with broad base and narrower distal region (Fig. 22A).

Cercopods. Long (ratio to body length approximately 0.30: 1). Outer mar-

gins weakly convex, and with setae along almost entire length. Distal region

342 ANNALS OF THE SOUTH AFRICAN MUSEUM

| B

aan B, C

(A Z ee ey

NY A, D

\ uy

Fig. 22. Branchipodopsis karroensis. A. Dorsal view of clypeus (SAM-A5919, syntype);

arrow indicates projection on median margin of inner lobe. B. Cercopods of male

(BMNH 1932.2.25.36-40). C. Left penis with apical region partially extended

(SAM-A5919); arrows indicate rounded projection on median margin and peg-like projection

on lateral margin. D. Female second antenna (SAM-A5919). Scales = 0.5 mm.

with three spines. Proximal half of inner margins with widely spaced, short

setae followed by 8-9 spines (Fig. 22B).

Abdominal spines. Ventral surface of last abdominal segment with two short

but strong spines.

Penes. Basal part with distinct, rounded projection proximally on median

margin and prominent peg-like projection apically on lateral margin (Fig. 22C).

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 343

Description of female

Second antenna. Apical pointed process long (one-half the length of antenna)

(Fig. 22D).

Distribution

Branchipodopsis karroensis has only been recorded from the type locality in

the Karoo (Fig. 38).

Remarks

Brehm (1958) identified specimens from Blouberg in the Transvaal as

B. karroensis but this record is considered invalid, since his illustrations of the

clypeus show two distinct tubercles on the basal process. These specimens are

most probably B. wolfi.

The penes of B. karroensis have been illustrated in the everted position. The

apical part does appear shorter than in other species and it is possible that it is

not fully everted. However, with the limited number of specimens available, the

structure cannot be verified.

Branchipodopsis natalensis Barnard, 1929

Figs 23-24

Branchipodopsis natalensis Barnard, 1929: 196, fig. 5o.

Material

Syntypes. NM 1384, 1 ¢ (5 mm), 2 ? (no measurements); collected from

Natal, half mile from Van Reenen (border of Orange Free State and Natal)

(Barnard 1929). These specimens, apparently housed in the Natal Museum,

Pietermaritzburg, could not be found in the institute’s collections.

Material examined. SAM-40846, 10 6 (12.5 + 0.8 mm), 1 2? (12.3 mm);

collected from a small depression in a boulder, Bushman’s Nek-Sehlabatebe

Game Reserve area, Drakensberg, Natal, by M. Hamer and O. Wirminghaus,

23 April 1993. SAM-A40847, 4 3 (9.4-9.8 mm); collected from a small rock

pool, Sehlabatebe Game Reserve area, Drakensberg, Lesotho, by M. Hamer

and O. Wirminghaus, 24 April 1993. DB 818, 3 6 (9.3, 10.8, 11.4 mm), 2 ?

(both 12.3 mm); collected from Sehlabatebe Game Reserve, Lesotho, by

T. Rutherford, 1985.

Redescription of adult male

Clypeus. Basal process apically bilobed with the inner lobe narrow and with

a minute apical projection. Outer lobe rounded. Conical tubercle present on

dorsal surface near base of inner lobe (Figs 23A, 24A-B). Lamelliform process

small and oval. Apical joint curved inwards and with rounded apex. Median

ventral process narrow and ovate.

Cercopods. Moderate length (approximate ratio to body length 0.2: a:

Outer margins convex with plumose setae along entire length. Inner margins

with sparse setae along proximal two-thirds; these replaced by 4-5 short spines

on distal third (Fig. 23B).

344 ANNALS OF THE SOUTH AFRICAN MUSEUM

Abdominal spines. Ventral surface of last abdominal segment with pair of

small spines (Fig. 23C).

Penes. Proximal region of median margin with small, angular projection

(Fig. 23D).

Fig. 23. Branchipodopsis natalensis (SAM-A40847). A. Dorsal view of clypeus; arrow

indicates small projection on inner lobe. B. Cercopods of male. C. Ventral view of last

abdominal segments of male; arrow indicates small spinous process. D. Ventral view of

male genital segments showing basal part of penes; arrow indicates median rounded

projection. E. Dorsal view of genital segments of female showing lateral extensions of last

thoracic segment (indicated by arrow). F. Female second antenna. Scales = 0.5 mm.

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 345

Fig. 24. Branchipodopsis natalensis (SAM-A40847). A. Dorsal view of left half of

clypeus. B. Detail of apical region of left basal process.

Scales: A = 0.5 mm; B = 100 um.

Description of female

Thoracic segments. Last thoracic segment with slightly laterally and pos-

teriorly extended surface (Fig. 23E).

Remarks

It is impossible to confirm the identification of the Sehlabatebe material

without the type specimens of B. natalensis. However, since both sets of speci-

mens come from high-altitude pools and there is a resemblance between the new

material and Barnard’s (1929) illustration and description of B. natalensis, it is

possible that a single species is represented.

Some of the specimens had distinctly rounded apices of the apical joints,

with the anteroventral surface expanded into a peg-like projection. There is,

however, some intraspecific variation in this character and, since it was not

described by Barnard (1929), it has been excluded from the above species

description.

The cladistic analyses group B. natalensis with B. dayae, B. karroensis and

B. hutchinsoni (Fig. 41). These species all share the bilobed basal process, with

a small projection on the inner margin near the apex.

Branchipodopsis scambus Barnard, 1929

Figs 25-26

Branchipodopsis scambus Barnard, 1929: 199, fig. 5p.

346 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 25. Branchipodopsis scambus. A. Dorsal view of clypeus (BMNH 1972.1.27.31-33).

B. Cercopods of male (AMLEN 18). C. Ventral view of basal part of penes (AM LEN 18);

arrow indicates small rounded median projection. Scale = 0.5 mm.

Material

Type material. Barnard (1929) stated that the type specimens were collected

in the Cape, Grahamstown, and were deposited in the Albany Museum. These

have not been located.

Material examined. BMNH 1972.1.27.31-33, 1 ¢ (12.8 mm), 1 2

(15.2 mm); locality and date unknown, collected by K. G. McKenzie.

AM LEN 18, 2 ¢ (10.2, 11.8 mm), 1 2 (15.0 mm); collected from flat, flooded

grassland along Grahamstown-Cradock road, by F. de Moor, K. Martens and

H. Barber, 28 November 1989. AM LEN 19, 1 ¢ (10.1 mm), same collection

data as AMLEN 18.

Redescription of adult male

Clypeus. Basal process slender, spiniform and apically acute (Figs 25A,

26A-B). Lamelliform process large and bluntly oval (Figs 25A, 26A). Apical

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 347

joint weakly curved, slender and smooth with rounded apex (Fig. 25A). Median

ventral process oval and with longer spines apically (Fig. 25A).

Cercopods. Short (ratio to body length approximately 0.11: 1), straight and

with plumose setae along entire length of both inner and outer margins,

excluding distal quarter of inner margins which have four spines and apices

which have two spines (Fig. 25B).

Abdominal spines. Pair of spines on ventral surface of last abdominal

segment.

Penes. Basal part short and with small projection halfway along median

margin (Fig. 25C).

Remarks

Barnard (1929) made no comment on the cercopods of B. scambus. In the

specimens examined, these are unusual for the genus in terms of length and the

small number of spines on the inner margins. These features, together with the

simple shape of the basal processes, could indicate that this species is a primi-

tive form of Branchipodopsis (Fig. 41).

Distribution

The species is known only from the type locality of Grahamstown and the

additional locality along the Grahamstown-Cradock road in the eastern Cape

(Fig. 38).

Fig. 26. Branchipodopsis scambus (AMLEN 18). A. Dorsal view of clypeus.

B. Detail of left basal process.

Scales: A = 0.5 mm; B = 100 pm.

348 ANNALS OF THE SOUTH AFRICAN MUSEUM

Branchipodopsis simplex Barnard, 1924

Fig. 27

Branchipodopsis simplex Barnard, 1924: 217, pl. 26 (figs 2-3); 1929: 196, fig. Sb-c.

Material

Syntypes. SAM-A6006, 3 ¢ (7.0, 8.0, 8.1 mm); collected from Eunda

(about 100 miles west-north-west Ondangua), Ovamboland, Namibia, by K. H.

Barnard, date unknown.

Other material examined. BMNH 1932.2.25.11-15, 1 3 (7.0 mm), 1 2

(8.3 mm); collected from Ovamboland, Namibia, collector and date unknown.

Redescription of adult male

Clypeus. Basal process with inflated proximal region, apically narrower and

rounded (Fig. 27A). Dorsal keel present along length of basal process

Fig. 27. Branchipodopsis simplex (SAM-A6006, syntype). A. Dorsal view of

clypeus; large arrow indicates dorsal keel; small arrow indicates apex of keel.

B. Lateral view of clypeus; arrow indicates keel. C. Cercopods of male. D. Ventral

view of basal part of left penis. Scale = 0.5 mm.

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 349

(Fig. 27B). Lamelliform process small and rounded (Fig. 27A). Apical joint

with distinct bend, distal to which posterior margin inflated. Apex of apical joint

blunt (Fig. 27A). Median ventral process absent.

Cercopods. Moderate length (ratio to body length approximately 0.25 : 1)

with outer margins weakly convex (Fig. 27C). Short setae on proximal part of

inner margins and along length of outer margins. Distal two-thirds of inner

margins with approximately 10 evenly spaced spines (Fig. 27C).

Abdominal spines. Spines on ventral surface of last segment absent.

Penes. Proximal region of basal part with median margin bulbous and with

rounded projection about halfway along length (Fig. 27D).

Remarks

The keel of the basal processes is difficult to see, particularly in such small

specimens, and is only obvious in lateral view (Fig. 27B). This character is aut-

apomorphic, but B. simplex was consistently grouped with B. kalaharensis in

the cladistic analyses (Fig. 41), even though the two species do not share many

basal process features.

Distribution

Branchipodopsis simplex has only been collected from the type locality in

Namibia (Fig. 38).

Branchipodopsis tridens Daday, 1910

Figs 28-31

Branchipodopsis tridens Daday, 1910: 308, fig. 53. Barnard, 1924: 217; 1929: 197, fig. Sd;

1935: 487. Linder, 1941: 228, fig. 30b.

Material

Type material. Consists of two male and two female specimens and is

housed in the Senckenberg Museum; collected from the Kalahari, by

D. Schultze, date unknown (Daday 1910).

Material examined. SAM-A5980, 1 3 (12.1 mm), 1 2 (11.8 mm); collected

from Papkuil, near Kimberley, Cape Province, by Miss Wilman, date unknown.

SAM-A7255, 1 ¢ with cercopods missing; collected from Bak River, Cape

Province, by K. H. Barnard, 1925. SAM-A6290, 1 ¢, poor condition; collected

from between Keimoes and Upington, Cape Province, 1909, collector unknown.

SAM-A7257, specimens dehydrated; collected from Outjo, Kaokoveld,

Namibia, by K. H. Barnard, 1926. SAM-A7256, specimens in poor condition;

collected from Narugas siding, Cape Province, by K. H. Barnard, 1925. SAM-

A7258, 2 ¢ in poor condition; collected from Kamanyab, Kaokoveld, Namibia,

by K. H. Barnard, 1926. SAM-A7267, 3 ¢ (12.5, 12.6 mm; 1 with cercopods

broken), 5 2? (10.4-12.3 mm); collected from Cauas Okawa, Kaokoveld,

Namibia, by K. H. Barnard, March 1926. SAM-A5922, 1 ¢ (19.0 mm);

collected from Great Fish River, near Gibeon, Namibia, by R. W. Tucker,

January 1916. SAM-A7599, 1 6 (cercopods removed), 12 ? (poor condition);

350 ANNALS OF THE SOUTH AFRICAN MUSEUM

collected from Amadap Valley, Little Namaqualand, Cape Province, by Dendy,

1938. SMN 51291, 8 3 (6.5-7.9 mm), 10 2 (5.4-7.4 mm); collected from small

pool in a rock kaross, Etosha Park, Namibia, by E. Griffen, 7 October 1986.

SMN 51197, large number of specimens, 25 ¢ measured (9.0 + 0.6 mm), 22 2

measured (8.3 + 0.6 mm); collected from large shallow rock pool at the base of

a rock hill (Bakenkop), Namib Naukluft Park, Namibia, by B. D. Collahan,

17 March 1986. SMN 51334, 1 3 with broken abdomen; collected from Charl

Marais Dam (Sukses Dam), Etosha Park, Namibia, by M. & P. Lindeque,

26 January 1988. TM VLKE443, large number of specimens, 33 ¢ measured

(10.6 + 1.2 mm), 25 2 measured (10.8 + 1.2 mm); collected from Gori Pan

(possibly Goru, Kangara), Botswana, by the VLKE, 20 April 1930.

HNHM 1/A-78, 3 3 (9.1, 9.2, 10.2 mm), 2 2 (8.7, 10.2 mm); collected from

Kalahari, Botswana, collector and date unknown. AM LEN 150A, large number

of specimens, many immature, 14 d measured (9.8 + 3.0 mm), 20 ° measured

(8.8 + 0.8 mm); collected from Leeubron, Etosha Park, Namibia, by A. Bow-

land, November 1985. AM LEN 156A, 1 ¢ (11.5 mm); hatched from sediment

(5 February 1993) collected from rock pool on top of Moonrock, Augrabies

Falls National Park, Cape Province, by M. Hamer, 19 December 1992.

AM LEN 163A, 6 ¢ (8.1-11.8 mm), 2 2 (9.3, 10.9 mm); hatched from sedi-

ment (2 April 1993) collected from dry pool, 40 km south of Kalahari Gemsbok

Park, Cape Province, by M. Hamer, 24 December 1992. SAM-A40850, 7 3

(10.6-14.4 mm); hatched from sediment collected from granite rock cavity,

10 km north-east of Gobabeb, Kuiseb River bed, by J. A. Day, 24 June 1981.

SAM-A40851, 2 3 (15.6, 16.3 mm); hatched from sediment collected from

Amichab, 140 km west of Rheboth, Namibia, by J. A. Day, 5 July 1981. SAM-

A40852, 1 ¢ (14.0 mm); locality data as for SAM- A40851; hatched from sedi-

ment collected by J. A. Day, 24 June 1981. DB 765, 1 3, 1 ? (both poor con-

dition); collected from series of pools at Blutkoppie on granite outcrop, Namib

Desert, Namibia, by J. A. Day, 27 March 1982. DB 767, 1 3 (18.8 mm), 5 @

(10.9-13.8 mm); hatched from sediment collected from Gemsbokwater, 18 km

north of Ganab, Namib Desert, Namibia, by J. A. Day, 14 May 1980. DB 768,

1 3, 1 2 (both poor condition); hatched from sediment collected from 4 km

north-north-west of Zebra Pan, Namib Naukluft Park, Namib Desert, Namibia,

by J. A. Day, 2-12 May 1980.

Branchipodopsis cf. tridens, Namibia. DB 761, 2 ¢ (12.1, 12.5 mm);

hatched from sediment collected from large gravel pool in desert plain near

Heinrichsberg, Namib Desert, Namibia, by J. A. Day, 5 July 1981. DB 763,

4 5 (10.0-11.0 mm); collected from Mirabib, 2 sinkholes in granite inselberg,

Namib Desert, Namibia, by J. A. Day, 26 March 1982. DB 764, 9 3

(13.1-15.8 mm), 25 2 (12.6 + 0.8 mm); hatched from sediment collected from

Gemsbokwater, series of pools in granite desert floor, Namib Desert, Namibia,

by J. A. Day, April 1978.

Branchipodopsis cf. tridens, Northern Transvaal and Zimbabwe.

AM LEN 151A, 4 ¢ (2 poor condition; others 6.9, 7.9 mm); collected from pool

on rocky outcrop near Umzingwani River, Benfer Estates, Zimbabwe, by

O. Wirminghaus, 8 July 1988. AM LEN 134B, 1 ¢ (13.4 mm), 1 ? (12.3 mm);

collected from Mopane, 35 km south of Beit Bridge, Transvaal, by O. Wir-

minghaus, 31 December 1992.

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 351

Fig. 28. Branchipodopsis tridens. A. Dorsal view of clypeus, Etosha Park (SMN 51334).

B. Cercopods of male (SAM-A7267). C. Ventral view of last abdominal segments of male

(SAM-A5980); arrow indicates spinous projection. D. Basal part of penes (SAM-A5980);

arrow indicates median rounded projection. E-G. Variation in basal processes shown in

dorsal view. E. Kalahari (HNHMI/A-78). F. Papkuil (SAM-A5980). G. Amadap Valley,

Kalahari (SAM-A7599). H. Dorsal view of clypeus of HNHMI/A-78, showing shape of

basal processes of larger specimen. Scales = 0.5 mm.

352 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 29. A, C. Branchipodopsis tridens (AM LEN 163A, outside Kalahari Gemsbok Park).

A. Dorsal view of left half of clypeus. C. Detail of left basal process. B, D. Branchi-

podopsis cf. tridens (DB764, Gemsbokwater). B. Dorsal view of left half of clypeus; white

arrow indicates indentation in median margin; black arrow indicates projection on apical

joint. D. Detail of left basal process. Scales: A-B = 0.5 mm; C-D = 100 um.

Redescription of adult male

Clypeus. Basal process broad, distally with a large, angular inner lobe and a

short, conical outer lobe. Inner lobe usually apically flat or slightly convexly

rounded (Figs 28A, 29A-D) and with scale-like patterning on apex

(Fig. 30A-C). Median margin of basal process usually with a distinct

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 353

indentation at base of inner lobe (Fig. 29A-D). Large conical/digitiform

projection present proximally on dorsal surface of each basal process (Figs 28A,

29A-D). Lamelliform process narrow and ovate (Fig. 29A-D). Apical joint

with distinct bend, and strongly curved inwards, usually apically indented or

bifid (Fig. 28A). Median ventral process small and minutely setulose but in

most cases absent.

Cercopods. Moderate length (ratio to body length approximately 0.25: 1).

Outer margins convex with short, plumose setae along three-quarters of length,

distal quarter with 4-6 spines. Proximal third of inner margins with sparse,

moderate length setae, followed by 12-16 sharp spines (Fig. 28B).

Abdominal spines. Pair of spines on last abdominal segment usually

prominent (Fig. 28C).

Penes. Basal part broad, with distinct rounded projection proximally on

median margin (Fig. 28D).

Remarks

A large amount of variation is evident in the clypeus of B. tridens from

different localities. In some cases the variation may be attributed to the speci-

mens not being fully mature or a result of poorly preserved material. The

HNHM specimens are, presumably, part of the type material. The clypeus of

even these three specimens shows variability in terms of the median margin

indentation, shape of the inner lobe (Fig. 28E, H) and apex of the apical joint,

and this is probably size related. Most of the specimens from the other localities

are, however, fully developed and in fair condition but they still exhibit a

certain amount of interpopulation variation in the shape of the lobes of the basal

process, their relative lengths, the proportion of the dorsal, conical projection to

that of the basal process and in the shape of the apex of the apical joint. The

indentation in the median margin of the basal process is also variable in extent.

In some specimens it is so pronounced as to form a narrow third lobe, whereas

other specimens have a completely straight median margin. There are a number

of sets of specimens with the median margin in a state intermediate between

these two extremes. An attempt to divide the large number of specimens into

morphological groups, based mainly on proportions of inner and outer lobes and

dorsal process, failed because of a large amount of overlap of characters.

Further comment on this intraspecific variation is made in the discussion

section.

Certain of the Namib Desert specimens (DB 761, 762, 763, 764) do, how-

ever, have characters that allow them to be distinguished from the other

B. tridens material. These specimens have a distinct, short and broad spinous

projection just proximal to the bend in the apical joint (Figs 29B, 30D). This

structure is absent in all other B. tridens specimens examined and from both

Daday (1910) and Barnard’s (1929) descriptions and illustrations. In addition,

these specimens have a deep indentation in the median margin of the basal

process, such that a narrow third lobe is formed (Fig. 32B, D), and the dorsal

projection is broad and blunt as opposed to the conical and apically narrow

process of other specimens. These differences could indicate that this material

represents a species or population intermediate between B. tridens and

B. kaokoensis.

354 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 30. Branchipodopsis tridens. A. Detail of apex of inner lobe of left basal process

(Namib Naukluft, SMN51197). B. Detail of apex of inner lobe of right basal process

(AM LEN 163A). C. Detail of apex of inner lobe, left basal process of Goru Pan specimen

(TM VLKE443). D. Projection on apical joint of Gemsbokwater specimen (DB764).

Scales = 50 pm.

The Zimbabwe and Mopane specimens show the greatest difference from

Daday’s (1910) figures and specimens, and are easily distinguished from the

remainder of the B. tridens material. The inner lobe is broadly hook-shaped

with a convex median margin and the apex of the apical joint is tapered and

narrow (Fig. 31A-B). For this reason, and because identical specimens have

been collected from two localities, it is quite possible that they represent a

closely related, but separate species from B. tridens. However, no material has

been collected from the area between Namibia and northern Transvaal—southern

Zimbabwe and the possibility of intermediate forms exists.

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 355

Fig. 31. Branchipodopsis cf. tridens (AMLEN 151A). A. Dorsal view of left half of cly-

peus, Zimbabwe specimens. B. Detail of basal processes. Scales = 100 ym.

Until such time as collecting over greater areas of southern Africa has been

done and further research into sexual isolation and molecular biology has been

undertaken, it seems prudent to simply describe the variation observed without

drawing any conclusions.

Distribution

Branchipodopsis tridens is widely distributed in the arid western region of

southern Africa. A large number of records from northern Namibia exist, and

this species has also been collected from the southern part of that country as

well as the north-western Cape (Fig. 39). The Zimbabwe specimens represent

the most easterly record of the species and B. tridens has also been collected

from Bloemfontein in the central region of South Africa (Seaman ef al. 1991).

Branchipodopsis underbergensis sp. nov.

Figs 32-33

Material

Holotype. SAM-A40837, 1 3 (14.4 mm); collected from rock pool on

Bamboo Mountain, near Underberg, Drakensberg, Natal, by T. Clarke &

N. Crouch, December 1993.

Paratypes. SAM-A40838, 15 6 (14.3 + 0.6 mm), 8 2 (12.0-15.0 mm);

same collection data as holotype. .

Other material examined. SAM-A40848, 24 6 (10.7 + 0.6 mm), 29 2? (10.1

+ 0.7 mm); collected from Underberg, Drakensberg, Natal, collector and date

unknown (specimens were previously in the teaching collection of the Zoology

Department, University of Natal, Pietermaritzburg).

356 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description of male

Clypeus. Basal process positioned low on basal joint (apex not reaching past

anterior median margin of basal joints) on inflated base, slender and digitiform

(Figs 32A, 33A-B). Prominent short and conical projection on proximal, dorsal

surface of basal process (Figs 32A-B, 33A-B). Apical region of basal process

and projection patterned by series of rounded ridges and depressions (Fig. 33C).

Median ventral process ovate and spinous with blunt apex (Figs 32A, 33A).

Lamelliform process very large and round-ovate (Figs 32A, 33A). Apical joint

curved inwards and with inflated apex (Figs 32A, 33A).

Cercopods. Long (ratio to body length approximately 0.35 : 1). Outer mar-

gins proximally convex, distal halves slightly concave and with short, spinous

setae along entire length. Small patch of setae proximally on inner margins

followed by about 14 short spines (Fig. 32C).

Abdominal spines. Pair of small spines on ventral surface of last abdominal

segment (Fig. 32D).

Penes. Basal part short, with moderate-sized, rounded projection on median

margin (Fig. 32E).

Description of female

Pair of small pouches on dorsal surface of penultimate thoracic segment and

lateral surfaces of last thoracic segment extended outwards (Fig. 32F).

Egg morphology

Egg surface with irregular, four- to five-sided depressions, separated by

narrow, sharp ridges (Fig. 33D).

Differential diagnosis

Superficially, B. underbergensis resembles B. drakensbergensis. This is

largely because of the proximal projection of the basal process and the slender

shape of the basal process. However, the fact that the basal process of the

former species is single as opposed to bilobed in the latter, suggests two distinct

species. Further differences include the very large lamelliform process in

B. underbergensis and the small basal process positioned low on the basal seg-

ment of the clypeus of this species. The median ventral process also differs in

shape, as do the cercopods. In addition, the females of B. drakensbergensis lack

the thoracic pouches (Fig. i0F) that are distinct in B. underbergensis females.

Distribution

Branchipodopsis underbergensis has only been collected from the Under-

berg area of the Natal Drakensberg (Fig. 38).

Remarks

In spite of the general similarity between the basal processes of B. drakens-

bergensis and B. underbergensis, these two species were not linked in the

cladistic analyses (Fig. 41). Branchipodopsis underbergensis was more fre-

quently grouped close to B. hodgsoni. This may be possible since the dorsal

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 357

Fig. 32. Branchipodopsis underbergensis sp. nov. (SAM-A40838, paratype). A. Dorsal

view of clypeus. B. Lateral view of clypeus; e—eye; s—spinous projection of basal process;

Ip—lamelliform process; 1—labrum. C. Cercopods of male. D. Ventral view of last

abdominal segments of male; arrow indicates spinous projection. E. Ventral view of basal

part of penes. F. Dorsal view of genital segments of female; small arrow indicates dorsal

extensions; large arrow indicates posterior lateral extensions. Scales = 0.5 mm.

process on the basal process of B. underbergensis could be homologous to the

dorsal lobe of the basal process of B. hodgsoni rather than the spinous process

in B. tridens, B. kaokoensis and B. drakensbergensis. The low position of the

basal process in B. underbergensis and in B. hodgsoni could provide further

evidence for the relationship between these two species.

358 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 33. Branchipodopsis underbergensis sp. nov. (SAM-A40838, paratype). A. Dorsal

view of left part of clypeus. B. Detail of left basal process, arrow indicates spinous

projection. C. Detail of apical region of basal process lobe. D. Egg.

Scales: A = 0.5 mm; B, D = 100 um; C = 10 pm.

Etymology

Branchipodopsis underbergensis is named after the type locality,

Underberg.

Branchipodopsis wolfi Daday, 1910

Figs 34-37

Branchipodopsis wolfi Daday, 1910: 304, fig. 52; 1913: 4. Barnard, 1924: 217; 1929: 197,

fig. 5g.

al i

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 359

Material

Type material. Collected in the Kalahari and housed in the Senckenberg

Museum. No further information available.

Material examined. SAM-A7268, 1 3 (11.9 mm), 1 2 (10.3 mm); collected

from Altmark, Outjo district, Namibia, by K. H. Barnard, January 1926. SAM-

A6004, 18 ¢ (6.5 + 0.5 mm), 15 2 (6.5 + 0.4 mm); collected from Onganjera,

Ovamboland, Namibia, by K. H. Barnard, 1926. SAM-A5920, 5 6 (11.9-

15.6 mm); collected from Beaufort West, Karoo, Cape Province, by W. F. Pur-

cell, 24 September 1905. SAM-A7253, 20 3 (6.3 + 0.6 mm), 11 2 (5.8 +

0.5 mm); collected from Kamanyab, Kaokoveld, Namibia, by K. H. Barnard,

1926. SAM-A7273, 8 5 (10.0-12.0 mm), 9 2 (10.0-11.5 mm); collected south-

east of Choabendus, Kaokoveld, Namibia, by K. H. Barnard, January 1926.

SAM-A5921, 19 3 (12.8 + 1.5 mm), 13 2 (12.1 + 0.9 mm); collected from

Great Fish River, near Gibeon, Namibia, by R. W. Tucker, January 1916.

SAM-A4248, large number of specimens, 23-¢ measured (7.8 + 0.8 mm),

15 2 measured (7.9 + 0.8 mm); collected from Kimberley, Cape Province, by

J. H. Power, 1917. SAM-A7254, 9 ¢ (12.8-14.8 mm), 1 2 (15.0 mm); col-

lected from Kimberley, Cape Province, by J. H. Powers, 1917. SAM-A6005,

5 ¢ (8.5-10.5 mm), 21 2 (8.6 + 0.7 mm); collected from Waterberg, Namibia,

by R. W. Tucker, 1920. SMN 51217, 5 ¢ (8.0-9.1 mm), 6 2? (7.8-8.8 mm);

collected from rain pool near Otjituuo, Hereroland West, Namibia, by B. A.

Curtis, 28 April 1987. HNHM I/A-77, 1 6 (8.9 mm), 1 2 (8.2 mm); collected

from Kalahari, Botswana, collector and date unknown. AM LEN 99B, 4 ¢

(14.8-15.3 mm), 8 2 (9.5-11.3 mm); collected from Pumbe Pan, Kruger

National Park, Transvaal, by M. Hamer, 28 October 1990. AM LEN 154A, 3 6

(8.1, 9.6, 10.0 mm), 5 2 (8.9-9.5 mm); collected from Nylsvlei Nature

Reserve, Transvaal, by C. Pitzke-Widdig & T. Widdig, February 1990.

DB 668, 1 ¢ (12.5 mm), 5 2 (10.0-13.0 mm); collected from Machoarane

Mountain, about 2 km south of Morija (Moraija), Lesotho, by T. C. Ruther-

ford, 8 November 1985. DB 667, 1 ¢ (abdomen removed), 2 2? (10.1,

12.8 mm); collected from Brakfontein, 17 km south of Mohales Hoek, Lesotho,

by T. C. Rutherford, 12 December 1985. DB 669, 8 ¢ (8.1-13.9 mm), 7 2

(11.3-12.8 mm); collected from Leloaleng near Quting, Lesotho, by T. C.

Rutherford, 13 December 1985. DB 817, 3 ¢ (14.0, 16.4, 17.0 mm), 3 2 (13.0,

13.9, 15.6 mm); collected from Outward Bound, Leribe, Lesotho, by T. C.

Rutherford, 4 January 1986.

Branchipodopsis cf. wolfi, Transvaal. AM LEN 69B, 3 ¢ (8.5, 8.8,

9.0 mm), 2 2 (8.0, 9.8 mm); collected from Mala Mala Game Reserve, Trans-

vaal, by W. A. Taylor, 25 June 1989. AMLEN 155A, 3 6 (9.9, 11.1,

11.3 mm); cultured from dried sediment (February 1990) collected from Mala

Mala Game Reserve, Kirkmans Kamp area, Transvaal, by M. Hamer, Septem-

ber 1989. AMLEN 120A, 7 ¢ (7.8-10.0 mm), 2 2 (6.8, 9.5 mm); collected

from region south of Skukuza, Kruger National Park, Transvaal, by A. & J.

Bowland, 14 October 1990. AM LEN 121B, 15 6 (9.8 + 0.6 mm), 22 2 (8.7 +

0.5 mm); same collection data as AM LEN 120A.

Branchipodopsis cf. wolfi, Zimbabwe. NMZ/Cr 29, 1 & (damaged), 3 2

(damaged); collected from Matopos, Zimbabwe, December 1964, collector

360 ANNALS OF THE SOUTH AFRICAN MUSEUM

unknown. NMZ/Cr 6, 9 ¢o (15.0-20.6 mm), 14 2 (17.9 + 2.0 mm); collected

from Matopos, Zimbabwe, by A. N. Smith, December 1981. USNM 184629,

3 6 (9.0, 9.3, 9.4 mm), 2 ? (8.1, 8.6 mm); collected from Bulawayo,

Zimbabwe, by D. H. Eccles & W. G. Moore, 10 February 1955.

USNM 102318, 3 ¢ (8.8, 9.0, 10.6 mm), 1 2 (9.5 mm); collected from pools at

isoyds of the tomb of Cecil John Rhodes, 25 miles south-south-west of

Bulawayo, Zimbabwe, by D. H. Eccles, 10 February 1955.

Redescription of adult male

Clypeus. Basal process wide with single large, conical or triangular lobe

positioned on inner side of basal process and two apical tubercles (Figs 34A,

35A-D). Lateral tubercle digitiform/conical and positioned slightly ventrally

(Fig. 36A-D). Median tubercle usually similar in size and shape to outer

Fig. 34. Branchipodopsis wolfi. A. Dorsal view of clypeus of specimen HNHM I/A-77

from Daday’s collection. B. Cercopods of male (AMLEN99B). C. Ventral view of last

abdominal segments of male (AM LEN 99B) showing spinous processes (indicated by

arrows). Di. Dorsal view of basal part of right penis (SAM-A5921); arrow indicates blunt

projection. Dii. Ventral view of apex of basal part of right penis (SAM-A5921); arrow

indicates blunt projection on lateral surface.

Scales = 0.5 mm.

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 361

Fig. 35. Branchipodopsis wolfi. A. Left half of clypeus, Lesotho specimen (DB 669).

B. Left half of clypeus of specimen from Pumbe, Kruger National Park (AM LEN 99B).

C. Left half of clypeus of Great Fish River specimen (SAM-A5921). D. Clypeus of

Altmark specimen (SAM-A7268). Scales = 0.5 mm.

tubercle and situated dorsally at base of inner lobe (Figs 36A-D, 37B-C, F). A

small, flattened projection often present on median apex of lobe (Figs 36A, C,

37D, F). Lamelliform process small and round or oval (Figs 34A, 35A-D).

Apical joint curved inwards with apex varying from flattened (Fig. 35B) to

slightly indented (Fig. 35D). Median ventral process very small to large and

prominent, usually oval and with surface covered by short spines (Fig. 35A-D).

362 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 36. Branchipodopsis wolfi. A. Detail of left basal process of DB 669 specimen;

il—inner lobe; It—lateral tubercle; mt—median tubercle; pr—projection on inner lobe.

B. Detail of left basal process of AMLEN99B. C. Detail of left basal process of

SAM-A5921. D. Detail of right basal process of SAM-A7268 (arrow indicates lateral

tubercle). Scales = 100 um.

Cercopods. Average length (ratio to body length approximately 0.27: 1).

Outer margins weakly convex and with plumose setae of moderate length but

these replaced by 2-4 short spines distally. Proximal third of inner margins with

plumose setae followed by 10-16 spines of unequal size (Fig. 34B).

Abdominal spines. Last abdominal segment with two strong ventral spines

(Fig. 34C).

Penes. Basal part slender, with proximal rounded projection on median

margin. Peg-like projection near apex on lateral margin (Fig. 34D-E).

Remarks

Much variation is evident in the clypeus of B. wolfi from different localities.

This was observed and commented on by Barnard (1929). Variation is most

obvious in the following features of the clypeus: the apex of the apical joint, the

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA

Fig. 37. Branchipodopsis cf. wolfi. A. Dorsal view of clypeus of southern Kruger

National Park specimen (AMLEN 121B). B. Left basal process of AM LEN 121B.

363

C. Right basal process of Mala Mala specimen (AMLEN 155A); outer tubercle.

indicated by arrow. D. Detail of apical region of basal process (AM LEN 121B);

il—inner lobe; mt—median tubercle; pr—projection on inner lobe. E. Basal

processes, Zimbabwe specimen (NMZ/Cr6). F. Detail of apical region of right

basal process of NMZ/Cr6. Scales: A-B, E = 0.5 mm; C, F = 100 um;

D = 50 pm.

364 ANNALS OF THE SOUTH AFRICAN MUSEUM

shape of the tubercles and the inner lobe, the presence and shape of a projection

on the inner lobe, the relative lengths of the tubercles, the position and size of

the median tubercle, and the shape of the median margin of the basal process.

The HNHM specimens are possibly the types, as they are from Daday’s

collection and from the same locality as the types. In these specimens, the

tubercles and inner lobe are rather short and rounded and all of similar shape. A

small apical projection is present on the inner lobe. The median margin has an

inflation just proximal to the base of the inner lobe. Most of the other specimens

show some variation on this basic arrangement. The median tubercle of the

Lesotho specimens is small and lies in the same plane as the inner lobe and the

apical process is very prominent and rounded (Fig. 36A). In the Altmark speci-

men, the inner lobe is roughly triangular and lacks the apical projection

(Fig. 36D). The Kruger Park (Pumbe) specimens have an angular median mar-

gin and a very broad inner lobe (Fig. 36B). It is, however, the Zimbabwean and

southern Kruger Park specimens that show the greatest morphological diver-

gence from the ‘types’ of B. wolfi. The median tubercle is small and triangular

and is situated about halfway along the length of the inner lobe (Fig. 37B, F).

These Zimbabwean and southern Kruger Park specimens could be regarded as a

separate species but have been included with B. wol/fi because there is a series of

specimens in which the basal process morphologies are intermediate between

these and the other, more typical representatives. For example, the Mala Mala

specimens have a larger median tubercle than the other two sets of specimens,

those from the Kruger Park have a slightly smaller tubercle, and the Zimbabwe

specimens have an even smaller one. These specimens, therefore, form a series

from a prominent to a small median tubercle and yet all three sets share a very

similar basal process shape. The specimens from Nylsvlei have a basal process

morphology intermediate between the Kruger Park, Mala Mala and Zimbabwe

specimens and other, more typical, B. wolfi specimens, and this makes it

impossible to divide, with any confidence, these specimens into separate species

or subspecies at this stage.

The structure of the basal process of B. wo/fi can be interpreted in a number

of different ways and this further complicates any attempt to identify species or

subspecies, and to determine relationships with other species of the genus. It is

uncertain whether the median tubercle illustrated in Figure 36B and D is

homologous to that illustrated in Figures 36A and 37B and F. In addition, the

inner lobe could be interpreted as a third tubercle and the lateral tubercle could

represent a second lobe. The latter interpretation was included in one of the

cladistic analyses but did not result in B. wo/fi being grouped differently to those

patterns illustrated in Figure 41.

Distribution

As Barnard (1929) stated, B. wolfi is the most widely distributed species of

Branchipodopsis. This species occurs in arid northern and southern Namibia,

the northern Cape and Karoo, in the sub-tropical eastern Transvaal and

Zimbabwe, and in the high-altitude, mountainous habitats of eastern Lesotho

(Fig. 40). Branchipodopsis wolfi was also collected from Umfolozi Game

Reserve in north-eastern Natal, but these specimens had dried out and they were

not listed under specimens examined. An additional set of specimens from pools

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 365

outside Gaberone was examined and identified as B. wolfi for Dr M. Cantrell,

formerly of the University of Botswana. Barnard (1929) referred to the presence

of B. wolfi in Kenya but illustrations or specimens from there have not been

seen and so this locality cannot be verified. A set of specimens in the BMNH

from ‘40 km north of Soroti in the eastern province of Uganda’ were examined

and their identification as B. wolfi confirmed in terms of the features of the

species as presented here. This record may indicate a wider distribution of

B. wolfi than for the other southern African Branchipodopsis species.

KEY TO THE SOUTHERN AFRICAN SPECIES OF THE GENUS BRANCHIPODOPSIS

(using features of the male clypeus)

1A. Basal process distinctly bilobed apically, lobes of similar height and

positionedsadjacent:toreachiothen..c-cc ees cns cee orescence waceete eneeese se 2

1B. Basal process with one to three lobes; if bilobed, lobes of distinctly

different heights or not positioned adjacent to each other ................... 5

2A. Basal process with tubercle on median dorsal surface and smaller projec-

fHOnTON Median marcim OP NNeEMlODE. osc. 0h ceceee ce aeee tees acer eee 3

2B. Basal process with small projection on median surface of inner lobe only

Cc Og SENOS aUBES ASU EE GAS SO SRO RE NEET A ee nthHcoeiiNn sina Ieelne ral ine Ra nOADAnirua Why a: ere 4

3A. Basal process lobes equal in size and shape, apex of apical joint blunt,

lamelliform processes broadly oval, with scalloped margin ..................

CBSE BETS aT SEL eae thsi B. hutchinsoni sp. nov. (Figs 17-18)

3B. Inner lobe of basal process narrower than outer, apex of apical joint

peg-like or rounded, lamelliform processes narrow oval with smooth

PULA PUT Se cies sista ssiecrs ease B. natalensis Barnard, 1929 (Figs 23-24)

4A. Inner lobe of basal process more conical than rounded, lamelliform

process small, distal region of apical joint foot-shaped ...................066+

Bee eee toes valoda ade manent eaieeuacee se DeNecme ae aioe B. dayae sp. nov. (Figs 8-9)

4B. Both lobes of basal process rounded, lamelliform process distinct, distal

region of apical joint with slight inflation and with blunt tip .................

Re eee Nee ace Tee aa nee haenaer B. karroensis Barnard, 1929 (Fig. 22)

5A. Basal process with large, spine-like or conical projection proximally on

GOESAMESUELACE Hi ieee cots o Saar taco eaiaceone Sane ou connie enous ea nec aa» 6

5B. Basal process without proximal spine-like or conical projection .......... 9

6A. Basal process with three distinct lobes, inner and outer lobes similar in

Sizeraridesmallerthanimedianlobeyn aes nee es see c cece ce es ee

BERR ARC ORE EE EEE DE RUE ERRES orate eTs B. kaokoensis Barnard, 1929 (Fig. oe

6B. Basal process with only one or two distinct lobes ..................seeeeeeeee

7A. Basal process with one or two lobes, narrower than high, median ventral

PEOCESSPLOMIMC te era risk of ks eee tease bon nats duce nee ottateltuansimmeminua ae er auioias 8

7B. Basal process wider than high, median ventral process vestigial or absent

SN teh iat Mhe ters eA ara x oidieetal B. tridens Daday, 1910 (Figs 28-31)

366

8A.

8B.

9A.

OB.

10A.

10B.

11A.

11B.

12A.

12B.

13A.

13B.

14A.

14B.

ISA.

ISB.

ANNALS OF THE SOUTH AFRICAN MUSEUM

Basal process bilobed and almost as high as division between basal and

apical joints, lamelliform process smaller than basal process ................

SGLGD ICOSH AS ACRES LAMM aan Nes B. drakensbergensis sp. nov. (Figs 10-13)

Basal process with single lobe and lower than anterior median margin of

basal joint, lamelliform process very large (larger than basal process) ....

Matinee na nee aes 5 oi elle A Seah Tomine B. underbergensis sp. nov. (Figs 32-33)

Basal process with single lobe, dorsal or apical tubercles or projections

PRESENE voici deleieviecieanisincisrneidind as szelsecaisioaya diineiatols ace yale alee ee TEC AE ARERR EERE 10

Basal process single or bilobed, if bilobed, lobes of distinctly different

heights, dorsal or apical projections or tubercles absent .................. Wg

Basal process with inflated base, rounded apex and with keel-like

Structuredorsallly <2. 4... cee. ceescceeee B. simplex Barnard, 1924 (Fig. 27)

Basal process without keel-like structure dorsally, but with one or more

tubercles or projections Near apeX ...2.....5...+0cces0 sence eens cee eRe 11

Basal process with two digitiform or conical tubercles apically, projecting

beyond anterior margin of basal process, lamelliform processes small or

MOGEFAtE-SIZER! sci sced ea dos. ducheeteic ewes atonsirnc ee aes een eR eeE eee 12

Basal process without apical tubercles projecting beyond anterior margin,

lamelliform processes enlarged ...... B. browni Barnard, 1924 (Figs 6-7)

Two tubercles present, second positioned ventrally to first, distinct inner

lobejabsent iseesc ie eelncnt Sosa ete ees B. hodgsoni Sars, 1898 (Figs 15-16)

Two tubercles present, one median and one ventrolateral, large inner lobe

PRCSEM teases yensenstes os sees cure pose ctneenee B. wolfi Daday, 1910 (Figs 34-37)

Basal process with single lobe of spiniform shape ......................eeeeeee

Lie Oe NRC eh Ai SBN IN ede tan B. scambus Barnard, 1929 (Figs 25-26)

Basal process bilobed, outer lobe distinctly shorter than inner lobe .... 14

Width of basal process greater than or equal to height, apical joint stout

and strongly inwardly curved with distinct bend ....................sceeeeeeees

Bi hic PCIE CCR ne RG ENG B. kalaharensis Daday, 1910 (Figs 19-20)

Basal process longer than wide, apical joint slender, weakly curved and

Without distinct DENG’... 25.05. ceccséacs Ses0cses sooeee se ceee saccuneenee Ree Eeeeeeeee 15

Apical joint distally hook-shaped, median ventral process present .........

RRA ery a SouNle ry ec eas Went a AEE ae bak lye ta B. drepane Barnard, 1929 (Fig. 14)

Apical joint distally smooth and straight, median ventral process absent ..

SOE ORO NORE BO cen Sonat mr aur B. barnardi sp. nov. (Figs 4-5)

DISCUSSION

Of the approximately 21 known species of Branchipodopsis, 16, including

the five new species described here, occur in southern Africa and 15 of these

are apparently restricted to this region. There is little morphological variety

among the three Asian Branchipodopsis species and it has even been suggested

that they may belong to a single species (Hartland-Rowe 1968; Brtek et al.

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 367

1984). The status and morphology of the two remaining Branchipodopsis

species, namely B. abiadi from Mauritania in West Africa and B. candea from

central Africa, are uncertain since these specimens are unavailable, and either

the descriptions are inadequate or, in the case of the latter species, the original

description cannot be located.

Taxonomic characters

The second antennae of the male, as in most other anostracan genera, is the

character of greatest taxonomic value within the genus Branchipodopsis. The

male anostracans use the enlarged, two-jointed antennae to clasp the female

during mating. Belk (1991) found that, within the Anostraca, mating depends on

female choice, with males attempting to mate completely unselectively. The

female does not, however, always successfully recognize conspecific males by

the structure of the antenna or by body armature or cercopod structure. This

statement was based on Wiman’s (1979) findings that hybridization occurred

between North American species of Streptocephalus both in the laboratory and,

under sympatric conditions, in the field. Selection, therefore, is not for anti-

hybridization mechanisms. Belk (1991) suggested that the second antennae,

Fig. 38. Map of southern Africa illustrating the distribution of Branchipodopsis browni

(open circle), B. drepane (open star), B. hodgsoni (open square), B. kalaharensis (closed

triangle), B. kaokoensis (closed square), B. karroensis (closed circle), B. natalensis (flower),

B. simplex (open, inverted triangle) and B. scambus (closed star).

368 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 39. Map of southern Africa illustrating the distribution of Branchipodopsis tridens

(closed circle), B. hutchinsoni sp. nov. (closed star), B. drakensbergensis sp. nov. (open

square) and B. barnardi (flower).

because of their function in mating, are ideal candidates for runaway sexual

selection by female choice and their ‘elaborate structure should be viewed as

arbitrary additions to the complexity of structures providing tactile clues’.

Speciation in anostracans appears to be the result of the slow accumulation of

genetic differences in allopatric populations (Wiman 1979).

In contrast to the streptocephalids, in which the male cercopod setation,

frontal appendage, abdominal armature and external egg morphology are fre-

quently species-specific, these characters either vary little between species of

Branchipodopsis or they are largely absent. Authors such as Linder (1941)

emphasized the structure of the penes as taxonomic characters at the familial

and generic levels, but between species these do not differ to any extent. There

is some interspecific variation in the prominence of the rounded projection on

the median margin of the penes in the Branchipodopsis species examined but

this is difficult to quantify and is often dependent on the preservation method or

condition of the specimens. The peg-like projection at the apex of the proximal

part of the penes is very obvious in some species, whereas in others only a blunt

obscure projection is present. In other species, no distal lateral projection of any

kind is visible. In certain cases, preservation state is responsible for the varia-

tion but this structure is quite clearly absent in some species. Unfortunately,

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 369

these structures do not appear to reflect relationships between species. Within

Branchipodopsis there is little interspecific variation in cercopod shape or

setation; the number of spines on the inner margin does vary to some extent

between species but there is a significant amount of overlap and intraspecific

variation. External egg morphology is also of limited value for the same reasons

(pers. obs.). Body armature, such as that seen in certain Streptocephalus

species, is absent in Branchipodopsis with the exception of the two spines on the

ventral surface of the last abdominal segment. However, this character does not

appear to reflect species relationships. It is often present in species with very

different antennal morphologies and, since these spines are present in

approximately half the described species, they cannot be used for species

identification. A character not mentioned by other authors but used in this

review is the presence, position and shape of pouches proximal to the genital

segments of the females of some species. The significance of these structures is

uncertain but, since they only occur in high-altitude species (B. barnardi,

B. underbergensis and B. natalensis), they may be of physiological importance.

Until proved otherwise, these structures provide a useful additional character in

separating species with similar basal process morphologies, such as B. natal-

ensis and B. hutchinsoni, and B. drakensbergensis and B. underbergensis.

32°

igus .

5 34°

i NES jue [

E 12° 16° 20° 24° 28° 32° 36°

Fig. 40. Map of southern Africa illustrating the distribution of Branchipodopsis wolfi (closed

square), B. underbergensis sp. nov. (open triangle) and B. dayae sp. nov. (open circle).

370 ANNALS OF THE SOUTH AFRICAN MUSEUM

Species groups and systematic analyses

An attempt was made to divide the southern African species of Branchipod-

opsis into species groups. This was recently done for the African strepto-

cephalids (Hamer et al. 1994a, 1994b) as a step towards understanding relation-

ships between species and in examining the biogeography of the genus Strepto-

cephalus. For the southern African species of Branchipodopsis, a combination

of cladograms was used to achieve this division. The cladistic analyses and the

groupings resulting from them must be seen as tentative, since inadequate data

are available on the morphology, ontogeny and evolution of the genus and

family. For example, it is almost impossible to determine, with any degree of

confidence, homologous structures, incidences of homoplasy or to be certain of

the pleisiomorphic and apomorphic states. Outgroup selection is also problem-

atic since the characters and members of the Branchipodidae require total

revision. The limited number of characters (Table 1) and, in certain species,

specimens, that can be used, is a further limitation to these analyses. The

relatively low RI and CI indices obtained from the Hennig86 analyses indicates

a large amount of homoplasy and a low confidence level in the trees produced.

Changing the interpretation of some characters and excluding others did not

improve these values nor did it produce groupings notably different from the

two illustrated in Figure 41. This could indicate that the species groups have

some credibility.

TABLE |

Tentative ordering of the southern African Branchipodopsis into species groups based on

cladistic and phenetic analyses.

Group Characters Species

1 Basal processes simple, spine-like, apical joint weakly = B. scambus

curved.

2 Basal processes with dorsal and ventral lobe, and set B. hodgsoni

low on basal joints B. underbergensis

3 Basal processes with single lobe, small projection on B. browni

median, apical margin, second projection/tubercle on B. wolfi

dorsal median surface

4 Basal processes with two lobes, almost equal in size, B. natalensis

one or more projections on basal processes. B. hutchinsoni

B. karroensis

B. dayae

5 Basal processes narrow, bilobed, with smaller outer B. drepane

lobe than inner B. barnardi

6 Basal processes broad, apical joints strongly bent, B. kalaharensis

median ventral process absent : B. simplex

7 Basal processes with dorsal spinous projection, 2 or B. kaokoensis

3 unequal-sized lobes. B. tridens

B. drakensbergensis

371

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA

aehep ‘g

sisuaosey ‘g

juosulya}ny “g

sisuajejeu “g

LJOM “@

jumosg “g

juosbpoy ‘g

sisuabsaquapun ‘g

sisuaoyoey ‘g

suapu} ‘g

xajduis ‘g

sisuaseyejey “g

Jpseuseg ‘g

sisuabsaqsuayesp °g

auedasp ‘g

snquweogs ‘g

eIWapsesed

sisuaoyoey “g

suapi} ‘g

xajduis ‘g

sisuaseyeey “g

Ipseuseg ‘g

auedasp “g

sisuabsaqsuayesp “g

aehep ‘g

sisuaouey ‘g

juosulyayny “g

sisuajejeu ‘g

jumolq ‘g

HOM “g

juosbpoy ‘g

sisuabsaqiapun ‘gd

snquegs ‘g

e/WUajIesed

Cladograms of the southern African species of the genus Branchipodopsis.

A produced from the command ‘mhennig’, with length of 35, ci value of 42, and ri of 66.

Fig. 41.

B produced by the command ‘ie-’, with length of 45, ci of 42 and ri of 62.

372 ANNALS OF THE SOUTH AFRICAN MUSEUM

Intraspecific variation

The large amount of variation observed in the basal processes of B. wolfi

and B. tridens presented a major problem in this study. Specimens of these two

species from some localities differed considerably from the original description

and the type material in terms of antennal and, particularly, basal process mor-

phology. However, other sets of specimens had basal process morphologies

intermediate between these and the original material. Attempts were made to

group populations of B. tridens or B. wolfi with the most similar basal process

structure morphometrically. However, these results indicated a large degree of

overlap and no distinct clusters, which would have been the case if different

species were involved. Thus, the variation was simply described and/or illus-

trated. In the case of B. wolfi, antennal variation was evident in almost every

population and it was impossible to illustrate all of this. Similar intraspecific

variation was also observed and illustrated in the Asian species B. affinis (Brtek

et al. 1984). In the case of these three species, more advanced taxonomic

methods, most important of which will certainly be molecular techniques, are

necessary to determine the relevance of this variation. At this stage, however,

lack of sufficient quantities of material of all representative populations presents

the major obstruction to such a study.

Distribution and ecology

An in-depth discussion of the biogeography of Branchipodopsis will be

published elsewhere and only a brief comment will be made here about distribu-

tion and habitats. The majority of Branchipodopsis specimens were collected

from small waterbodies, such as roadside ditches, shallow rain pools and small

rock pools, where conditions would be expected to be extreme. Observations

under both laboratory and field conditions have revealed that a number of

Branchipodopsis species (B. tridens, B. wolfi, B. dayae and B. browni) grow

very rapidly and reach sexual maturity 4-6 days after hatching. Their life span

is only 2-4 weeks. This contracted life history allows members of the genus to

exploit habitats not available to the other main group of southern African

anostracans, the streptocephalids, which generally have a slower, longer life

history. Species of the genus Streptocephalus have never been collected from

high-altitude rock pools, thus allowing Branchipodopsis to dominate such

habitats. Belk (1991) found a similar situation with §. texanus and Branchinecta

packardi in central Texas. In temporary pools in southern Africa where

Streptocephalus occurs, Branchipodopsis species are either absent, or are found

in low numbers and complete their life history before the streptocephalids reach

maturity, or they occupy a different niche. This may imply a reduced competi-

tive ability for the genus. Further evidence for this is the rarity of the presence

of more than one species in a single habitat. Barnard (1929) reported that two

male B. drepane were found among a sample of B. tridens and B. wolfi but this

is the only known case of multispecies existence. This could be a result of the

restricted resources in habitats such as montane rock pools; but the forces

separating species such as B. barnardi and B. natalensis, which inhabit the

same type of rock pools in the Drakensberg in an area of less than 5 km?, are

not known. Perhaps, as has been suggested (Wiman 1979; Williams 1985;

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 373

Brendonck ef al. 1990), branchiopods are adapted to resisting, rather than pro-

moting, dispersal, which may be a rare occurrence.

Williams & Busby (1991) suggested that only in episodically filled tempor-

ary pools could one expect to find widespread and easily dispersed species. The

reason for this is that the irregularity of filling of such habitats does not allow

the evolution of local adaptations and subsequent speciation. Many Branchi-

podopsis habitats, however, because of their small size, are likely to undergo a

number of wet and dry phases during a single rainy season. As a result, many

Branchipodopsis generations could be expected to hatch, as opposed to a single

generation in large pools that remain full for the duration of the rainy season, or

those that only fill every decade or less often. This would allow relatively rapid

differentiation and could account for the large amount of intraspecific variation

in species such as B. wolfi and B. tridens.

Conclusions

Southern Africa has the greatest diversity of species of Branchipodopsis. As

suggested by Banarescu (1990), this may be as a result of the genus having a

Gondwanaland origin, or the result of a wide adaptive radiation having occurred

in response to the habitat diversity present in southern Africa. However, of the

16 species now described, four species (B. drepane, B. kaokoensis, B. karro-

ensis and B. simplex) are known only from a single collection (the type material)

consisting of fewer than 5 male specimens. For an additional species

(B. hutchinsoni), only the type material has been collected, although this is in

larger quantities than for the former species. Most species appear to have

restricted distributions and it is possible that some of the species that were last

collected early this century have become extinct. Further collecting will

certainly reveal a number of undescribed species, as will research into

intraspecific variation using molecular techniques to examine mitochondrial

DNA. The distribution of species also needs to be reassessed as previously

unexplored areas, such as the northern and western Transvaal, Zimbabwe,

Botswana and Mozambique, are sampled. Detailed investigation of species

relationships and of the position of the genus within the Branchipodidae needs to

be undertaken to answer the taxonomic questions raised here.

ACKNOWLEDGEMENTS

The senior author was in receipt of a Foundation for Research Development

(FRD) bursary. The directors and curators of the museums from which material

was loaned, in particular Dr F. de Moor and Mrs H. James of the AM, Mrs

M. van der Merwe of the SAM, Ms B. Curtis of the SMN, Dr L. Forr6 of the

HNHM, Ms S. Halsey and Dr G. Boxshall of the BMNH and Dr T. Bowman of

the USNM, are thanked for their co-operation. Dr J. A. Day of the University

of Cape Town, and Dr D. Belk of Our Lady of the Lake University, San

Antonio, provided material collected in Namibia, the Cape and Lesotho. The

National and Natal Parks Boards gave permission to collect in areas under their

control and field staff gave necessary advice and assistance on a number of

occasions. The staff of the Electron Microscope Unit of the University of Natal,

374 ANNALS OF THE SOUTH AFRICAN MUSEUM

Pietermaritzburg, are thanked for their assistance with electron microscopy,

particularly Mrs P. Donnelly who printed the micrographs. Dr J. Brtek kindly

provided valuable literature, references and a species list. Prof. G. Mura and

Drs D. Belk and B. Cook made valuable comments on the manuscript. Mr

O. Wirminghaus, Dr N. Rayner and Mr R. Struckmeyer are thanked for

various forms of assistance rendered during this study.

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BARNARD, K. H. 1935. Scientific results of the Vernay-Lang Kalahari Expedition,

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BREHM, V. 1958. Crustacea: Phyllopoda und Copepoda Calanoida. Jn: HANSTROM, B.,

BRINCK, P. & RUDERBECK, G. eds. South African animal life 5: 10-39.

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APPENDIX 1: GAZETEER

Localities from which southern African species of the genus Branchipodopsis

were recorded.

BOTSWANA

Gaberone 24°42’S 25°54’E

Gori (Goru?) Pan 18°S 22°E

Kalahari 20°S 24°E

Kanke Pan, 90 miles west of Molepole 24°24’S 25°32’E

Sunnyside 21°40’°S 22°03’°E

376 ANNALS OF THE SOUTH AFRICAN MUSEUM

LESOTHO

Brakfontein, 17 km south of Mohale’s Hoek

Leloaleng, near Quting

Machoarane Mountain, + 2 km south of Morija (Moraija)

Outward Bound, Leribe

Sehlabatebe National Park

NAMIBIA

Amichab, 140 km east of Rheboth

Etosha Park

Etosha Park, Charl Marais (Sukses) Dam

Etosha Park, Leeubron

Great Fish River, near Gibeon

Great Namaqualand, Kalkfontein South (Karasburg)

Hereroland West, near Otjituuo

Kaokoveld, Choabendus, 115 miles north-west of Outjo

Kaokoveld, Cauas Okawa

Kaokoveld, Kamanyab

Kaokoveld, Outjo

Kaokoveld, Outjo district, Altmark

Namib Desert, Blutkoppie

Namib Desert, Gemsbokwater, 18 km north of Ganab

Namib Desert, 10 km north-east of Gobabeb, Kuiseb River

Namib Desert, near Heinrichsberg

Namib Desert, Mirabib

Namib Naukluft Park, base of Bakenkop

Namib Naukluft Park, 4 km north-north-west of Zebra Pan

Ovamboland, Eunda, + 100 miles west-north-west of

Ondangua

Ovamboland, Onganjera

Waterberg

SOUTH AFRICA

Cape Province

Amadap Valley

Ashton

Augrabies Falls, Moonrock

Bak River

Bredasdorp, road to Struisbaai

Carnarvon, 10 km south

Eland’s Bay

Fraserburg, 8 km north-east

Gordonia, between Keimoes and Upington

Grahamstown

Grahamstown to Cradock road

Kalahari Gemsbok Park, 40 km south

Karoo, Beaufort West

Karoo, Carnarvon, 10 km south on road to Loxton

Karoo, Fraserburg, 8 km north-east

30°14°S 27°23’E

30°24’S 27°42’°E

29°38’S 27°33’E

28°58’S 28°09°E

29°50’26”S 29°06’52”E

23 1372S lao 25E.

18°S 14°E

19°00’32”S 15°26°46”E

19°03°48”S 15°48°54”E

25°13’S 17°42°E

28°0O1’S 18°45°E

19°39’S 18°34V’E

19°50’S 15°41V’E

19°30’S_15°05’E

19°40’S 14°50’E

20°06’S 16°09°E

20°06’S 16°09’E

22°48’S 15°22’E

22°96:S oe 3igE

23°377S 15°05S°E

23°14’S 15°3V’E

23°26’S,15°18°E

23°11’S 15°34¢’E

23-31 S15 285

17°31’S 14°39°E

17°54’S 15°04’E

20°32’S 17°08°E

25°14’S 18°43’°E

33°49’S 20°03’E

28°35’S 20°19"E

28°31°S 17715%E

34°35’S 20°00’E

31°10’S 22°08’E

327 18°S 1821978

31°52’S 21°538°E

28°30’S 21°14°E

3301775 262315

33°16°40”S 26°29’05”E

26°10’S 20°38’E

3222235) 227550,

31°10°S 22°08’E

S152 S21 eve

THE GENUS BRANCHIPODOPSIS IN SOUTHERN AFRICA 377

Karoo, Hoogeveld, south-west of Beaufort West

Karoo, Hutchinson, 30 km south on road to Richmond

Karoo, Williston

Kenton-on-sea

Kimberley

Koppieskraal Pan

Narugas siding

Papkuil, near Kimberley

Port Elizabeth

Ugie, Prentjiesberg

Williston

Natal

Drakensberg, Giant’s Castle

Drakensberg, Loteni Nature Reserve

Bushman’s Nek/Sehlabatebe Game Reserve border

Drakensberg, Underberg

Drakensberg, Underberg, Bamboo mountain

Umfolozi Game Reserve

Orange Free State

Bloemfontein

Van Reenen, 0.5 miles from

Transvaal

Benoni, Witkoppies

Kruger National Park, Pumbe Pan

Mala Mala Game Reserve

Mopane, 35 km south of Beit Bridge

Nylsvlei Nature Reserve

ZIMBABWE

Benfer Estates, near Umzingwani River

Bulawayo

Bulawayo, 25 km south-south-west, tomb of Rhodes

Matopos

33°10’S 21°46’E

31°30’S 23°29°E

31°20’S 20°50’E

33°41’S 26°40’E

28°45’S 24°45°E

26°56’26”S 20°18°47”"E

28°08724”S 20°18'47"E

30°12’S 18°21’E

34°S 25°E

31°09°09”S 28°07°48”E

31°20’S 20°50’E

29°16’S 29°30’E

29°26’21”S 29°32’48”E

19°50’S 29°06’E

29°02’S 29°30’E

29°43’S 29°20°E

28°17’S 31°49°E

29°07’S 26°14’E

28°21’S 29°22’E

26°11’S 28°16’E

24°10’S 31°5S’°E

24°45’S 31°30’E

22°36’S 29°52’E

24°39’S 28°42’E

22°12’S 29°56’E

20°11’S 28°35°E

20°30’00”S 28°31711”E

20°33’S 28°20°E

6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature (particu-

larly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed

by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov.,

etc.

An author’s name when cited must follow the name of the taxon without intervening punctuation

and not be abbreviated; if the year is added, a comma must separate author’s name and year. The

author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is trans-

ferred from its original genus. The name of a subsequent user of a scientific name must be separated

from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published scientific

names by which the species previously has been designated are listed in chronological order, with all

references to that name following in chronological order, e.g.:

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).

Nucula largillierti Philippi, 1861: 87.

Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:

comma separates author’s name and year

semicolon separates more than one reference by the same author

full stop separates references by different authors

figures of plates are enclosed in parentheses to distinguish them from text-figures

dash, not comma, separates consecutive numbers.

Synonymy arrangement according to chronology of bibliographic references, whereby the year is

placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable.

In describing new species, one specimen must be designated as the holotype; other specimens

mentioned in the original description are to be designated paratypes; additional material not regarded

as paratypes should be listed separately. The complete data (registration number, depository, descrip-

tion of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.:

Holotype

SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach, Port Eliza-

beth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

Note standard form of writing South African Museum registration numbers and date.

7. SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text

e.g. ‘. . . the Figure depicting C. namacolus ...’: ‘. . . in C. namacolus (Fig. 10)...’

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded by

initials or full names

e.g. DuToit but A.L. du Toit; Von Huene but F. von Huene

e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should preferably be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a book or

article, such as

‘Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial

capital letter, provided the same generic name is used consecutively. The generic name should

not be abbreviated at the beginning of a sentence or paragraph.

Name of new genus or species is not to be included in the title; it should be included in the abstract,

counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts.