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ANNALS OF THE ANNALE VAN DIE

SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

VOLUME 102 BAND 102

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

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SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

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D2109

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Page

BROOKE, R. K.

Annotated catalogue of the Aves type specimens in the South African Museum.

(RublishediRebruanyalOO3 ie vee ee nal mone te aust ess Serer ees eee ae 327

Davison, P.

Human subjects as museum objects. A project to make life-casts of ‘Bushmen’ and

‘Hottentots’, 1906-1924. (Published February 1993.) ...................----- 165

DINGLE, R. V.

Quaternary ostracods from the continental margin off south-western Africa. Part 1.

Dominant taxa, (Bublished(October 19922 )mam == a4 sed.) sense see eee - 1

KENNEDY, W. J. & KLINGER, H. C.

On the affinities of Cobbanoscaphites Collignon, 1969 (Cretaceous Ammonoidea).

(RublishedWManuwanysl993.)\0ar 9 ee ee tani. mars turin e acs oh a oho ores Moacl set obele > > 265

KENNEDY, W. J. see KLINGER, H. C.

Kina, G. M.

How many species of Diictodon were there? (Published February 1993.) .......... 303

KLINGER, H. C. & KENNEDY, W. J.

Cretaceous faunas from Zululand and Natal, South Africa. The heteromorph

ammonite genus Eubaculites Spath, 1926. (Published January 1993.) .......... 185

KLINGER, H. C. see KENNEDY, W. J.

MARTENS, K.

Taxonomic revision of African Cypridini. Part 2. Description of Ramotha gen. nov.

(@rustaceas Ostracoda). (Published'@ctoben199223)—.55- 5.42.5: 6655-22222 o- 91

ROOKMAAKER, L. C.

Additions and revisions to the list of specimens of the extinct blue antelope (Hippo-

magus leucopnaeus). (Published October 1992.) — 7-2-2... -. snc 2s ee--+ > 131

SHAW, E. M.

The basketwork of southern Africa. Part 2. Basketwork of the Khoisan and the

Dama-= (RubhishediPebruaryl993))\ 225. a2 cs noes e see eth seer tessa Sees’ 273

STEINER, K. E. see WHITEHEAD, V. B.

WHITEHEAD, V. B. & STEINER, K. E.

Two new species of oil-collecting bees of the genus Rediviva from the summer rain-

fall region of South Africa (Hymenoptera, Apoidea, Melittidae). (Published

INOVEMOC TBO OZ) Meiyeee re eee sie ot era Nt. cine oGbe a MENT ORGS 143

Volume 102 is complete in 10 parts.

Py P

|! 1

ney

nat

aed

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pal

—s

1 (oan

- it fh at

» ES ae 'D 73

Ot, a . —_—s

“ae,

m0 fy

1 1

iat

a) ce

7 it =)

) :

NEW GENERIC NAMES PROPOSED IN THIS VOLUME

FESCUCOKEHCH AUD IOC sO) a uniter ietate = arnte ehmewe re nye oa ueieds eine jaws a aiale aa Bs

Ramotha Martens, 1992

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

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ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 102 +#2zBand

October 1992 Oktober

Part 1 Deel

QUATERNARY OSTRACODS FROM

THE CONTINENTAL MARGIN OFF

SOUTH-WESTERN AFRICA

PART I. DOMINANT TAXA

R. V. DINGLE

Cape Town Kaapstad

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QUATERNARY OSTRACODS FROM THE CONTINENTAL MARGIN

OFF SOUTH-WESTERN AFRICA

PART I. DOMINANT TAXA

R.V. DINGLE

Micropalaeontology Research Unit, South African Museum, Cape Town

(With 59 figures and 6 tables)

[MS accepted 28 February 1990]

ABSTRACT

Eighteen species belonging to eleven genera account for 87 per cent of the total fauna of benthic

ostracods from the continental shelf and upper slope between the Kunene River and Cape Peninsula.

Seven new species, one new subspecies and a new genus are described. These are: Cytherella

namibensis, Palmoconcha? walvisridgensis, Kuiperiana angulata, Neocytherideis boomeri, Ambo-

stracon (A.) keeleri, Paracypris lacrimata, and Xestoleberis hartmanni; Bensonia knysnaensis robusta;

and Pseudokeyella. Eleven of the species have been previously recorded from the area. The remaining

14 per cent of the total ostracod fauna, none of which occur as dominants, will be described in Part II

of this report.

The faunas have been subdivided into ‘modern’ and ‘relict’ assemblages and, on a regional scale,

the dominant taxonomic groupings in both assemblages are the same. The shelf north of Walvis Bay

has a loxoconchid—Cytherella—Bensonia fauna, whereas south of the Orange River a Ruggieria

cytheropteroides—Pseudokeijella lepralioides fauna occurs. The intervening area has a mixed

assemblage.

CONTENTS

PAGE

Iinithoduch One peeare Ae bra ht cay dS ee Nar fons 1

Dominan tgtaxare eerie en see eal es 5

Ibiste Ole net Aaland SPECIESHa a Aue aay ee, oes Mesa 16

SyStemaltC@eschiptlons gaye ee sass) ee Sb ss cee u ee 17

SUIMDINETNY ste c'c BONG bid OES Ce Ces MEE RS ee 85

ZACKNOWIEG SEMEN S eae vac ea gusuoeen ek wee noes 85

INCTERCTICCSR Nan r Peter teh mantra ian Rc MOR kN» 85

INTRODUCTION

Sea-floor sediment samples from the continental shelf and upper slope (<950 m

water depth) off south-western Africa have been investigated for their benthic ostra-

cod fauna. Of the 269 samples examined, 192 contained ostracod valves, whereas 77

were barren (Fig. 1). Details of these faunas will be presented in three parts: Part I

(present report) deals with the dominant taxa, Part II (in press) deals with those

species that occur as minor constituents of the fauna, and Part III (in preparation) is a

discussion of the ecology of the ostracod populations and palaeo-oceanographic impli-

cations. Earlier publications have dealt with the fauna from a further 45 sediment

samples collected in water depths >950 m (Dingle et al. 1989, 1990).

The continental shelf off south-western Africa extends latitudinally from 17°S to

35°S, a distance of nearly 2 000 km. To the north, surface waters are affected by the

Ann. S. Afr. Mus. 102 (1), 1992: 1-89, 59 figs, 6 tables.

2 ANNALS OF THE SOUTH AFRICAN MUSEUM

15°E 20°E 25°E

PAG 9

+++ ++ #Walvis Bay

NAMIBIA

Namib Desert

Lideritz

beth

FTI, as Orange R.

Orange Shelf —

a ++

30°S +"* x

"+3 7m om Namaqualand

Pat SOUTH AFRICA

+H +H “

St Helena Bay

Saldanha/C. Columbine

Cape Peninsula

ome |

a eres Bank

Fig. 1. Ostracod-bearing samples from water depths less than 950 m

along the continental margin off south-western Africa. Those on the

eastern Agulhas Bank were reported on by Keeler (1981).

subtropical Angola Current, whereas most of the region lies under the influence of the

northward flowing Benguela System, which is subject to intense upwelling in quasi-

permanent zones (Lutjeharms & Meeuwis 1987). This phenomenon gives rise locally

to steep water-temperature, salinity, and dissolved-oxygen gradients, as well as

strongly influencing local substrates. Much of the area lies adjacent to an arid hinter-

land with low and episodic fluvial runoff. The extreme southern part of the region is

periodically influenced by inflow of subtropical Agulhas Current water from across

and around the Agulhas Bank. A more detailed discussion of the oceanography and

substrates will be given in Part III.

Modern marine Ostracoda were first reported from this region by Brady (1880),

who identified 14 species of podocopid Ostracoda from two samples collected in 1873

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 3

during the 1873-76 HMS ‘Challenger’ expedition: Station 140 (15-20 fm) (27-37 m):

False Bay) and Station 142 (150 fm (274 m): off the Cape of Good Hope). Brady’s

‘Challenger’ collection (including types of the South African species; see Table 1) was

re-illustrated, and lectotypes established by Puri & Hulings (1976). Subsequently,

several authors have worked on ostracod collections taken mostly from coastal sites

(Table 1): Miller (1908) reported on the fauna from one sample in Simonstown

Harbour (in the course of describing material collected on the German South Polar

Expedition 1901-03); Klie (1940) reported on ostracods from the vicinity of Lideritz

and Swakopmund; Benson & Maddocks (1964) described the fauna from Knysna

Lagoon (and illustrated a specimen from False Bay); and Whatley & Dingle (1989)

reported from the margin off south-western Africa the first known sighted species of

the genus Poseidonamicus. Also, in a major regional survey, Hartmann (1974)

recorded faunas from numerous localities on the coast between northern Angola and

Mozambique.

Unpublished theses dealing with modern marine ostracods from the area have

been produced by Keeler (1981—eastern Agulhas Bank) and Boomer (1985—conti-

nental margin, south-western Africa). Both these studies used samples collected by

the Marine Geoscience Unit at the University of Cape Town.

Tankard (1976), in his account of the Pleistocene deposits of the coastal plain

between Saldanha Bay and Elands Bay (Cape Deseada), identified 18 species of

marine Ostracoda, 13 of which he was able to refer to previously described taxa

(Table 1).

TABLE 1

Previous records of ostracods from south-western and southern Africa".

Walvis Liideritz Saldanha Cape False Knysna

Bay* area Pen. Bay Lagoon

BRADY 1880

Pontocypris subreniformis sp. nov.

Macrocypris maculata Brady

Bairdia ovata Bosquet

Cythere exilis sp. nov.

Cythere flabellicostata sp. nov.

Cythere lepralioides sp. nov. X

Cythere craticula sp. nov.

Cythere melobesioides Brady X

Cythere cytheropteroides X

Cythere stolonifera

Loxoconcha subrhomboidea sp. nov.

Xestoleberis africana sp. nov.

Cytherura mucronata sp. nov.

Cytherura clausi sp. nov.

Cytherella dromedaria sp. nov.

AK mM RK OM

aM mK OM

MULLER 1908

Macrocypris dispar sp. nov.

Macrocypris africana sp. nov.

Pontocypris gaussi sp. nov.

Pontocypris flava sp. nov.

Xestoleberis capensis sp. nov.

Xestoleberis ramosa

A MO

4 ANNALS OF THE SOUTH AFRICAN MUSEUM

Table 1 (cont.)

Walvis

Bay*

KLIE 1940

Pontocypris flava Miller

Eucythereis mirabilis sp. nov. X

Eucythereis levetzovi sp. nov.

Procythereis serrata sp. nov.

Procythereis major sp. nov.

Procythereis minor sp. nov.

Xestoleberis ramosa Miiller

Xestoleberis crenulata sp. nov.

Xestoleberis ferax sp. nov.

Xestoleberis baja sp. nov.

Xestoleberis humilis sp. nov.

Sclerochilus incurvatus sp. nov.

Sclerochilus meridionalis Miiller

Cytherois minor Miller

Paradoxostoma caeruleum sp. nov.

Paradoxostoma griseum sp. nov.

Paradoxostoma angustissimum sp. nov.

Paradoxostoma auritum sp. nov.

Paradoxostoma reflexum sp. nov.

Paradoxostoma semilunare sp. nov.

BENSON & MADDOCKS 1964

Cytherella aff. punctata Brady

Bairdia villosa? Brady

Paracypris westfordensis sp. nov.

Aglaiella railbridgensis sp. nov.

Perissocytheridea estuaria sp. nov.

Sulcostocythere knysnaensis sp. nov.

Cytheretta knysnaensis sp. nov.

Loxoconcha parameridionalis sp. nov.

Loxoconcha megapora sp. nov.

Xestoleberis capensis Miller

Hemicythere? sp.

Nereina? sp. A

Nereina? sp. B

Aurila dayii sp. nov.

Urocythereis sp.

Mutilus sp.

Bradleya? sp.

| HARTMANN 1974

Cytherella aff. punctata Brady

Bairdoppilata sp. 44

Perissocytheridea estuaria B & M

Sulcostocythere knysnaensis B & M

Cyprideis limbocostatc sp. nov. X

Cyprideis remanie Klie

Mutilus bensonmaddocksorum sp. nov.

Aurila kliei sp. nov.

Aurila levetzovi (Klie)

Aurila petricola sp. nov.

Procythereis major Klie

Procythereis minor Klie

Procythereis serrata Klie

Procythereis foveata sp. nov.

Caudites knysnaensis sp. nov.

Cytheretta knysnaensis B & M

Loxoconcha walvisbaiensis sp. nov. x

Loxoconcha megarora vat. magna n. vat.

Loxoconcha parameridionalis B & M

Australoxochonca favornamentata sp. nov.

Australoxochonca parafavornamentata sp. nov.

Semicytherura dayi sp. nov.

Hemicytherura? kazmaierae sp. nov.

Liideritz

PS PS PS PS PS OPS OPS OM Od OO OS OS Od Ot OO OO

mS PS PS PP OR OO

Saldanha

Area

Cape

Pen.

False Knysna

Bay Lagoon

X X

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 5

Table 1 (cont.)

Walvis Liideritz Saldanha Cape False Knysna

Bay* Area Pen. Bay Lagoon

TANKARD 1976

Paracypris westfordensis B & M

Aglaiella railbridgensis B & M

Cytheretta knysnaensis B & M

Cyprideis cf. limbocostata Hartmann

Perissocytheridea estuaria B & M

Cytherura sp.

Hemicytherura parvifossata Hartmann

Bairdia cf. villosa Brady

Aurila dayii B & M

Caudites knysnaensis Hartmann

Procythereis sp.

Urocythereis sp.

Loxoconcha parameriodionalis B & M

Loxoconcha peterseni Hartmann

Cytheromorpha sp.

Bradleya sp.

Xestoleberis capensis Miller

Cytherella punctata Brady

AK KK KK KK OK OR SO

"—excludes Myodocopida

*—includes Swakopmund and Sandwich Harbour

B & M—Benson & Maddocks

Additional relevant studies of modern and late Tertiary ostracod faunas from

adjacent areas include the following: Skogsberg (1939—Subantarctic), Benson

(1964— Antarctica), Bold (1966—Gabon), Neale (1967—Antarctica), Maddocks

(1969, 1977—Southern Ocean), Valicenti (1977—Patagonia), Babinot & Kouyou-

montzakis (1986—Congo estuary), Hartmann (1986, 1987, 1988— Antarctica),

Whatley et al. (1987, 1988—Antarctic and south-western Atlantic), and Coimbra &

Pinto de Ornellas (1989— Brazil).

DOMINANT TAXA

Approximately 120 species have been identified from the continental shelf and

upper slope off south-western Africa within a total collection of 24 058 specimens

from 192 ostracod-bearing sediment samples. In this paper I discuss the 11 dominant

groups (10 genera and one family) that individually account for 2 per cent or more of

the total specimen count (Table 3). This is merely a convenience for handling the

large data set, but the eighteen species concerned account for 87 per cent of the

overall population and give a first order impression of regional taxa dominance. Dis-

tribution patterns for the total fauna will be discussed in Part III.

Table 2 lists the locations and faunal contents of the 183 ostracod-bearing sedi-

ment samples that contain one or more of the species described in this paper

(Table 3). The eleven dominant taxa constitute a mean of 82 per cent of the fauna of

each sample but it is notable that, in relation to the depth distribution of the total

fauna, these taxa are most important in water depths 60—400 m (mean 86% — Fig. 2).

At inshore sites they constitute 74 per cent, and at depths >400 m 66 per cent,

although they are relatively more abundant between 800 m and 950 m than in the

TBD sample no.

Latitude (°S)

22,895,

34,0597

32,6833

26,05

34,2033

34,09

32,62

25,1166

26,05

27,95

34,74

34,45

33,1566

17,5333

19,9166

34,7733

25,1166

20,45

23,9333

33,9608

19,9166

31,0883

27,925

34,09

19,9166

34,0923

34,015

34,0267

32,7467

28,255

17,5333

20,45

17,5333

34.0925

19.9166

28.8

31,8333

E5833

34,5167

34,97

28,4333

22,9333

19,21

20,9166

33,995

29,7833

Longitude (°E)

14,4916

18,355

18,0833

14,9333

18,51

18,3333

18,0417

14,7466

14,9

15,6366

19,5216

18,575

17,955

11,6833

12,8933

19,5083

14,6666

13,1467

14,355

18,2733

12,86

17,6467

15,5166

18,2825

12,8233

18,2717

18,264

18,2

17,7617

15,79

11,6416

13,0367

11,6

18,215

12,6416

16.1833

17,8083

11,5666

18,6833

19.6

15,6

13,8883

12,1766

13,0366

18,1483

13,7133

15,1583

15,9666

12.4333

17,4333

19,5083

11,5333

14,1333

17,7766

15,1666

18,1733

13,3583

14,1066

13,7567

15,4

16,4167

11,5

13,45

16,2167

16,0167

Water depth (m)

ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 2

Location of ostracod fauna of sediment samples.

Total no. of valves

—

pf3o f)5 Ooo 0 oD DO

—

“4 4

WD si Crs tr me tr Nn

NO OU

ie.)

CORR) pees NAM: Ns es st

Pseudokeijella

lepralioides

'o1o4 On

Oo oo 8 oO 5 op Po ]n no 2

pare ep Me te t + CNet tt

bon SOH 9 0 6 nN 5 A

Ore

COR sr re Dar ts tt WO sr 1 Ws OHM,

lon

Ruggieria

= _cytheropteroides

horot ot Da an tp 8 eee Baie

CO ce St Te

fo)

rrrte NN

—_

—

170907 PND 90 On 0 kN 0 Oto o on OO

roroe Nr tr Ns st

(rNe st CH7ANMR se st

ie)

Ambostracon (A.)

z _ flabellicostata

®PNwOr ss Me

oop ma 4 COn ts

Ten Oe ei 0s Cee ero Oe ee es De eet te SP Cie thie!) 0 ie Oe Dit 0 SOs Oe 70

vs)

CO A

ee te Wir a We

— tN

1 CO}, NHWi ss 5

me B&D

Ambostracon {A.)

keeleri

reorounndo

Foo oto 00 0 00 0 FD ONS On op ORM op poo op Oyo pp Oto oO oo

1 0 7,0 Of oOo 4

Doo oO DDO 0 0 9 0

Ambostracon (A.)

levetzovi

0000 08 Dt oA DOD Oo oOo 8 Dp ol hl DO OD

bye eT te NE Oe SO TS ty ete Uw tee Je Deed tem meule eee te Mhatieet pede) cil fb cy ofan

o900 9 9 DOO DSO nN Do A oOo Dd ee Ao OI 6 on pg oO eH hl BD lblp hb ool lolUflhUvhlUn OU

Henryhowella

= _ melobesioides

909 00090 0 0097 097 © OD 8 DD DO YD

AS PAU AY AN esi emia OSL CRN MOT aNL ope Gun ghy) URI) 8) sare aie vy Ue kel bp Mare ays tul a fan seiera Ale yal estn sal, Sah ANN eb) tah peeks

v2)

rr Weaias

N io)

Oi mi in ts One eo ee 8 a em

v=o Se eee Oe ees Od Nea tern neo on <n oo eo

Q~r is bt at nie

Doratocythere

=z exilis

T= ee uahe eG

Veen aed at Aw Une Vem Veet ook urn Ue (te (femme tot STC TCT on) tt Deas!

CEO ET ee OR a Let Pi Re Yow tas ere <(MeenY be =X ORI eeeC ee tees eaten Wee itia Mme (jn ejuev) lye tr "le Ty) ol. Thoth ah

©Nepaas

con

DO a) OOK

nO —_—

PU ot Po tS a oO No Po 0 Hea SH Oty on no eo 8 0 6 on a eS Sn 6h oo

Neocytherideis

< boomeri

—

1 C1 Mi itn a Oe ein mee te bt bob tb tb tt ee

eK) TNs Jim bes Js Say a OTS URE ET Dear She hae Fen fam) ee ee id lm Jie COM Pee Thm Chee el mel tie Slarh ed eee ah. 20)

le.)

1 OOm 6 WM te tt me tt Ma tt

Tn ie Se

Db oe

Ona: 1: 1

OQ =

poe Dr ge tb NAG tt me OO me

—_

Palmoconcha

= walvisbaiensis

1 ee Ue ae ca) eee

Meet a tp aDMa NNe st tr DNs ete sett

7 O80 0 DISH 0 0 Oe Po 0 9 oo Sha eo 6 6 pb WF no ASRS) A oo

subrhomboidea

angulata

Palmoconcha?

= walvisridgensis

‘Kuiperiana

Palmoconcha

po Ra a oo te Oo) Ach tf 0° o O ao bo ooo 2

—

AS 208 SU Sa NN a ROE NE a A ee aa ap) Sp ae an a

(Pleas! Games! Wat Vaesas) Youmen) Tobey Nmmrt Wee Yes (igi (em (Yim Jem Dctst Yoon Yay Jone) Clty Framer Piet Demy Vue Pleats Tome TS fT |

Suet ee aay Ca) tay ey an) a Ses se a ae A aa al Say sine oe

Cane > iets et aec Jims meena Je YJ et Viens ome hun 7 be et emi June) Peet oe MU ets Sur, Psion arf)

IA AR path eR) (dieu aia ae aod ees) Sie ine met

—

er Teen ee Peer Jom yom james ey Yee > NY Jone oY fee (Oe: eee fer fee freer Dees et oe y(t jeer oo fete ope pote (1) (ee pore (= otis leet eet)

Pork kt WON 2 it tk ttt IN Ds wt Ms 5 tt st WM st tt a

eR ee ae ee ae nee ee Ga KG aS al sae a as ae ew ip an Seer Say cae ish Sey tas sn, aa ys

Fes ets Tih 10 me fame Lect Yoneurt Jems? Ne ee Veamee [Jemint ugar ca Soteart Julie Jeunes Ne Ti url Sandy [emir Flas Jet imsiRct Jr ur Jen amie Pomuty Quumeet JUlmecsy (ct omer Irae Tomes Doe” Dhl? Tamey Tenet}

th ee tr age a a se a ae a ay A a a Te a an ee sg ey a oe i sp tay a a}

foenah al SU 0 sh om te ae ea one AND car Gay ety ae Tap apie Ge) Gs) guy gt tak ce age) dp. Geleeniay ase ah

Yi icy Fetesc( fame) MT SUK SOY Jes Tees > Vr Gam JURY ET VR J] = Js) Tom Fen TUN Yomi Tmt a Vat UY GO Jo Jc) ToT Sal Pe (Oy TTT

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA

Cytherella

namibensis

Cytherella

= dromedaria

Paracypris

lacrimata

” Bensonia

knysnaensis

< _ knysnaensis

Bensonia

knysnaensis

robusta

Xestoleberis

hartmanni

ys]

Total Relict

ys)

rs)

to)

292 42

NWO

i>)

an —

AS = OT TD DD TD OO YT

Le)

—

—_

—_= WW

ao ~

Mii: ot

eS Oeste Jt te Thc th es De ot SNe aie teat ue He en bere pen ne ne or cre a ten

oUt ue th 0 oe Gee Te Rs oe Oo 0 ee to Soe ees

‘oO

ee AN a rere te See edie eee ii ber peat in | (emer brie oe ky Sb ues

{ heset Verte] Koutieg Hess <=) tetsu Jeicst enys JS! Ya gy Fosiets Ramet ine=t Jeet’ Bruny ome 7 Ve Yom, Feet Yc=t fusmmatt filmed (ome (un Fe 17]

CV fate ee ee Tey ey eee [fu etl bee mee teas Fae ha Wee tami hae emit clam iQe ih ee cheeth 94)

(hie ee Ip nomen utes ee oe OSes Gun OuerNe akesthea(mmhovieo) ath ove 4 ah 0

Oe Dea De Nee hk Di ST ie 1S 20 Oy te Se ee

—

Mt Wrest Ns tt Was st tt W

pre Bes tt HDs tt st Rt tN st

—

Cine Josue’ Josue ames foment ess emmvY fisert fmm aery Josie faces” ONT? Juss) Jun) YO Jone? Dusuiet JONNY Fett Ties Hamm W200? TRS Yas [oT (TT Bn Joly NT ORT rt Tene omic ae 0 «OJ J (Sy J Fa

—

Con es ts Wr On Sa cd Bae em et ee

Ne)

sei ce ate GCE ys mt Sarl filmes Ft Hom te it Tog Pamest baer Detect Famers ey Veco Inet Hem}

—

Tesoro eVect = Seto <> er Veer mie feet Tage Peunc( JoJo en uty Jest ert = Yor1 inti Yoo ose Jot Va mk HEC Deane Joc) omy JON Jo meet oe Youu Yee =f Dee =)

Ok ew athe Wi 0 a a =e Oy Ol i kOe oe a cor ee Sy a Oe Fe ae ee et HP cee ae pert Te ele eyes eC alin mei ee eee esti ses ie()-5 Mendy) etl ye ined

pei ee eet me a) ty Qe Oa gy a et ta ee a so

Se aie Abe ye ie ate ace Calpe ayn tavicaa) a) “Noga ene un vse ay SS 8. tah a Aa tee Ge) anak sh Gun) ee tah ue

4 Ape dpe Oa ae Gs: Ut) it OOO eee) nae AS Sie ee ue ue ey sy Ua ge Ee Al teh as

fpocth Mite Ue i th Leite tho Aho saoth sth ap ech eerie etearhe ) Sreenh ie arent tjucete me ttieaa

BSS

Wo nono on oro 6-6 oP bo ob oo op So oo lp ht hoy

Mr ir ra nr ms Ns te te tt Re tt tt Ss tt ltl WN a

Li lee duet (lie etl Pitcreeeet [i aha Khe Jur) CH He Seat) eT HNe ct Jin Ui: [Needsecm—arl Tolmer ct 100 set Fy Jou Font] fre joey Te ur (emus foment Yuna) Juin Deiat)

eo vine ei Oe toute Ui WN ese eeet ies tie at)

Total Modern

Subtotal

iSS)

FPrRONCUNMNNOOre

—

SOrDCOWRRFPONNNON CON STWAINDWOOA

—

Table 2 (cont.)

TBD sample no.

Latitude (°S)

34,5833

28,8833

28,4167

30,35

33,1916

35,1333

31,9583

28,4167

28,8833

34,7

28,9

29,1666

20,4333

30,775

27,1167

26,1166

29,95

30,9166

34,15

35,2366

30.4

23,93

2).

34,9333

25,6

30,9166

34,185

19,1916

31,9666

30,45

31,3133

29,3166

827833

30,5333

30,5166

31,2333

65535

23,4333

27,0833

22,25

24,4333

34,0666

34,2383

22,9283

26,1833

32,65

31,2333

32,758

20,45

30,5166

34,6

31,9333

25,7833

Longitude (°E)

18,5867

15,7666

16,8166

17,7033

19,2833

17,2366

14,5833

15,5666

18,6

15,3666

12,3417

18,45

14,8

17,23

14,96

14,3167

15,65

12,3233

13,5733

18,0317

14,9833

12,6833

15,925

14,8667

14,4666

15,4167

W5)57/

17,995

19,1466

16,4333

13,9066

15,4333

18,4833

13,2333

18,6

14,1333

17,945

11,9833

17,0416

16,0666

16,8583

14,7166

17,4333

15,6166

15,8333

16,625

19,1333

13,3967

14,6833

13,05

13,87

17,7216

17,8667

13,1633

14,1333

17,0883

16,3833

We?

12,34

15,2166

18,3667

16,6583

13,925

Water depth (m)

ANNALS OF THE SOUTH AFRICAN MUSEUM

Total no. of valves

Pseudokeijella

lepralioides

vs)

WwW a

WOR DHWDNND HM:

—

cytheropteroides

Ruggieria

1 Di Nn

Ambostracon (A.)

flabellicostata

—_

Ambostracon (A.)

keeleri

Ambostracon (A.)

levetzovi

Henryhowella

melobesioides

Lelie |

rOr Rest Rs COR

Doratocythere

Neocytherideis

boomeri

vs)

Palmoconcha

walvisbaiensis

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA

2 3

a. fF

35 s= se 5 2.4 2 a a

3 ey $s “ aS 28 aS < x se

£8 £3 aS Ss SS zs ss SVs Sas) SS S

S38 si 86 aS} 3 iy = =Sss Sof ss ESS} ray Oo

eS 9.8 0.8 2s LX as SS RSs QS SS S) =

oS SS Ss es ES = S85 $s S-2 SBS = }

& & 3 es SS aS 35 255 229 Ses SS O) =e =

= ss ss so = gs £©£ss £ os as as) ~ = =I =

33 Ss sa oF Sy Ss S ~~ Se VS Sirs ae = iS

< aX Qu a 4 a) ~< ~< 3 3 2 2

[o) ° 3

R M R M R M R M R M R M R M R M R M 1 i na ss

Bi w

—

—_—

Pe, ee eet eh meatier ema ete att = hot —S eT) ane iy

een 1 plea (eee ieeen Cee CU, ieee! Poeen Pem

gd Obs af SAT Tt tes Tee ere iT SS CL ane irre Chueh Te hea aya i et Jn fas my Jc] me (tt Cee Ta) ine ORY Vy Loy Yel ele P| > ee Je Yemy et Fon) — Wats Pe

[FS STs et a ee Ta J a ae Ue Fi a (ed

TSU Ue Beem tee Sur Gu Jee ster maa laf oMNDSas wih” fay Na ay cae ah Ga) Ue Se et) Qe tn) Bet mire? 0 jneern unr dee one

rey Re is eae es ee Kee ser Ge Oe tec ra We

0) Oe 0 De SU SO) 0 She SOP a ee Ole Cea)

1 pl ae Sa mel Aa Ga tay iep an aes Sa dee Pe esa Si Poti ay ae TA tee. Whee

ae el ae Nal wey Mer a ap a

= il

> 3

s 8 il

Saat

a 2

a il

an) 3

= i

Ct o> ee et

OD Or 0 ee 0 fm 0 0 Of 0 at foo

rr Nos

One 00 20 0 LF 0 Oe 0” a ore at ete ate = TL — fe TO es ON nOn yet Ee ee 90

CD DT

0 U0 OTYO Oo 1 O 0

Co SJ-o cist fists Woo 6 0 0 6 6 oo oo Oo Uo

cr _—

tO: SOTO Ou 0h Sn et st Dr mer

6 prob o og o o 6b 0 So 0 6 ll

Ott 7 71 1p

lee)

lon

114

nN. =>

I oS

BRON OONAMNAINOMO~)W

WwW Ww

— N i

ecoo°dcorcdon

SODTOCOWOW RRB NOR OWROCORHACONALO

CoonocPecoor

SOOOCOhRCTONOOMNrRCOCOOCOCOCOOCOrROCOOFRFCoO

iw) —

- (oe) SS)

BR CORR WOONDACONWOMC

Lo)

Table 2 (cont.)

ANNALS OF THE SOUTH AFRICAN MUSEUM

i) => => aN

S = 3 Ss = S ok Y bs ‘

— — ~ in}

a eee i aS 2s s8 S. NB Ss @ Se 8.

& ~ o a. © 2s 8 a3 as ax Ss Ss == 8:

5 2 3 = g S5 2s Ss Ss Ss Se Ss g§ =

A 2 @ 5 “= = gS =~ &* = a = S

(oa) os 3 bY io)

= SI =) = & R ™M R M Ren R R M R M R M R M R

4015 22,0833 12,9333 325 10 > Anata a ares uf ie So ae a ge A

3789 21,9 12,8166 325 8 > a Bee aye Eat cts aus Sb ae a yale Ba :

3522 22,9333 13,345 344 44 a) ee 33 ate aes ee ae ae, Ray ae tee BaD. =

2AT0) 9307550) 1555) 345 59 Seem Spy Ho ode an ae i. Bie ct gal :

3866 19,9166 11,9133 348 7 = 4 ii ie ae ea ieee Se ne ae = Se és

3171 27,0833 14,5167 349 4 Lon Be va fo Ete Cues < ee Se ree c

2447 31,925 16,4666 350 49 eS ee) ean nes ae Bs at ae Ve ye =a 5

2976 28,4167 14,3833 350 18 : = he ae eyes ee ae eee: é

3256 26,2166 13,95 352 ll - A ee A re eae. ian ee ae ae <i 56 pie i

3923 19,1566 11,5 368 8 > 2 aS Lies seus ey Le Da ee Bw coe «: 4

3943 19,7333 11,8167 373 10 se mace ae ele ae ee Eat ee Ste

2825 34,1 17,6 375 8 et es Ae > = 8 ceo age iy Oe 2. ae oe Ce i

3556 23,4333 13,2167 379 31 5s Taw: Sees See a var 4, ae 4 ae 5

3826 20,9333 12,4666 382 23 se eae ane ame aes Se tae ae We 5

3359 26,325 13,875 385 12 - Paes) Sire Eo ie Buia ce ae eae

3255 26,2166 13,7833 390 1 Se one er re ee vie ue ae ve 3

3562 24,6067 13,5833 391 69 > so MM 6 CAPs ae eo) a 24 ae ae £

2448 31,925 16,255 392 37 = 5 Bi) « 2 6 sc By Ses Q@ ol aes ae 4

3172 27,1167 14,3 403 3 a: Ine we ae BES age dite ae ae Ea: i

1694 34,75 18,3267 425 31 fl il os Shines es 6 = sel (Os 2 a Leslee 5

3462 23,9583 13,1916 430 40 BPE r eM BL, 1D ay Se ete a he BN i =. iS fe

2488 30,8416 15,4666 430 14 ee 1, wee a ee Sees ee a0 Tie eae Bea) e

3179 27,0333 14,1 437 79 ee ae AOR ES a8 an ee agree z)'adge 2 ae Oh a: i

2262 32,7917 16,8167 450 3 a ee paves =, ee aye te =, wa eS. ae :

2440 33,2416 17,15 450 18 = fl os eae ke brs iis % 7 Bee ea) if

3577 31,3667 16,0833 453 96 = = 8 2 > oe 5) he EN as i) es: snl oe fs

2700 30,2167 14.85 469 13 > 54 83 2 6 ie ce, ok i fe Pat cP ie é

2780 33,2416 17,25 475 42 2 a OM i > oe 2 > 6. soe ec -

3524 22,9333 12,9666 475 64 ag Bz Soa. ©) on pee Su oe Seek ee sie ee:

301 35,0333 18,5333 500 7 3s 5 8 ee Be oe Bn, ce a. ee ee Sr, Nae :

1698 34,7833 18,2167 502 10 So Pa a Eee xe ae Ge vee aie 3 ae 3

3225 25,1166 13,6 530 1 oe Bre ee So Fe Soraree a ee nt sores ae ne x

2879 29,4833 14,55 530 61 ey ee G8 > ¢ Sb & 8 Bh we = - = -

1697 34,7667 1825 545 112 ay = 8 a Sane 2 ate Gan ee ee oe

2S SPS KS SD 2 : rvs 2) ta oe se hae at tes oes Sere f

3845 20,5917 12,0833 566 1 : ee ay te LS es Ne A ieee ae Sy i

3555 23,4433 13,03 590 19 : Dit ie ca ae: = ie ae ais ee cua Cie e

3112 31,45 15,7667 648 11 - Ss a ae My cate = Su) 0c ay tena E

3561 24,58 13,4167 655 9 a eae are eae ae Otte ey: a tee G

3458 24,1 13,2 725 16 - eee cee BD ie ay Le aA 9 ae ae i

3346 31,12 15,575 730 23 0 ira eae ae ae ee ieee AG ae Bg i

2978 28,4 14,2 736 38 > : BT ya ae Se Ok WORMS ae Sale ‘

3921 19,1366 11,1666 738 4 Z ae fe ane ee Maes eh ge in Lae a 2 :

3885 17,5666 11,2833 779 10 Oi me ee ehatae ee ce ed 2 gee eS ee Pi i

3846 20,5833 11,8667 810 1 oe pa See ore upp, (Bs ee ae Sei ee Tyne sie Sa i

3869 19,9167 11,3667 825 1 fs = 6 ties ape ee op tS ha es an: ate 4

3113 31,3667 15,5833 840 16 seit Sw ee ts ple = whe a) aoe Salles a ie :

3525 22,9333 12,8 850 14 2 2 Se hes = 5 WM 2 > oc a gee fi

3461 23,9583 13,0166 850 13 5 8 Sees ave etate ae ee q OD <2 are ‘

3109 31,9 15,9 900 80) 2 os ae sai Pie Bhs sn ee Sea {an Ue 5 ie i

2697 30,2833 14,3167 940 12 - a M8 eS eae GO = ie ig ae “

3704 19,3666 11,06 941 16 > 8 s 6 - 5 Pus 5) | Wo. ee ars 2 oe :

3341 32,6833 16,5167 945 36 2s 3s an ve a MR oN ve Se es R

Total valves 24059 6235 1946 5266 223 442 48 989 35 11 8 415 14 637 O 486 25 414 6

% Relict 345 - 202 = 24 = 54 = 00 © 2° 22° (=) 3's ee Cee eo

% Modern - 68,0 = bi! = Ys = ip - 0,2 - 0,4 = @ - 0,8 22,

7% Subtotal 391 > 262 9- 23 >) 48 >, 00 =) 20) 1) 13:0 mee 2

% Total ostracod fauna 33,8 = WRT - 2,0 - 42 - 0,0 é 1,7 c DG > Dil = 1,9

Abbreviations: R = relict fauna, M = modern fauna

Kuiperiana

angulata

Gy Tt 0 6 OD 0! es TT So 0 ie oT UC act reset Ta st Deca feety (hd Frnt Heumey Yay Jey Piety TR J Fee Tint Ces Joe Jet Peet Some Yume (fa)

ETI StTs eUat Tots) Mr Fan feof RT Tier eum UL Me tk Oe ll Fame Ce ey DU Limes! lowest FOP Re 1 Leet (S-MT Loe at (ieee fear ol Fae Foemet a [Joma TSC hemes t Tiesgnl Demet Pleo acs Lume Foal Sent Fey oI tem Sm my cel eee She eC wet)

Palmoconcha?

walvisridgensis

(Pa Tee Meet sont ee Comet eer Neel jm att bileert Coan} Yoel fae ai 1eccl fume Toe ieee Cima Je -{0e eel Paka tone Te ou DR Mt Tmo Lees) cL) cat er ee Ramen ee eect Ls el Cem em Tem Yee OC ICE The af]

Teed Gas OPOy) LAGE Oa STG M ey WA Uae iahe TA WNRcTIM ES AOL ONE TAA COC aa) sia;c ar eS Prete MDD CUNT cer mide 10) Sil Ae eae teeta ce ae le a Saya KP OS FY ley engh rtyee:

Palmoconcha

subrhomboidea

C8 OT Oe eas Se Ae ST De BD) SC aT) Swe oe Dee ee 2 Famed el Fees (Tamer Item Let LG SL Ja a eset heed Suid ema Jed Cao Cece FON Jom eee a TU CY DT |

vs)

—<

yo)

—

DiriNDnDse ts NYVHi COMs ts 1s 1 DROna

—

Gory ts tt Ns

oro op be eb bh ob te et Ne bk tt Os Wes 2 tt

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA

namibensis

dromedaria

Paracypris

lacrimata

knysnaensis

robusta

Xestoleberis

africana

Xestoleberis

hartmanni

Cytherella

Bensonia

Total Relict

Cee NS SONS eae eM Vig Oe Se ORR RGM MUR Ny I CUCOTICee Oh byetemnat: GOln Iee ify cB tal oun ous! on8)) (Atom seewOl er wh keene? eb nny ah ual Tals teh wun es erg) ish oi an ty ae

Tee Aaa nebe cow aL EOP Ui ual ney vale ate Sth sie nit 20 v0 ORe sd CEL eRe NS ie NOs MeteRORM FL 62 8) 00 hee 8s lec as eh en a, aa a sa TB el eb np tn ae eh ne a

es Oe BO OO BO) MOL Oe tle Cie SCR tee) Us et Ved Tint Ts Cet eam homey Laan med Pied Porm Jeu Jemeany (ue en eet Hewes Fister aminl Teaieed Haunt Doles Jee en Ht ie at Pat (eet Peed e-ia chee 0

Kets Thee Gee ceney LAP ia cee ets) Ue MAN mMAN GOR Mnd? cee cp) OLN an. Cfo ere ces Copouss) ame ne Wipe lose tte hee teU Gh Cpeest ty gh Mee tele se Se; Wale) se) Te crap fa ce® Sa ap at ge

Co tee. “ODS k0 ee alee) De Cece a: ST iirc ce wel Cede Nee! 7 2 ete Tefal roid ied ee Cane eo Jagd Ve oe Ae ee et Neel be Dee eats etait ome eticn. Ti? of]

TPs at CR eect his es i Ce yer le CN ee Cree (Ie (eed ome a (0c pret Fel] farm Tite ieee F=f Yew a Doma 4 Vena Jove] Ss eet Gee Me ees eh ie co ie) et Jie MDa eC eee Tots Sameer ea Eee te m0)

ese ON SO eet OF et eee ee 0S a Tei De ery ye ie i mee of el eM Pe ed Peer ee eee ye ee eee em Sens 20S te)

OD Ce eT ie oe Ue Ah wile sO i ye 10 ar Yom tk eens Yes om ICRA tu Ld Pett Pee figs EM) eC Pent PO eTown Ved Dory WOH LOS Tut Pees eet) tt Vimeo Je ToT eee (ai hati ( Fame et)

Cite 0: 0G oO. 0) 0 0) 0 hs SO Os 0 POD =O VO DG 0h ea TT Oe eT, el Me set ee rhs ROB LTE Me AY OE) of]

[lead uth wet (hae dP yl heel? Me eds (he Cle, LHe teat eect lei Joacl em cA iy Pm i= et Pie Jeena He ul Viaced Piet fT henna Mn Foe eae aut oe ec ee a SCs Ee he mo Ci CO Oe er O

CO I ty OO he Oo Oi 0 0! 0 ee et TR 0 Se Ce eat hat DG Ss ae Casts ate Chai See Oi US i 0590

De oath Toe Oy He 80 a A 0 ee AC heehee ime eel Dee ME ome kay Jism Jat Pad Wee QoceeU er aC emi TOM Coat T gt Ii st Tia aa) Meal ad Ye) ae SPs Pool ea Poet

a OO ss OR ea os

Ne)

NA:si NSO

—

57 993-318 4

1‘

NY oN

a a)

Total Modern

SGNOHKHNNOCCOOROCONOCOCOSCCONCTDONNWOROWONOCTOARDCNONCOCOH—$CCCoCo°oo

2 18046 2861 20907

Subtotal

2 * P

a ‘

ee ‘

}

ry v

} .

‘ 6

? . i .

5 Kt

hs \

F, {

? 0)

' A=

‘

’

= 1

7 ?

ti ‘ x

‘ ; i

| J

)

4 in 5

4 \

} “

‘

1 A ‘ 4

{

; 2

\ Y,

)

: )

7 ~ \

ss é

C ha

‘

0) 5) 5

an ‘

j ; 1 ay’.

vo ‘| { ’

if r veh

? b 4

ae * t i \

aie ' e

fe pb pe v

ANNALS OF THE SOUTH AFRICAN MUSEUM

10 >

Table 2 (cont.) b

RU ke) oS = 3 x # af

- &§ = s ss 5 5 s3 $ 3 gs

: = e 2 sy 8 $8 s Ss 33 S 1, 25

Pg 22 2 Ss ae ee

Preece ec ome. ss os= Fs 8 §8 §5

OR ae a

= = Ss

2 g & = e Row RP ow RY OM RM R M RM Te ko R mM >

4015 22,0833 12,9333 325 10 oe a 4 = aan A uae er ek: es RS aS aes, |

3789 21,9 12,8166 325 8 Sake eat Sh Sa ara a a rate os ws

3522 22,9333 13,345 344 dt Sie Rd MESSI Biwi ven ayes ee ae atta

2470 30,755 (15,55 345 59 SPO Wee ae," ee Sse = lee tes eens a aes eee

3866 19,9166 11,9133 348 7 ae ot Le) “ois i wee ales ws Stes shee | a

3171 27,0833 14,5167 349 4 = 0S Seas = on Ss ap ae re es Sy te iat

2447 31,925 16,4666 350 49 Seal ay). any Bea te 5 eee jee ee ek eee

2976 28,4167 14,3833 350 18 a NS eee - : =r ye See: ak ae

3256 26,2166 13,95 352 7 =), aE ie ite Se Sofas oe aye Ss ules ee

3923 19,1566 11,5 368 8 2 a aS emt = ; eS. a ies Sait: :

3943 19,7333 11,8167 373 10 Ss - - CaS SP =i ee Has a ihe ehh eh

2825 34,1 17,6 375 8 eet ae So = 2 - - - z ae : =

3556 23,4333 13,2167 379 31 Bil, ete eg T iD - - - Se ae he Paes =e

3826 20,9333 12,4666 382 23 = = - - Etat =” fc - -

3359 26,325 13,875 385 12 Seater a8h 12 - - - - os as - Sw hs os Se

3255 26,2166 13,7833 390 1 - eta - + ee Sic - Seats a. = ues

3562 24,6067 13,5833 391 69 =) ois Oo t6, - - - as Sik a Say wus

2448 31,925. 16,255 392 37 = 2) 2D) oe - - - - 9 - Shit au aS - 38

3172 27,1167 14,3 403 3 J. = ee - - ata os ee we es Lp Te 2 es ae

1694 34,75 18,3267 425 31 bs 1 - ch Ass - - 2 os 49 8 oe SS on es a:

3462 23,9583 13,1916 430 40 SAS Mey ge) Ss = ae oie Saas? See Bee : ae

2488 30,8416 15,4666 430 14 = = 1 - = 8 ia e 8 eo) *e AS Sea” a |

3179 27,0333 14,1 437 79 ae eed ie ee Bere = Ee ae bie ata ie,

2262 32,7917 16,8167 450 3 Shee a ane te See a tas sO aes ee,

2440 33,2416 17,15 450 18 eae In age Fae 2 8 BY ons of ce ME Bo a om

3577 31,3667 16,0833 453 %6 Se Se =e cS So Ie WME ae Bee ahi

2700 30.2167 14.85 469 13 ee ee) - - ee eae it a Se ee ae

2780 33,2416 17,25 475 42 a a) et i fe a ae mal Pees ae a,

3524 22,9333 12,9666 475 64 eee Gye oy) Ore ce Ea ae ee LE Se he ELAas = ee

301 35,0333 18,5333 500 eS - + eee 3 a0 Ws ci cc eA Be an ae

1698 34,7833 18,2167 502 10 oe vs es aE ae at 6 Le Sie es 7 ie

3225 25,1166 13,6 530 1 eoee Ba bss AP ole se" seas Sees SAS Bais Aohs

2879 29,4833 14,55 530 61 Re Sn ae - - - - =i a Bi 2 oy ws Ae ee

1697 34,7667 18,25 545 112 - - Sp ee == Ser aacig. st ae ne Sas

2785 33.225 17,45 560 12 Bh ey x be ees ss Ske oes aia ye lee ae 2 a

3845 20.5917 12,0833 566 1 a) Styne ei Ne one Save ay mind a tS Ba fs

3555 23,4433 13,03 590 19 eroeyS 2h Nas yo aes Dake Sorta ce be eres

3112 31,45 15,7667 648 i a Shy ie =. ae 2 es P Mg 1 Kee BS ae

3561 24,58 13,4167 655 9 oe aaae aE Silas Ae (= ayer ie mae ee . 2B

3458 24,1 13,2 725 16 re Oe a ee alte ata eae oe = ae

3346 31,12 15,575 730 23 >: ee ae ame Se awe Soy oe aes Ber.

2978 28,4 14,2 736 38 ane HS ane ere he le =, ree Ris:

3921 19,1366 11,1666 738 4 os eee ero aoa ee ae ee see Uh 983 ee

3885 17,5666 11,2833 779 10 ah as fy BE oi as = 6 ye aie men

3846 20,5833 11,8667 810 I Bade ee at SS Bre ante +S ee ee

3869 19,9167 11,3667 825 1 Pigs Epes Ea at cae a ae tags ae at

3113 31,3667 15,5833 840 16 Es Sy ee ae eR “i ae pe eS aes phat. a

3525 22,9333 12,8 850 14 oo ¢ Bite aes ane: Se ae SO eae Sons ih yi

3461 23,9583 13,0166 850 13 zo3 cay Je args ayer DS ae eke ee nee pe

3109 31,9 15,9 900 80 Sa Sf tke ~ 4 a eee ale Eee 2, |= =p

2697 30,2833 14,3167 940 12 Sere es aps eee ce he Deus aig te eae _ +

SHU IDEM TM OL Gs ES yi |

3341 32,6833 16,5167 945 36 teu Sse ee ane Se ae ies Dilek = i

Total valves 24059 6235 1946 5266 223 442 48 989 35 1 8 4 7 0 486 25 414 61 >

% Mode BOOS yl a 9 ral ee ae ee a Ym 2

”e Modern - 68,0 = fil = 6 = itp - 02 2 an) - 0,8 = 24

% Subtotal yi R

% Total ostracod fauna So er Se AR = 00h em 20s 30; = 24 7 22 = ‘

33,8 DOs - 20 - 42 - 0,0 . 1,7 2 26 5 24 = 19 E

Abbreviations: R = relict fauna, M = modern fauna

Kuiperiana

angulata

Palmoconcha?

walvisridgensis

]

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA

23 2 aS £2 2 i

S é es 2s 22 2 2 25

§5 g§ SS RS 288 2§s Sz Sign) se

Ss SE s& Or S55 S52 2s Zé Ss 3

ES 35 2s 88 286 £88 tS fs) 3S 2) ae

aia o igs ia a il S 3S =“ ‘al cd E &

M oR M OR MR M OR Me” R IO R® Mee Ree ne cmc

OEM oN kee ek. os 30 0 iw 0

a err SS ke EG

=i, 104 ck cere dame Mc urd . -S - - 4 0 4@ 97

eee ee 8 a ee ea ae me

ass) aoa ees. ss ee mice me SS aK)

a ee re eae SE lm yO ae

ea EE Se a ec

ne eee Se fe es sO Gi

= . = 3 & 5 & = 2 = = = - - - i 0 7 100

ee ee ee a eek els no ae) MB FB

5, MRO) Se ee ae 20 ee os ay ae 2 = 6 OW WU

5 5 = Z s 5 = = < = = = - - - 8 0 8 100

ee ee ee eS ee ec GH. 35 MD

a oo Crees eee kl OD SS

= a 4 < 2 < = = : = - = - - - 8 2 10 83

Se ee ee ee eS Sse OG NR

a bess kes, = = OM 6) &

2 = = S = = = = = = > © - - - 29 0 29° «78

= 1 3 ee 1 E ef is - - - = = 4 3 100

SE ah ae a en aN fe ie vio e ia Key e ee Ss At Rl

ee ee Serene Sra As lO | 2h

= S s = = 2 = 2 = : = - - - - 1 0 ih 7g

ee ee en eS ee SG gk gn

ee ees eee A ee ee oa ik Se

ee emer ENE gi st ke ss i CR Oe

_ s & = = 5 S S = = = = - - - 36 0 360 «(37

ee le AO

ee ce ae ai SR ae ee cots ey hy

i 43 G S z & s 3 4 = = = = = - 49 2 Sil 7h}

e = 5 = a = 2 = = = - - - - - 6 0 6 8

5 s as = 3 = = E: = 2 2 - - - - 6 0 6 60

au) asia St eel Re Cree tr clog sc ee Soe alt Oe a tiet00

Se ante Suet yhte cs eee Lome eos. (6 Re GA, Ga 10

iii Sie C20 Moe a) Bence ee eee Set A Bol OR Oe 28. 20

ee ee ee eS ee GR eG

BE Raape Sno ay eee Pe eee. Sac Oe eepteetOD

Seco S eteiee SciCeS Se er TE SOE Sat Caen Om

a = = ~ = = > = s = - - - - - 10 0 10 90

; Et Ca es aaa Ee te aeA ee Buy os = = %9) <0) 91 100)

z eae es ee Se zo wy, Pee =.) tet | ES Se RO mea

5 2 P = & = = é = = = - - - - 13 0 18036

ee a ee Se AS, og a i 2° Sa

a cs “, = 3 2 2 = > = é > - - - 1 1 2 50

Pre) oe ee eS eee ee DS iy OS. ay

ty > ee ee eM pre Nee Se =, = ceo OMRMmTeRTOO!

- Ss i s 2 = = a 5 = = = - - - 1 0 1 100

= Sal SME I ed ae 2) A) Oe =) if O. 12.16

= = 5 = - = = = = = - - - - - Bl 2 13° 92

« é re x a = = s = 2 - - - - - 8 2 10 76

: 20 ee EE ed cing 3 tediy Vass” | ee are wm aiime hoes)

S cf SN re Paes ee ee ees, oe SG) 10 Oh 9B

i any wt «Rot rs a 6 =) RE) es oe Gam

: | Ope le ee ae = 0 CG

16 396 27 695 7 490 57 993 318 43 0 436 64 18 2 18046 2861 20907 87

= 2 - 38 Poi > 5,5 - 0,2 = 24 - 0,0

0,5 - 09 - 0,2 - 1,9 - 11,1 = 0 = 22) - 0,0

- 2,0 : 3,3 - 26 ) 6,2 = 02 - 23 - 0,0 -

ive SPOR yy Oe OE = A = Oe

12 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 3

Taxa described in this paper, and their abundance as a percentage of the fauna of the 183 samples

in which they occur.

Percentage of fauna

Total Modern Relict

Pseudokeijella lepralioides (Brady) 39,0 67,9 34,5

Ruggieria' cytheropteroides (Brady) 26,2 Well 223).

Bensonia Sp. 6,4 lpia 5,6

Bensonia knysnaensis knysnaensis (Benson & Maddocks) 6,2 UI 5,4

Bensonia knysnaensis robusta subsp. nov. 0,2 0 0,2

Ambostracon spp. WZ 3,0 7,9

Ambostracon keeleri sp. nov. 4,8 1,6 5,4

Ambostracon flabellicostata (Brady) Pd) ee 2,4

Ambostracon levetzovi (Klie) 0,1 0,4 0,1

Henryhowella melobesioides (Brady) 2,0 0,4 Dyld

Cytherella spp. 353 ell 5,9

Cytherella dromedaria Brady 353) 0,2 3,8

Cytherella namibensis sp. nov. 2,0 0,9 DJ

Paracypris lacrimata sp. nov. 26) Le) Dee

Doratocythere exilis (Brady) 3,0 0 aD)

Neocytherideis boomeri sp. nov. 2,4 0,9 16

Xestoleberis spp. 2,4 253 2,4

Xestoleberis africana Brady D3} ey, 2,4

Xestoleberis hartmanni sp. nov. 0,1 0,1 0,1

Loxoconchidae Dell 2,9 2,8

Palmoconcha walvisbaiensis (Hartmann) 2,3 Pog Mt D3}

Palmoconcha? walvisridgensis sp. nov. 0,1 0,5 0,1

Kuiperiana angulata sp. nov. 0,2 0,2 0,3

Palmoconcha subrhomboidea (Brady) 0,1 0,1 0,1

100

mean 74%

mean 66%

50 T Sir

0 200 400 600 800 1000m

Fig. 2. Distribution with depth across the continental margin of south-western Africa of the combined

abundances of the dominant species (5-point running mean of percentage of total ostracod fauna).

Mean values for various shelf zones are shown by horizontal lines. The faunas described herein are

seen to be primarily typical of the continental shelf (60-400 m water depths).

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 13

range 400-800 m. In summary, the species dealt with in this paper are dominantly a

mid-shelf to outermost shelf assemblage (60—400 m).

Using criteria based on shell preservation, all individual specimens have been

classified into one of two categories: ‘modern’ (i.e. living, or dead but well preserved,

translucent valves) and ‘relict’ (opaque valves that are abraded and/or corroded). In

terms of overall population abundance, the rankings of the dominant taxa are similar

for the two categories (Fig. 3), the main difference being the relatively greater import-

ance of Bensonia knysnaensis in the modern fauna (2nd) compared to the relict

populations (Sth). However, the dominance of Pseudokeijella lepralioides is much

greater in the modern (67,9%) than in the relict fauna (34,5%), and contrasts with a

RELICT

—

BK DE Ey eA NB 2G” iM

rE RG A C Be OE Ea PAL NB X HM

Fig. 3. Rankings of the dominant taxa in the total relict and modern ostracod

assemblages from the continental margin of south-western Africa (shown as a per-

centage of the fauna of the samples in which they occur). Abbreviations: PL—

Pseudokeijella lepralioides, RC—Ruggieria cytheropteroides, A—Ambostracon

(A.) spp., C—Cytherella spp., BK—Bensonia knysnaensis, DE—Doratocythere

exilis, _L—loxoconchids, PAL—Paracypris lacrimata, NB—WNeocytherideis

boomeri, X—Xestoleberis spp., HM—Henryhowella melobesioides.

14 ANNALS OF THE SOUTH AFRICAN MUSEUM

HENRYHOWELLA MELOBESIOIDES (0,4%)

20 30

NEOCYTHERIDEIS BOOMER]! (0,9%) | |

' ! | 1

20 30

| CYTHERELLA 2 SPP. (1,1%)

20 30

PARACYPRIS LACRIMATA (1,9%)

20 30

XESTOLEBERIS 2SPP. (2,4%) | |

| | | |

LOXOCONCHIDS 4 SPP. (2,9%)

20 30

AMBOSTRACON (A.) 3 SPP. (3,0%)

RUGGIERIA

CYTHEROPTEROIDES (7,7%)

Trace Siocon! amen came amen amen imma Somme Gnemen ise) ona anen Uaneee me

20 30

| BENSONIA KNYSNAENSIS (11,1%)

Ss}

0 SU

20 30

7 PSEUDOKEIJELLA

LEPRALIOIDES (67,9%)

20 30

| 1 od

WR WB L, OR S CP

Fig. 4. Latitudinal abundances of species

modern populations expressed as percentages of

the modern fauna, arranged in descending order of

overall abundance (in parenthesis). Data are

double 5-point running means. Vertical scale

percentage, horizontal scale = degrees of latitude.

Abbreviatons: WR—north edge of Walvis Ridge

abutment shelf; WB—Walvis Bay; L—Lideritz;

OR—Orange River; S—Saldanha; CP—Cape

Peninsula.

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA

| HENRYHOWELLA MELOBESIOIDES (2,2%)

20 30

| XESTOLEBERIS 2 SPP. (2,5%) | |

eel

5 20 30

NEOCYTHERIDEIS BOOMER! (2,6%) | |

20 30

_| PARACYPRIS LACRIMATA

(2,7%)

20 30

LOXOCONCHIDS 4 SPP. (2,8%)

20 30

| PSEUDOKEIJELLA

LEPRALIOIDES (34,5%)

41 I

Fig. 5. Latitudinal abundances of species in relict

populations expressed as percentage of the whole

fauna; arranged in descending order of overall

abundance a parenthesis). Data are double

5-point running means. Vertical scale = percent-

age, horizontal scale = degrees of latitude. Abbrevi-

ations: WR—north edge of Walvis Ridge abutment

shelf; WB— Walvis Bay; L—Lideritz; OR— Orange

River; S—Saldanha; CP—Cape Peninsula.

16 ANNALS OF THE SOUTH AFRICAN MUSEUM

corresponding diminution in importance in Ruggieria cytheropteroides (7,7% and

29,1%, respectively). In addition, Doratocythere exilis (6th ranking in relict popu-

lations), has not been identified in the modern fauna. Invariably, the regional

distribution of the modern and relict populations of each species show differences

(Figs 4, 5), details of which will be presented for individual species in the taxonomic

notes, with a brief summary in the discussion section at the end of the paper.

Sediment samples used in this study were retrieved from the top 10 cm of the sea-

floor using a Van Veen grab. They were collected from the University of Cape Town’s

research vessel “Thomas B. Davie’ (sample numbers have a TBD prefix), and the orig-

inal samples are retained in the store of the Marine Geoscience Unit, University of

Cape Town.

Microfossils were picked from 125-micron and larger sieve fractions, and num-

bered specimens are lodged in the micropalaeontology collections at the South

African Museum, Cape Town, under the catalogue prefix SAM—PQ-MF.

LIST OF GENERA AND SPECIES

The genera and species of Ostracoda discussed in this work are given below:

PAGE

Cytherella Jones, 1849 ° 2.00. 6s. pcs a ek Pe ee 7/

Cytherella dromedaria Brady, 1880

Cytherella namibensis sp. nov.

Paracypris: Sats, 1866.6 (25 sks gee so ee UD

Paracypris lacrimata sp. nov.

Doratocythere McKenzie; 1967. 9o400..2 pea. oe a ee eee 29

Doratocythere exilis (Brady, 1880)

Bensonia Rossi de Garcia, 1969 . : os. on ee ee Sy

Bensonia knysnaensis knysnaensis (Benson & Maddocks, 1964)

Bensonia knysnaensis robusta subsp. nov.

Neocytherideis Puri) 1952 o.oo Cece oa ee, es eee 35

Neocytherideis boomeri sp. nov.

Ambostracon Hazel, 1962 niin bs Ge os ob eh oe ee eee 42

Ambostracon (Ambostracon) flabellicostata (Brady, 1880)

Ambostracon (Ambostracon) levetzovi (Klie, 1940)

Ambostracon (Ambostracon) keeleri sp. nov.

Palmoconcha Swain. & Gilby: 1974... ee ee eee 54

Palmoconcha walvisbaiensis (Hartmann, 1974)

Palmoconcha? walvisridgensis sp. nov.

Palmoconcha subrhomboidea (Brady, 1880)

Kuiperiana Bassiouni; 1962: oct ee oe Be et ee 61

Kuiperiana angulata sp. nov.

Ruggierta Keij: V9 7 i205 don sea ey ep en eee Se Ee 63

Ruggieria cytheropteroides (Brady, 1880)

Henryhowella Puri, 19572 sai) .00 es os ae ee ee ee 68

Henryhowella melobesioides (Brady, 1869)

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 17)

FESCTGORCUIC AWC MMONE te, ieete es eran aries wae yinlatali and oi Earoin aa 6 lela aly a eles Vi

Pseudokeijella lepralioides (Brady, 1880)

KEBAQDETES, SBIB, WS SOM ie A OL MRI A cools) Het a ane ee a a Vi

Xestoleberis africana Brady, 1880

Xestoleberis hartmanni sp. nov.

SYSTEMATIC DESCRIPTIONS

The classification used here is based on (Moore 1961), with various additions

necessitated by subsequent work.

Abbreviations used: AM = anterior margin; ATE = anterior terminal element; C

= carapace; CA = cardinal angle; DM = dorsal margin; LV = left valve; MA = mar-

ginal area; ME = median element; MPC = marginal pore canal; MS = muscle scars;

NPC = normal pore canal; PM = posterior margin; PTE = posterior terminal

element; RV = right valve; SCT = subcentral tubercle; VM = ventral margin.

In the discussion of ostracod distributions, UDL, and LDL = upper depth limit,

and lower depth limit, respectively.

Class CRUSTACEA Pennant, 1777

Subclass OSTRACODA Latreille, 1806

Order PopocoripA Muller, 1894

Suborder PLATYCOPINA Sars, 1866

Family Cytherellidae Sars, 1866

Genus Cytherella Jones, 1849

| At least twelve species of Cytherella have been reported from the late Cenozoic

of southern Africa, seven from the Quaternary. Four of these occur on the continental

shelf of south-western and southern Africa: Cytherella dromedaria Brady, 1880—west

coast shelf; Cytherella namibensis sp. nov.—west coast shelf; Cytherella aff. C. punc-

tata Brady, 1866 (Benson & Maddocks 1964)—Knysna Lagoon; and Cytherella

omatsolai Hartmann, 1974—coastal zone, Luanda to Sandwich Harbour.

Cytherella dromedaria Brady, 1880

Fig. bA—D

Cytherella dromedaria Brady, 1880: 173, pl. 43 (figs 6a—b). Puri & Hulings, 1976: 312, pl. 24 (fig. 14).

Cytherella sp. aff. C. cuneiformis Hartmann, 1974. Keeler, 1981: 185-187, pl. 11 (figs 1-3).

Illustrated material.

SAM-PO-MEF-0509, LV, TBD 6824, 90 m

SAM-PQ-MEF-0510, RV, TBD 6824, 90 m

SAM-POQ-MF-0511, RV, TBD 6824, 90 m

SAM-PQ-MF-0512, C, TBD 6824, 90 m

Material

702 valves.

18 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 6. A-D. Cytherella dromedaria Brady, 1880, TBD 6824, 90m. A. SAM—PQ-MEF0509, LV

SEM 2575. B. SAM—PQ-MF0510, RV, SEM 2573. C. SAM-—POQ-MF0511, RV, SEM 2577.

D. SAM-—PQ-MF0512, carapace, dorsal view, SEM 2582. E-F. Cytherella namibensis sp. nov.

TBD 2975, 180 m. E. SAM—PQ-MF0513, holotype, LV, SEM 2643. F. SAM—PQ-MF0514, RV,

SEM 2640. Scale bars = 100 microns.

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 19

Remarks

This distinctive species has a strongly gibbous DM, and numerous large normal

pore openings on the lateral surface. The AM in each valve has a very narrow rim that

is clearly seen in dorsal view.

15 longitude 20 95

latitude

Ga ,

ele Ns

C. dromedaria

Fig. 7. Distribution of Cytherella spp. Squares—C. namibensis sp. nov.

(southern limit shown as dotted line south-east of the Cape Peninsula);

crosses—C. dromedaria (Brady) (seaward limit shown by solid line); dia-

monds—C. serratula (Brady) ianaivatd limit shown by dashed line, after

Dingle et al. 1990). Abbreviations for this and subsequent distribution

maps: K—Kunene River; WR—Walvis Ridge abutment shelf; WB—

Walvis Bay; L—Liideritz; OR—Orange River; S—Saldanha; CP—Cape

Peninsula.

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

latitude °S

0 200 400 600 800m

= dromedaria + ‘punctate’ 2 ‘smooth’

Fig. 8. Latitudinal water-depth distribution of sites with Cytherella dromedaria and

C. namibensis sp. nov. Field outlines: solid line = ‘punctate’ variety of C. namiben-

sis; dashed line = ‘smooth’ variety of C. namibensis.

Distribution

Brady (1880) originally recorded this species from ‘Challenger’ Station 104 in

False Bay. The present data indicate that it is confined to the west and south coast

continental margins south of 29,5°S (Fig. 7).

Modern specimens were recovered from two sites: west of Hout Bay (94 m) and

west of Cape Agulhas (73 m).

Relict populations are confined to two areas. The northernmost lies on the middle

to outer shelf (170-300 m—Fig. 8) between the Orange and Olifants rivers, where

abundances are generally low (>5% at one of four sites). A southern population

(central Cape Peninsula to the eastern Agulhas Bank) has UDL and LDL of

40—220 m, and includes the two modern sites. Abundances in the southern sector are

relatively high (>5% at four of eleven sites), and the preferred water depth of the

relict populations of Cytherella dromedaria was c. 100 m (Fig. 11).

Cytherella namibensis sp. nov.

Figs 6E-F, 9A—-D, 10A

?Cytherella sordida Miller, 1894. Bold, 1966: 158-159, pl. 1 (fig. 10).

?Cytherella aff. C. sordida Miiller, 1894. Dingle, 1976: 39, fig. 12 (39).

Cytherella spp. Boomer, 1985: 12-13, pl. 1 (figs 16-17).

Derivation of name

From the type locality of the species adjacent to the Namib Desert, south-western

Africa.

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 2h

Fig. 9. A-D. Cytherella namibensis sp. nov. A-C. TBD 2971, 162 m. A. SAM-PQ-MF0515, LV,

SEM 2648. B. SAM-PQ-MF0516, RV, SEM 2647. C. SAM-PQ-MF0517, LV, SEM 2650.

D. SAM-PQ-MF0518, RV, TBD 2975, 180 m, SEM 2645. E-F. Paracypris lacrimata sp. nov.

TBD 6846, 95 m. E. SAM—PQ-—MF0519, holotype, LV, SEM 3047. F. SAM—PQ-MF0520, LV

SEM 3045. Scale bars = 100 microns.

DD, ANNALS OF THE SOUTH AFRICAN MUSEUM

Holotype

length height

SAM-PO-MEF-0513, LV, TBD 2975, 180 m 0,68 mm 0,39 mm

Paratypes

length height

SAM-—PQ-MEF-0514, RV, TBD 2975, 180 m 0,70 mm 0,42 mm

SAM-POQ-MEF-0515, LV, TBD 2971, 162 m 0,75 mm 0,42 mm

SAM-PO-MF-0516, RV, TBD 2671, 162 m 0,93 mm 0,60 mm

SAM-—PQ-MEF-0517, LV, TBD 2971, 162 m 0,80 mm 0,48 mm

SAM-—PQ-MEF-0518, RV, TBD 2975, 180 m 0,72 mm 0,50 mm

Material

423 valves.

Diagnosis

Punctate species of Cytherella with greatest height in anterior third. AM outline

has a bulbous appearance, posterodorsal outline slopes steeply posteriorly, DM is

straight, but the dorsal outline in lateral view is distinctly concave. Posteroventral area

of valve is inflated, and has a curved, angular outline.

Description

The AM is broadly and symmetrically rounded. Bulbous in lateral view because

the greatest height lies in the anterior third of the valve. PM outlines differ: in LV the

outline is asymmetric, with a distinctive posterodorsal slope, whereas in RV the

outline is more symmetric, although there is slight acumination along the line of great-

est length. In both valves the DM and VM are essentially straight, but the outline of

the former in lateral view is distinctly concave around mid-length. In both valves there

is a small but distinctive posteroventral angularity, that in RV forms a slight angular

projection.

Surface ornamentation punctate, it grades from punctate overall, to specimens in

which the main punctation is confined to the PM, with rare, scattered puncta in the

anterior half of the valve.

Internal features are typical for the genus.

Remarks

The end members of the ‘punctate’ and ‘smooth’ series are easily distinguished,

and overall their geographical distributions are distinct. It is tempting to split the

group into two species but, in the case of numerous individuals, the distinction would

be arbitrary.

Brady (1880) did not record Cytherella from ‘Challenger’ Station 142 on the con-

tinental shelf off the Cape of Good Hope, but illustrated several punctate specimens

from widely separated localities under the name C. punctata Brady, 1866. The types

of the latter are from the eastern Mediterranean and have a distinct dorsomedian

depression, and have a straighter DM than C. namibensis. Of the specimens allocated

to C. punctata in the ‘Challenger’ report (Brady 1880), those from Port Jackson (Aus-

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 23

tralia) are closest to the types, whereas all others all have a distinct reniform outline.

In the latter category he presumably included material from Tristan da Cunha.

Benson & Maddocks (1964) recorded living punctate specimens of a species refer-

red to Cytherella aff. C. punctata Brady, 1866, from the Knysna Lagoon. These differ

significantly in lateral outline from C. namibensis. Their highest point is in the poste-

rior half and they have relatively narrow AM outlines and a straighter DM. Similarly,

Hartmann’s (1974) species C. cuneiformis (coastal Angola) has a strikingly different

DM outline to my species (Fig. 10).

In their survey off the River Congo estuary (R. Zaire, 11 m to at least 50 m water

depth), Babinot & Kouyoumontzakis (1986) found Cytherella sp. aff. C. punctata

Brady to be the most abundant ostracod taxon. From their clear illustrations it is

confidently concluded that they were not dealing with C. namibensis because the

highest point in the valve outline lies in the posterior part.

Two previously reported Tertiary species from the region are close to C. namui-

bensis. These are C. sordida Miller (Bold 1966) from Gabon, and Cytherella aff.

C. sordida Miller (Dingle 1976). Both are punctate, and differ only slightly in lateral

outline from C. namibensis, with which they may be conspecific.

Distribution

Cytherella namibensis sp. nov. occurs along the entire south-west African conti-

nental shelf (Fig. 7), but usually there is a subdivision into areas populated by the

‘punctate’ and ‘smooth’ varieties.

Modern specimens occur at locations scattered along the shelf from 17,5°S to 28°S

(Orange River), where they occupy a depth range 115-295 m. The main cluster lies

on the Orange Shelf. Both ‘punctate’ and ‘smooth’ varieties occur here.

Relict specimens occur from 17,5°S to the Cape Peninsula, but Keeler (1981) did

not find the species on the eastern Agulhas Bank. Almost invariably, the ‘punctate’

A B

Fig. 10. Outlines of LV of Cytherella species. A. C. namibensis sp. nov.,

SAM-POQ-MF0513, holotype, TBD 2975, 180 m. B. Cytherella aff. C. punc-

tata Brady, 1866, Benson & Maddocks 1964, Knysna estuary, traced from

Benson & Maddocks (1964, pl. 1 (fig. 2)). C. C. cuneiformis Hartmann, 1974,

Angola, traced from Hartmann (1974, pl. 13 (fig. 104)). D. C. omatsolai Hart-

mann, 1974, Angola, traced from Hartmann (1974, pl. 11 (fig. 82)).

Scale bars = 200 microns; other scales not known.

24 ANNALS OF THE SOUTH AFRICAN MUSEUM

variety of the species occurs at shallower water depths than the ‘smooth’ variety. The

‘punctate’ variety also predominates off the Cape Peninsula; only two sites south of

31°S yielded the ‘smooth’ variety. The depth-related partition between the two var-

ieties is well-illustrated in Figure 8. This shows that the ‘punctate’ type has UDL and

LDL between 115 m and 280 m off the Walvis—Orange sector and between 120 m and

345 m off the south-western Cape, in contrast to a 130-590 m range for the ‘smooth’

variety. (The two sites at 535 m and 736m, with ‘punctate’ and ‘smooth’ types,

respectively, are considered allochthonous.) The depth-related preferences of the two

morphotypes is further emphasized when abundances are plotted against depth

(Fig. 11). The ‘punctate’ variety peaks at about 220 m, and the ‘smooth’ variety at

about 380 m. Presumably this distribution is linked to depth-related environmental

factors.

General remarks on the distribution of the genus Cytherella

Three species are widely distributed on the continental margin of south-western

Africa: C. dromedaria occurs on the continental shelf south of 28°S, C. namibensis

occurs on the continental shelf between 17,5°S and the Cape Peninsula (34°S), and

C. serratula (Brady, 1880) occurs on the continental slope in water depths >1 000 m.

Less widely distributed are two coastal species, C. omatsolai (as far south as Sandwich

Bay, 40 km south of Walvis Bay), and Cytherella aff. C. punctata (Knysna Lagoon),

and the deep-water taxon Cytherella sp. 3027 (Dingle et al. 1990—2 916 m).

The shelf species C. dromedaria and C. namibensis both have restricted modern

distributions: the former is confined to the Cape Peninsula area, and the latter to

areas north of 28°S. In relation to the abundances of other taxa, the modern popu-

lation of C. dromedaria is insignificant but, between the Orange River and Lideritz,

and north of the Walvis Ridge abutment shelf, C. namibensis is an important compo-

nent of the modern fauna (Fig. 4). This distribution suggests that C. dromedaria and

100

0 200 400 600 800m

= dromedaria + ‘punctate’ « ‘smooth’

Fig. 11. Abundance of Cytherella dromedaria and C. namibensis sp. nov. as percentage

of ostracod fauna plotted against water depth. Field outlines: solid line = C. dromedaria,

dashed line = ‘punctate’ variety of C. namibensis, dot-dashed line = ‘smooth’ variety of

C. namibensis.

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA DS

C. namibensis can be considered characteristic of shallow, warm, Agulhas water, and

cold west-coast mid-shelf conditions, respectively.

The range of both species is extended in the relict faunas; C. dromedaria to the

north, and C. namibensis to the south, although this was mainly in the ‘punctate’

variety, exhibiting the tendency of this morphotype to favour inner shelf areas. Con-

sidering their relative abundance by latitude (Fig. 5), C. dromedaria locally forms an

important component of the relict population in the Cape Peninsula—Saldanha Bay

area, whereas C. namibensis is either the dominant, or one of the dominant species, in

the relict ostracod populations over a wide sector of the continental shelf from north

of the Walvis Ridge abutment to between Walvis Bay and Luderitz. South of this it is

generally a minor component.

There is a sharp break between the distribution of the neritic (shelf) species and

the bathyal C. serratula. This has been attributed to the environmental barrier formed

by the core of the salinity minimum zone in the Antarctic Intermediate Water at about

750 m (Dingle et al. 1989, 1990, fig. 3).

Superfamily CyPRIDACEA Baird, 1845

Family Paracyprididae Sars, 1923

Genus Paracypris Sars, 1866

In the open literature, only one Quaternary species of this genus has previously

been recorded from southern Africa: Paracypris westfordensis Benson & Maddocks,

1964, from Knysna Lagoon. No records were made by Brady (1880), Miller (1908),

Klie (1940), and Hartmann (1974) in their studies of the region, nor was it recorded

by Bold (1966) in the Neogene of Gabon.

Fossil records of the genus in southern Africa have been made from the Creta-

ceous (e.g. Dingle 1981) and Tertiary (Dingle 1976).

Paracypris lacrimata sp. nov.

Figs 9E—-F, 12C, F

Paracypris sp. aff. P. polita Sars, 1866. Keeler, 1981: 34-35, pl. 1 (fig. 14).

Paracypris sp. Keeler, 1981: 35-36, pl. 1 (fig. 15).

Pontocypris sp. Boomer, 1985: 16-17, pl. 4 (fig. 64).

Derivation of name

Lacrima—Latin, tear. Refers to tear-shape of valve.

Holotype

length height

SAM-POQ-MEF-0519, LV, TBD 6846, 95 m 1,24 mm 0,50 mm

Paratypes

length height

SAM-PQ-MF-0520, LV, TBD 6846, 95 m 1,20 mm 0,50 mm

SAM-PO-MF-0521, RV, TBD 6846, 95 m 1,31 mm 0,48 mm

SAM-—POQ-MF-0522, LV fragment, TBD 6846, 95 m

26 ANNALS OF THE SOUTH AFRICAN MUSEUM

D E F

ilies

= & atte

o4% se ==

Fig. 12. A-F. Paracypris. A, D. P. polita Sars, internal view and MS, LV, north-western

Europe, traced from Sars (1928, pl. 31). B, E. P. bradyi McKenzie, internal view and

MS, RV, south-eastern Australia, traced from McKenzie (1967, fig. 2d). C, F. P. lacri-

mata sp. nov., TBD 6846, 95 m. C. SAM-—PQ-MF0521, internal view, RV.

F. SAM-PQ-MF0522, MS, LV. Scale bars: A-C = 200 microns; D-F = 100 microns.

Material

547 valves.

Diagnosis

Species of Paracypris with long, asymmetrical, pointed PM whose apex is strongly

ventrally directed. DM strongly convex and VM concave. Highest point lies at about

one-quarter valve length. There are c. 15 branched anterior radial pore canals, and a

MS pattern in which the four anterior-most scars are longitudinally aligned.

Description

External features. Broadly rounded AM that is somewhat ventrally directed, and

a strongly acuminate PM that is also ventrally directed. DM strongly arched, with the

highest point at about one-quarter valve length. VM concave. Overall the outline is

tear-drop or comma shaped. RV outline has an anterodorsal step.

Internal features. Wide anterior and posterior vestibulae with c. 15 simple and

branched anterior radial canals that are larger and more complex anteroventrally. The

MS consist of five scars, all elongate, with the four anterior-most aligned longitudi-

nally. The hinge is adont.

Remarks

Many of the penultimate instars have a reddish-brown ?chitinous lining to the

outer lamella, which gives the valves a reddish hue seen externally. Also, because

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA Dil

these instars appear to be particularly susceptible to decalcification in deeper waters,

the presence of the species is recorded as residual brown ‘husks’. Sars (1923: 70) in his

re-appraisal of his genus mentions that the type species P. polita has a reddish brown

hue along the ventral side of the valves and that the marginal areas are ‘highly chiti-

nized’.

Paracypris lacrimata sp. nov. is very close in both outline and internal features to

the holotype P. polita Sars, 1866. The new species is slightly more arched along the

DM and concave on the VM, giving it a more pronounced tear-drop or comma-like

appearance. The definitive differences are in the greater asymmetry and complexity of

the AM radial pore canals of the new species, and differences in MS pattern (see

15 longitude 20 25

Dal sp \

xe) \

= + |

& YL

mre |)

ee “OR

30 nf \

O ‘

+ H+ =

Ba me }

wea ?

modern CP

1 a ee

Fig. 13. Distribution of Paracypris lacrimata sp. nov. Squares = sites with

calcareous valves; crosses = sites with red/brown husks. Dotted line is the

seaward limit of modern populations. See Fig. 7 for abbreviations.

28 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 12). Paracypris lacrimata lacks the prominent eye spot described by Sars (1923)

for P. polita.

A further close relative is P. bradyi McKenzie, 1967, from south-east Australia.

This species has more numerous AM radial pore canals and a different outline in both

the AM and PM inner lamella. There is a slight difference in MS disposition between

the two species (Fig. 12).

Although it occurs geographically close by, P. westfordensis Benson & Mad-

docks, 1964, from the Knysna Lagoon is easily distinguished by being less acuminate

posteriorly, and having a less broadly rounded AM outline.

Distribution

Specimens of P. lacrimata occur along the continental shelf from 19°S (Walvis

Ridge shelf) to the south-western Cape and across to the eastern Agulhas Bank

(Fig. 13).

Modern populations all lie in a depth range 15—133 m between Saldanha Bay and

Cape Agulhas.

Relict distribution is complicated by post-mortem decalcification (?by oxygen-

depleted waters). These populations extend as far north as the Walvis Ridge but, with

two exceptions, all the specimens north of 33°S consist of chitinous husks. These pre-

sumably represent decalcified material, because the two calcareous valves mentioned

are in a poor state of preservation. Off the south-western Cape, relict populations

consist of calcified material, with only a small proportion of chitinous ‘valves’. The

depth ranges over which these relict populations are found varies with latitude

(Fig. 14). North of 27°S, the UDL is >200 m (with the exception of the calcite valve

from 24°S at 161 m), but farther south, the UDL of the chitinous material is c. 150 m,

and off the south-west the shallowest depth with ‘husks’ is 94 m. The LDL increases

latitude °S

calcareous valves

0 500 1000 m

Fig. 14. Latitudinal water-depth distribution of sites with Paracypris lacrimata sp. nov.

Dashed line shows LDL of specimens with calcareous valves.

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 29

northwards, from 450 m in the south, to 590 m near Walvis Bay. The three records of

single specimens at >800 m are probably allochthonous because they are so much

farther downslope.

Superfamily CYTHERACEA Baird, 1850

Family Campylocytherididae Puri, 1960

Subfamily Campylocytheridinae Puri, 1960

Genus Doratocythere McKenzie, 1967

McKenzie (1967) erected this genus for certain species that occur at inshore loca-

tions along the coast of southern Australia. It has since been reported from Japan

(Ishizaki & Matoba 1985), but the illustrations in this publication do not appear very

close to the types illustrated by McKenzie (1967). Kempf (1988) cited no additional

records of the genus, so currently available records probably confine the genus to the

Southern Hemisphere.

Doratocythere exilis (Brady, 1880)

Fig. ISA-F

Cythere exilis Brady, 1880: 69, pl. 16 (figs Sa—h). Puri & Hulings, 1976: 276, pl. 10 (figs 1-11).

Doratocythere exilis (Brady, 1880) Keeler, 1981: 39-41, pl. 1 (figs 20-22).

Reymentia exilis (Brady, 1880) Boomer, 1985: 49-S0, pl. 2 (fig. 21).

Illustrated material

SAM-—PQ-MF-0523, LV, TBD 2975, 180 m

SAM-—PQ-MF-0524, RV, TBD 2975, 180 m

SAM-PO-MF-0525, LV, TBD 2975, 180 m

SAM-POQ-MF-0526, RV, TBD 2975, 180 m

SAM-—PQ-MF-0527, C, TBD 6824, 90 m

Material

673 valves.

Remarks

Brady’s (1880) species accords well with McKenzie’s (1967) descriptions and illus-

trations of the type species (D. foveata). There are three points of difference:

1. In D. exilis there is no appreciable thickening of the valve wall in the vicinity of the

NIB

2. The small ‘micropunctate interscar’ area of D. foveata could not be identified in

D. exilis, although in well-preserved specimens a small spot lies in this position.

3. Doratocythere exilis has a prominent fulcral point anterodorsal to the MS. No

similar feature was mentioned in the species described by McKenzie.

Of these points, the first appears to be the most significant because McKenzie

(1967) emphasized its presence as generically diagnostic. However, we do not consider

the difference important enough to warrant the erection of a new genus for the South

African taxon.

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 15. A-F. Doratocythere exilis (Brady, 1880). A-D. TBD 2975, 180 m. A. SAM—PQ—MF0523,

LV, SEM 2476. B. SAM—POQ-MF0524, RV, SEM 2478. C. SAM-—PQ-MEF0525, LV, SEM 2489.

D. SAM-—PQ-MF0526, RV, SEM 2480. E-F. SAM-—PQ-MF0527, carapace, TBD 6824, 90 m.

E. Dorsal view, SEM 2494. F. Detail of ornamentation, dorsal surface, mid-length, LV.

Scale bars: A-E = 100 microns; F = 10 microns.

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 31

SEM photographs of D. exilis reveal that the crests of the prominent longitudinal

ribs consist of a narrow, smooth cord, and that the flanks and non-punctate intercostal

areas have a delicate secondary reticulation.

Distribution

No modern specimens of D. exilis were recovered.

Relict populations occur on the continental shelf from 22°S to south-western

Cape, and Keeler (1981) reported the species on the eastern Agulhas Bank (Fig. 16).

The main distribution lies between 28° and 31,5°S (Orange—-Namaqualand

shelves), where, in several samples, the species constitutes >10 per cent of the total

ostracod population. The depth range in this area is 158-305 m (Fig. 17), with the

greatest abundances between 155 m and 200 m.

15 longitude 20 25

latitude

Fig. 16. Distribution of Doratocythere exilis (Brady). See Fig. 7 for

abbreviations.

32 ANNALS OF THE SOUTH AFRICAN MUSEUM

latitude °S

100 200 300 m

Fig. 17. Latitudinal water-depth distribution of sites with Doratocythere exilis. Horizontal

lines delimit across-shelf populations.

A barren zone occurs between the mid-Namaqualand shelf and Saldanha, to the

south of which the south-western Cape population occurs in a depth range 40—227 m.

Although in this area the UDL is much shallower than farther north, the greatest

abundances occur in a similar depth range to the Orange—Namaqualand population.

This centre may be continuous with that on the eastern Agulhas Bank, which is within

the same depth range.

A single, worn valve occurred anomalously at 22,25°S at 223 m.

Comparing the latitudinal abundance of D. exilis with the other dominant taxa

(Fig. 5), shows that it is never more than a secondary component of the overall

ostracod populations, although locally west of the Cape Peninsula it is the fourth most

abundant taxon. In addition, together with Ambostracon spp., D. exilis is a character-

istic element of the northern part of the Namaqualand shelf fauna.

Family Cytherettidae Triebel, 1952

Subfamily Cytherettinae Howe, 1961

Genus Bensonia Rossi de Garcia, 1969

This genus appears to be confined to the Atlantic Ocean. It probably first

appeared in southern Africa in the Upper Eocene.

Bensonia knysnaensis knysnaensis (Benson & Maddocks, 1964)

Fig. 1SA—F

Cytheretta knysnaensis Benson & Maddocks, 1964: 22-23, text-figs 11-12, pl. 2 (figs 7-11).

Bensonia knysnaensis (Benson & Maddocks, 1964) Keeler, 1981: 43-45, pl. 2 (figs 2-4).

Cytheretta sp. Boomer, 1985: 24-25, pl. 3 (fig. 44).

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 33

Fig. 18. A-F. Bensonia knysnaensis knysnaensis (Benson & Maddocks, 1964), TBD 2224, 58 m.

A. SAM-PQ-MF0528, RV, SEM 2455. B. SAM—POQ-MF0529, LV, SEM 2452. C. SAM-—PQ-MF0530,

RV, SEM 2462. D. SAM-PQ-—MF0531, LV, SEM 2459. E. SAM-PQ-MF0535, carapace, dorsal

view, SEM 2456. F. SAM—POQ-—MF0530, RV, MS, SEM 2464. Scale bars = 100 microns.

34 ANNALS OF THE SOUTH AFRICAN MUSEUM

Illustrated material

SAM-PQ-MF-0528, RV, TBD 2224, 58 m

SAM-PQ-MF-0529, LV, TBD 2224, 58 m

SAM-POQ-MF-0530, RV, TBD 2224, 58 m

SAM-PQ-MF-0531, LV, TBD 2224, 58 m

SAM-PQ-MF-0532, C, TBD 2224, 58 m

Material

1 311 valves.

Remarks

Our material is identical to that described by Benson & Maddocks (1964) from

Knysna Lagoon. The marginal areas are not typical of the genus Cytheretta (as men-

tioned by Benson & Maddocks in the original description), and the species is best

accommodated in Bensonia, erected by Rossi de Garcia (1969) for material from the

Miocene of Argentina.

A, species with very similar ornamentation (slightly coarser ribbing), lateral

outline, and hinge structure was recorded as ?Leguminocythereis sp. 1 from the Upper

Eocene of the JC-1 borehole offshore Natal (Dingle 1976). This specimen should

probably also be referred to the genus Bensonia.

Distribution

Benson & Maddocks (1964) recorded this species from Knysna Lagoon, where it

was the dominant taxon (30%) at Leisure Island. They quoted the high- and low-

water salinity ranges at this site as 33-350, suggesting that B. k. knysnaensis can tol-

erate salinities slightly below ‘normal’ marine values.

Modern populations of B. knysnaensis knysnaensis have been encountered in two

main offshore areas (Figs 20, 21): off Liideritz (31-88 m) and between the south-

western Cape and the eastern Agulhas Bank (15-94 m), and also in Knysna Lagoon,

where Hartmann (1974: 296) reported a relatively sparse fauna at Leisure Island. This

pattern suggests that south of 25°S the species has a modern distribution of inshore to

mid-shelf. I record a geographically isolated single modern valve at 19°S at 236 m on

the Walvis Ridge abutment, the presence of which is difficult to reconcile with the

main population centres. Smoothed plots of latitudinal variations in abundance of

modern B. k. knysnaensis (Fig. 4) show that, together with Ruggieria cytheropte-

roides, it is the dominant taxon in the Orange River—Liideritz sector, and the

southern part of the Liideritz—Walvis Bay shelf sector. It is also locally one of the

main elements in the modern fauna in the Saldanha—northern Cape Peninsula region.

Relict specimens have a wide distribution, from 19°S on the Walvis Ridge abutment,

to the eastern Agulhas Bank. The LDL increases from 172 m on the Agulhas Bank, to

236 m on the Walvis Ridge, whereas the UDL varies from inshore (15-31 m) off the

south-western Cape and Liideritz, to 139-149 m in the Orange River and Walvis

Ridge areas. Plotting water depth against the percentage of the species in the total

ostracod assemblage (Fig. 22) suggests a preferred water depth of 35-95 m. Latitudi-

nal distributions (Fig. 5) show that B. k. knysnaensis is an important element in the

relict faunas immediately north of Lideritz, and immediately north of Saldanha.

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 35

Bensonia knysnaensis robusta subsp. nov.

Fig. 19A—D

Derivation of name

robustus, Latin, strong—reference to thickened rib ornamentation.

Holotype

length height

SAM-PO-MEF-0532, LV, TBD 3972, 200 m 0,71 mm 0,37 mm

Paratypes

length height

SAM-PO-MEF-0533, RV, TBD 3972, 200 m 0,68 mm 0,35 mm

SAM-—PO-MEF-0534, RV, TBD 3972, 200 m 0,70 mm 0,35 mm

Material

43 valves.

Diagnosis

A heavily celated subspecies of Bensonia knysnaensis with thickened rib orna-

mentation and a smooth anterodorsal surface.

Description

Overall valve architecture and ornamentation are the same as in B. knysnaensis

knysnaensis, but in the new subspecies individual longitudinal ribs are thicker, with a

consequent diminution in size of intercostal grooves and pits. In addition, the antero-

dorsal region is heavily calcified, resulting in a smooth, plate-like area centred on the

eye spot. Similarly calcified, but less extensive areas occur posterodorsally, and over

the subcentral region that overlies the MS. Internal features are identical with

B. knysnaensis knysnaensis.

Remarks and distribution

The morphological features that differentiate the two subspecies of B. knysnaen-

sis are attributed to environmental factors. No modern specimens of B. knysnaensis

robusta have been recovered, and the geographical distribution of the subspecies is

restricted to water depths of 140-200 m in a narrow latitudinal range between 20° and

24°S, seaward of the modern mud belt to the west and north-west of Walvis Bay. Only

two samples from this area contained B. knysnaensis knysnaensis (both relict), and in

only one were both subspecies found together.

Family Cytherideidae Sars, 1925

Subfamily Neocytherideidinae Puri, 1957

Genus Neocytherideis Puri, 1952

This genus has world-wide distribution in shallow-water environments. Its taxo-

nomic status has recently been reviewed by Athersuch (1982).

36 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 19. A-D. Bensonia knysnaensis robusta subsp. nov. TBD 3792, 200 m. A. SAM—PQ—MF0533,

RV, SEM 2467. B. SAM-PQ-MF0532, holotype, LV, SEM 2469. C-D. SAM-—PQ-MF0534, RV.

C. Internal view, SEM 2472. D. MS, SEM 2473. E-F. Neocytherideis boomeri sp. nov., TBD 6836, 80 m.

E. SAM—PQ-MF0536, holotype, RV, SEM 2435. F. SAM—PQ-—MF0537, LV, SEM 2433. Scale bars

= 100 microns.

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 37

Neocytherideis boomeri sp. nov.

Figs 19E-F, 23A-D, 24

Neocytherideis sp. Keeler, 1981: 56-57, pl. 2 (fig. 19).

Copytus sp. Boomer, 1985: 58-59, pl. 3 (fig. 43).

Derivation of name

Named for Dr I. D. Boomer (University of East Anglia) who first noted the

species off south-western Africa.

15 longitude 20 25

latitude

Knysna

eee

a“

Fig. 20. Distribution of Bensonia knysnaensis (Benson & Maddocks, 1964).

B. k. knysnaensis shown as squares (relict) and crosses (modern: dashed

line is seaward extent); B. k. robusta subsp. nov. shown as triangles. See

Fig. 7 for abbreviations.

38 ANNALS OF THE SOUTH AFRICAN MUSEUM

Holotype

SAM-PO-MF-0536, RV, TBD 6836, 80 m

Paratypes

SAM-PQ-MF-0537, LV, TBD 6836, 80 m

SAM-PQ-MF-0538, LV, TBD 6836, 80 m

SAM-PQ-MEF-0539, RV, TBD 6836, 80 m

SAM-PQ-MF-0540, C, TBD 6836, 80 m

SAM-PQ-MF-0541, RV, TBD 6847, 94 m

Material

511 valves.

Diagnosis

length

0,98 mm

length height

096mm 0,29 mm

0.97 mm 0,29 mm

0,.95mm 0,27 mm

1,00 mm —

height

0,28 mm

width

0,23 mm

Species with acuminate AM outline that is ventrally directed in RV, relatively

strong narrow surface ridges sub-parallel to AM and PM, and a prominent fulcral

point. The adductor MS lie on an elongate platform.

Description

External features. Elongate, cylindrical valves with strongly acuminate AM, par-

ticularly in the RV, where the line of greatest length is directed ventrally. PM is

acutely rounded. Surface ornamentation consists of narrow ridges extending sub-

parallel to the valve margins. These are most prominent anteriorly and posteriorly.

latitude °S

0 100

200

300 m

Fig. 21. Latitudinal water-depth distribution of sites with Bensonia knysnaensis (Benson

& Maddocks, 1964). Horizontal lines delimit across-shelf populations. Crosses and

dashed line = LDL of field with modern B. k. knysnaensis, triangles and dot-dashed line

= field with B. k. robusta subsp. nov.

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA

100

0 100 200 300m

Fig. 22. Abundance of Bensonia knysnaensis (Benson & Maddocks, 1964) as percentage

of ostracod fauna plotted against water depth.

Fig. 23. A-D. Neocytherideis boomeri sp. nov., TBD 6836, 80 m. A. SAM-PQ-—MF0538, LV,

SEM 2443. B. SAM—PQ-—MF0539, RV, SEM 2447. C. SAM-—PQ-MF0540, carapace, dorsal view,

SEM 2439. D. SAM—PQ-—MF0539, RV, MS, SEM 2449. Scale bars = 100 microns.

40 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 24. Neocytherideis boomeri sp. nov.,

SAM-PQ-MF0541, RV, detail of MA, TBD

6847, 94 m. Scale bar = 100 microns.

15 longitude 20 25

latitude

Fig. 25. Distribution of Neocytherideis boomeri sp. nov. Squares = relict sites; crosses =

modern sites. See Fig. 7 for abbreviations.

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA Al

Internal features. Normal pore canals are prominent, but their surface expression

is faint. MA are broad anteriorly, with a wide triangular-shaped vestibulum, and mod-

erately wide posteriorly and posteroventrally. There are approximately ten short,

straight anterior radial pore canals. Hinge weakly lophodont. MS are prominent;

adductors consist of a closely adjacent row of four, set on a raised platform that is

linked to the prominent fulcral point by a ridge. The anterior scar is ovate and lies

relatively far anterior to the adductors.

Remarks

Although species of Neocytherideis have been reported world-wide (Kempf 1986),

the genus has until now not been formally recognized from the South Atlantic area.

Two species of the related genus Copytus have, however, been recorded from the

south-western Atlantic and nearby Antarctic regions: C. caligula Skogsberg, 1939, and

C. elongatus Benson, 1964 (e.g. Skogsberg 1939; Benson 1964; Neale 1967; Hartmann

1986). Copytus does not occur off southern Africa but is widespread in the Austral-

asian area, where C. rara McKenzie, 1967, occurs in the Eocene to Recent in

Australia and New Zealand (McKenzie 1967; Swanson 1969). The two genera are dis-

tinguished by AM shape, and hingement.

Neocytherideis boomeri sp. nov. has all the generic characters, but its AM outline

and surface ornamentation serve to distinguish it from previously described species of

the genus. It is overall slimmer and more acuminate than the type (N. subulata

(Brady, 1867)), and has a longer DM than N. cypria Athersuch, 1982, which lacks the

posterior ornamentation of N. boomeri. The two species of Neocytherideis that have

been recorded from the Southern Hemisphere (N. mediata Swanson, 1969— Miocene

New Zealand; and N. muehlenhardtae Hartmann, 1982—Recent New Zealand) both

differ from N. boomeri in ornamentation and AM outline.

A similar taxon is that illustrated by Hartmann (1978) from Wilie Creek on the

west coast of Australia as Copytus aff. C. rara McKenzie, 1967. This specimen has a

modern

latitude °S

100 200 m

Fig. 26. Latitudinal water-depth distribution of sites with Neocytherideis boomeri sp.

nov. Horizontal lines delimit across-shelf populations. Crosses and dashed line delimit

LDL of modern fauna.

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

similar AM outline and surface ornamentation (somewhat more reticulate) to N. boo-

meri, but a different hinge and MS.

Distribution

Neocytherideis boomeri has been found on the continental shelf between Chamais

Bay (28°S) and at least as far east as 23°E on the Agulhas Bank (Figs 25, 26). It was

not recorded by Brady (1880), Klie (1940) or Hartmann (1974). Modern populations

are confined to an inner shelf region between Hout Bay, False Bay and Quoin Point

(19,3°E) on the south-western Cape coast in water depths between 40 m and 94 m.

Here, modern valves make up between 2 and 100 per cent of the total N. boomeri

assemblage. Relict populations occur in two regions. Between Chamais Bay and the

Olifants River the species has UDL and LDL of 52 m and 227 m, whereas south of

Hout Bay, the UDL and LDL are 40 m and 227 m, respectively. Although the popu-

lations in both areas have similar depth limits, their abundance distributions are

different; the northern populations favour deeper water (>173 m), whereas in the

south the species is most abundant in <100 m.

100 200 m

Fig. 27. Abundance of Neocytherideis boomeri sp. nov. as percentage of ostracod fauna

plotted against water depth. Dashed line delimits modern fauna.

Family Hemicytheridae Puri, 1953

Subfamily Hemicytherinae Puri, 1953

Genus Ambostracon Hazel, 1962

The genus is interpreted in the sense of Valicenti (1977), who used the strength of

the ocular ridge as a criterion for distinguishing between the two subgenera Ambostra-

con Hazel, 1962, and Patagonacythere Hartmann, 1962.

Three species of the genus occur off south-western Africa, all belonging to

A. (Ambostracon).

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 43

Subgenus Ambostracon (Ambostracon) Hazel, 1962

Ambostracon flabellicostata (Brady, 1880)

Figs 28A—D, 29C, F

Cythere flabellicostata Brady, 1880: 88-89, pl. 1 (figs 6a—h). Puri & Hulings, 1976: 276-277, pl. 8

(figs 1-4).

Ambostracon sp. B Keeler, 1981: 113-115, pl. 6 (figs 9-10).

Ambostracon sp. D Keeler, 1981: 116-118, pl. 6 (figs 13-14).

Ambostracon sp. 2 Boomer, 1985: 45-46, pl. 4 (figs 62, 65).

Ambostracon (Patagonacythere) sp. A468 Frewin, 1987: 40, pl. 13A.

Illustrated material

SAM-PQ-MF-0542, LV, TBD 2973, 173 m

SAM-PQ-MF-0543, RV, TBD 2973, 173 m

SAM-PQ-MF-0544, C, TBD 2224, 58 m

SAM-PQ-MEF-0545, RV, TBD 2224, 58 m

Material

490 valves.

Remarks

In Brady’s (1880) material, the strong ocular ridge crosses the eye tubercle and

extends sub-parallel to the AM. In the anteroventral corner it curves and is continu-

ous with a ventrolateral ridge. The SCT is prominent and joined to the anteroventral

corner by a short ridge. The main features of the ornamentation and MS pattern are

shown in Figure 29.

As Valicenti (1977, table 1) has shown, the genus is mainly represented in the

South Atlantic—Antarctic area by species of A. (Patagonacythere), and the only record

of A. (Ambostracon) from the area outside southern Africa is from the Miocene of

Argentina (Ambostracon (A.) sp. 1 Rossi de Garcia, 1970). This is a relatively elon-

gate species with a prominent diagonal ridge extending between the posterodorsal and

anteroventral corners. No records of the genus were made from the Tertiary of Gabon

by Bold (1966), but Frewin (1987) has recorded two (possibly three) species from the

Lower Tertiary of the Agulhas Bank. One of these (described as A. (Patagonacythere)

sp. A468,) appears identical in ornamentation to A. (A.) flabellicostata, and occurs in

a sample of Lower Palaeocene—Upper Eocene age (TBD 1275). A second specimen

from the Upper Eocene (described as Ambostracon (P.) sp. B1457 by Frewin (1987)

may be conspecific, but this has stronger ornamentation, and the dorsolateral rib

pattern differs slightly from Brady’s types.

Distribution

Brady (1880) recorded this species only from ‘Challenger’ Station 140 (30-40 m)

in False Bay.

Modern specimens are restricted to nearshore sites off the south-western Cape

between Saldanha Bay (33,16°S: 58 m) and Cape Agulhas (34,77°S: 73 m), where they

have UDL and LDL of 15 m and 131 m, respectively (Fig. 30A). Keeler (1981) did

not differentiate modern specimens from the eastern Agulhas Bank.

44 ANNALS OF THE SOUTH AFRICAN MUSEUM

CMM

YY Vy

Fig. 28. A-D. Ambostracon (A.) flabellicostata (Brady, 1880). A. SAM-PQ-—MF0542, LV,

TBD 2973, 173 m, SEM 2515. B. SAM-POQ-MF0543, RV, TBD 2973, 173 m, SEM 2500.

C. SAM-PQ-MF0544, carapace, dorsal view, TBD 2224, 58 m, SEM 2522. D. SAM—POQ—MF0545,

RV, TBD 2224, 58 m, SEM 2516. E-F. Ambostracon (A.) levetzovi (Klie, 1940), TBD 3089, 18 m.

E. SAM-—PO-MF0546, LV, SEM 2502. F. SAM—PQ-MF0547, RV, SEM 2505. Scale bars =

100 microns.

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA

B | ee

Fig. 29. Ambostracon (Ambostracon): comparison of three species. A-F. Main rib patterns of

RV, and MS. A. A. (A.) keeleri sp. nov., SAM-PQ-—MF0552, TBD 6823, 120 m. B. A. (A.)

levetzovi (Klie, 1940), SAM—PQ-—MF0548, TBD 3089, 18 m. C. A. (A.) flabellicostata (Brady,

1880), SAM—PQ-MF0543, TBD 2973, 173 m. D. A. (A.) keeleri sp. nov., SAM—PQ-—MF0555,

LV, TBD 6835, 100 m. E. A. (A.) levetzovi (Klie, 1940), SAM—PQ—MF0549, RV, TBD 3089,

18 m. F. A. (A.) flabellicostata (Brady, 1880), SAM—PQ-MF0545, RV, TBD 2224, 58 m.

G. A. (A.) keeleri sp. nov., SAM—PQ—MF0556, LV, MA and radial pore canals, TBD 6823,

120 m. Scale bars: A-C, G = 200 microns; D-F = 100 microns.

46 ANNALS OF THE SOUTH AFRICAN MUSEUM

Relict populations occur in a narrow zone between latitude 20,43°S and the

eastern Agulhas Bank (Fig. 31A). Although this zone is widest on the Orange Shelf, it

is here that abundances are lowest (<5%). Two continuous areas with relatively high

abundances (>5%) lie along the coast: southern Namaqualand and south-western

Cape, and the vicinity of Walvis Bay. Over the latitudinal range of relict specimens,

the UDL falls into two well-defined groups: off Walvis Bay and on the Orange Banks

they are 142 m and 158 m, respectively, whereas off Luderitz and south-western Cape

they are 31 m and 40m, respectively (Fig. 32A). The latter values are similar to

modern UDL. In all areas the LDL lies between 184 m and 223 m. Plotting abun-

dance against water depth suggests that there is an abundance minimum between

about 70 m and 90 m water depth off the south-western Cape (Fig. 33A).

Ambostracon (Ambostracon) levetzovi (Klie, 1940)

Figs 28E-F, 29B, E, 34A—C

Eucythereis levetzovi Klie, 1940: 419-421, figs 23-29.

Aurila tevetzovi (Klie, 1940) Hartmann, 1974: 284, pl. 149 (fig. 7).

Illustrated material

SAM-PQ-MEF-0546, LV, TBD 3089, 18 m

SAM-PQ-MEF-0547, RV, TBD 3089, 18 m

SAM-POQ-MEF-0548, RV, TBD 3089, 18 m

SAM-—PQ-MF-0549, RV, TBD 3089, 18 m

SAM-PO-MF-0550, LV, TBD 3089, 18 m

Material

19 valves.

Remarks

Klie (1940) and Hartmann (1974) both recorded this species on algae at inshore

sites in Lideritz Bay (no depth given). The present material is from one site

(TBD 3089: 18 m) in St Helena Bay, along with relict and modern A. (A.) keeleri sp.

nov. and relict A. (A.) flabellicostata. It is easily distinguished from the latter by the

straightness and strength of the ocular ridge, and from A. (A.) keeleri by the fact that

the ocular ridge crosses the eye tubercle.

Ambostracon (Ambostracon) keeleri sp. nov.

Figs 29A, D, G, 34D-F, 35A-B

Ambostracon sp. C Keeler, 1981: 115-116, pl. 6 (figs 11-12).

Ambostracon sp. E Keeler, 1981: 118-119, pl. 6 (figs 15-17).

Ambostracon sp. F Keeler, 1981: 119-120, pl. 6 (figs 18-19).

Ambostracon sp. 1 Boomer, 1985: 45-46, pl. 4 (figs 67-69).

Derivation of name

Named for Mr N. P. Keeler, formerly of University College of Wales, Aberyst-

wyth, who first recovered the species.

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 47

17 1g longitude 9 17 18 longitude 19

2 St Helena Bay

fe St Helena Bay

Saldanha

33 al

5h,

latitude

Hout Bay

u False Bay

34 1

is |

Ambostracon (A.) flabellicostata

Fig. 30. Distribution of Ambostracon (A.) spp. off the south-western Cape. A. A. (A.) flabellicostata

(Brady). B. A. (A.) keeleri sp. nov. Solid squares are modern sites.

Holotype

length height

SAM-PQ-MF-0551, LV, TBD 6823, 120 m 0,70 mm 0,38 mm

Paratypes

length height width

SAM-PQ-MF-0552, RV, TBD 6823,120m 0,68mm 0,33 mm —

SAM-—PO-ME-0553, C, TBD 6823, 120 m 0,73 mm — 0,30 mm

SAM-PQ-MF-0554, RV, TBD 6823,120m 0,68mm _ 0,35 mm —

SAM-—PQ-MEF-0555, LV, TBD 6835,100m 0,69mm 0,38 mm —

SAM-PQ-MEF-0556, LV, TBD 6823, 120 m

Material

1 024 valves.

Diagnosis

Species with a strong ocular ridge that runs anterior to the eye tubercle, and is

not continuous with the ventrolateral ridge. Ribs radiate centrally from SCT.

48 ANNALS OF THE SOUTH AFRICAN MUSEUM

A 15 longitude 29 25 B 15 longitude 99 25

Ambostracon (A.) flabellicostata Ambostracon (A.) keeleri

20 20

g 2

5 iS

30 30

Fig. 31. Distribution of Ambostracon (A.) spp. A. A. (A.) flabellicostata (Brady). B. A. (A.) keeleri

sp. nov. Squares = relict sites, crosses = modern sites. See Fig. 7 for abbreviations.

Description

External features. Sub-quadrate lateral outline, with males more elongate than

females. AM has numerous small spines, PM has three short stout spines postero-

ventrally. Central ornamentation consists of ribs radiating from a SCT with intercostal

reticulation, and a semi-elliptical ridge in the posterodorsal area. Peripheral ribs

consist of: a prominent, almost vertical rib that extends across the anterior area, and

does not cross the eye tubercle, which it skirts anteriorly with a sickle-shaped deflec-

tion; a ventrolateral rib that runs from the PM and which, in the LV, abuts the

anterior ridge, and in RV, is continuous with the anterior ridge; a thin rib that extends

from the eye tubercle along the DM; and a curved rib that loops from a position near

the SCT, via the posterodorsal shoulder down across the valve almost to the ventro-

lateral ridge. The eye tubercle is a large dome. In juveniles it is linked to the SCT by a

prominent short curved rib.

Internal features. Typical of the genus. AM areas are avestibulate, with numerous

(at least 30) straight, hair-like marginal pore canals. Hinge amphidont, with a promi-

nently lobed PTE in the RV. MS could not be clearly seen, despite the large number

of specimens available. They appear to be simpler than those in A. flabellicostata and

A. levetzovi, with a total of six scars (Fig. 29D).

Remarks

Ambostracon (A.) keeleri is easily distinguished from A. (A.) flabellicostata, with

which it usually co-occurs, by the difference in AM ridge pattern (see Fig. 29).

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 49

A A. (A.) flabellicostata

= ro] = = @ o "8 mg

t_|

30 5

” a | ae

xe)

= 79 lg

a = >

20 :

B A. (A.) keeleri

(o>)

(o)

latitude °S

0 100 200 300

Fig. 32. Latitudinal water-depth distribution of sites with Ambostracon (A.) spp. Hori-

zontal lines delimit across-shelf populations. A. A. (A.) flabellicostata (Brady, 1880).

B. A. (A.) keeleri sp. nov.

Ambostracon (A.) sp. A463 Frewin, 1987, from the Upper Palaeocene—Lower

Eocene of the Agulhas Bank has a similar ocular rib to A. (A.) keeleri, but a different

central area ornamentation, which is closer to that of A. (A.) flabellicostata.

Its closest relative is probably A. (A.) longiducta (Skogsberg, 1928), which also

has a similar ocular rib that is deflected anterior to the eye tubercle. The two species

differ in the course of the ocular rib, which is parallel to the AM margin in A. (A.)

longiducta; in the disposition of ribs posterodorsally and in A. (A.) longiducta being

somewhat plumper in outline. The latter species has been reported from various

localities in Antarctica and the Subantarctic area: Ross Sea, 57 m (Benson 1964);

Bransfield Strait, 133 m (Hartmann 1986, 1987); and South Georgia, 12-52 m (Skogs-

berg 1928).

Distribution

Modern specimens of A. (A.) keeleri are confined to nearshore sites off the south-

western Cape between an isolated occurrence in St Helena Bay (32,68°S: 18 m) and

50 ANNALS OF THE SOUTH AFRICAN MUSEUM

A. (A.) flabellicostata

A. (A.) keeleri

0 100 200 300 m

Fig. 33. Abundance of Ambostracon (A.) spp. as percentage of ostracod fauna plotted

against water depth. Dashed lines = LDL (A), and UDL and LDL (B) of modern

faunas. A. A. (A.) flabellicostata (Brady, 1880). B. A. (A.) keeleri sp. nov.

Cape Agulhas (34,77°S: 73 m). The main concentration of samples lies off Hout Bay

on the Cape Peninsula. UDL and LDL are 15 m and 140 m, respectively (Fig. 30B).

Relict specimens are recorded between latitude 20,43°S and the eastern Agulhas

Bank (Fig. 31B). A narrow zone of relatively high abundance (>5% total ostracod

fauna) stretches from the south-western Cape, to the vicinity of Liideritz (25,6°S),

where it forms a broader zone on the mid-shelf. A further narrow zone lies between

20,43°S and 23,43°S on the Walvis shelf. Over the latitudinal range of the relict speci-

mens, UDL falls into two well-defined groups: off Walvis Bay and on the Orange

Banks they are 160m and 158m, respectively, whereas off Luderitz and south-

western Cape they are 31 m and 15 m, respectively (Fig. 32B). The latter values are

similar to those for modern UDL. Off the Walvis Shelf the LDL is at 200 m, whereas

in all the southern areas the LDL lies between 252 m and 303 m. Plotting abundance

against water depth suggests that there is an abundance peak between 90 m and 160 m

water depth off the south-western Cape (Fig. 33B).

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 51

Fig. 34. A-C. Ambostracon (A.) levetzovi (Klie, 1940), TBD 3089, 18 m. A. SAM—-PQ-MF0548, RV,

SEM 2498. B. SAM-PQ-MF0549, RV, SEM 2506. C. SAM-POQ-MF0550, LV, SEM 2525.

D-F. Ambostracon (A.) keeleri sp. nov., TBD 6823, 120 m. D. SAM—PQ-MFO0551, holotype, LV,

SEM 2529. E. SAM—PQ-—MF0552, RV, SEM 2532. F. SAM-PO-MF0553, carapace, dorsal view,

SEM 2533. Scale bars = 100 microns.

52 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 35. A-B. Ambostracon (A.) keeleri sp. nov. A. SAM—PQ-MF0554, RV, TBD 6823, 120 m,

SEM 2544. B. SAM—PQ-MF0555, LV, TBD 6835, 100 m, SEM 2540. C-—D. Palmoconcha walvis-

baiensis (Hartmann, 1974), TBD 3940, 184 m. C. SAM—PQ-—MF0562, LV, SEM 2420. D. SAM—PQ-

MF0563, RV, SEM 2417. Scale bars = 100 microns.

Summary of the distribution of the genus Ambostracon

Ambostracon (A.) flabellicostata and A. (A.) keeleri have very similar distribution

patterns, particularly for modern specimens, where they are both confined to the

south-western Cape (Fig. 4), and best developed off the middle part of the Cape Pen-

insula in almost identical depths: 15-131 m and 15-140 m, respectively.

There are subtle differences in their relict distributions. A. (A.) keeleri is best

developed in a more or less continuous zone from the south-western Cape to the vicin-

ity of Liideritz, with the Walvis Shelf populations relatively sparse. In contrast,

A. (A.) flabellicostata is best developed off the south-western Cape and on the Walvis

shelf, with only a relatively sparse representation on the Namaqualand—Orange

shelves. Both species show a very similar latitudinal UDL variation, with the areas off

the Walvis and Orange shelves having values c. 100 m deeper than areas to the north

and south. Regionally, relict Ambostracon faunas are an important component of the

ostracod populations at the northern end of the Cape Peninsula, and immediately

south of Walvis Bay (Fig. 5).

Depth versus percentage plots (Fig. 33) suggest that A. (A.) keeleri prefers mid-

shelf environments (c. 120 m), whereas A. (A.) flabellicostata has peaks in the near-

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 53

shore (c. 40m) and mid- to outer-shelf (c. 130-200 m) zone. Ambostracon (A.)

levetzovi is confined to coastal and inshore sites between Luderitz and St Helena Bay.

Family Loxoconchidae Sars, 1925

Taxonomically the family Loxoconchidae is complex. It has a relatively long geo-

logical history (late Cretaceous to Recent) and the various genera within it have

world-wide distribution. The generic classification followed here is an emended

version of that discussed by Athersuch & Horne (1984).

The family is relatively well-represented in the Quaternary around southern

Africa with 12 species, eight of which occur off the south-western and southern coasts

(Fig. 36):

Loxoconcha megapora Benson & Maddocks, 1964. Reported from Knysna Lagoon

(Leisure Island—Benson & Maddocks 1964), whereas the variety L. megapora magna

occurs at Liideritz and Kommetjie (Cape Peninsula— Hartmann 1974).

L. parameridionalis Benson & Maddocks, 1964. Reported from Knysna Lagoon, asso-

ciated with sandy substrates with Zostera and a maximum salinity of 30°/.. (Benson &

Maddocks 1964; Hartmann 1974).

Australoloxoconcha favornamentata Hartmann, 1974. A subtropical coastal species

living on fine sand substrates. Reported from coastal sites at Knysna, St Lucia, and

Maputo.

Australoloxoconcha parafavornamentata Hartmann, 1974. Reported only from Knysna

Lagoon.

Palmoconcha walvisbaiensis (Hartmann, 1974). Reported from coastal and inner shelf

areas, Walvis Bay to south of Luderitz.

Palmoconcha? walvisridgensis sp. nov. An inner shelf species reported only from the

Walvis Ridge abutment area.

Palmoconcha subrhomboidea (Brady, 1880). An inner shelf species reported between

the Cape Peninsula and eastern Agulhas Bank.

Kuiperiana angulata sp. nov. An outer—inner shelf species reported from the Walvis

Ridge Shelf to south of the Cape Peninsula.

Fossil representatives of the family from south-western Africa have been reported

from the Agulhas Bank (4 species, Palaeocene-Eocene—Frewin 1987), the Natal off-

Australoloxoconcha favornamentata eu=======

Loxoconcha parameridionalis em»

Australoloxoconcha parafavornamentata em

Palrnoconcha subrhomboide?2 ===

? Kuiperiana angulata

? Palmoconcha walvisbaiensis

? emu 2Palmoconcha walvisridgensis

a Ne ee eee, Ce

WRA WB L OR CP K

Fig. 36. Distribution of coastal and shelf species of the family Loxoconchidae around

south-western Africa. Abbreviations: WRA—Walvis Ridge abutment shelf; WB—

Walvis Bay; L—Lideritz; OR—Orange River; CP—Cape Peninsula; K—Knysna.

54 ANNALS OF THE SOUTH AFRICAN MUSEUM

shore (1 species, Oligocene—Dingle 1976), and Gabon (2 species, Mio—Pliocene—

Bold 1966).

Genus Palmoconcha Swain & Gilby, 1974

This genus is distinguished by its gongylodont hinge with smooth ME, its

Y-shaped anterior MS and a fulcral point adjacent to the second or third adductor scar

(Swain & Gilby 1974; Horne & Kilenyi 1981; Athersuch & Horne 1984). Its known

geographical range is Europe (including the Mediterranean and Black seas), the east

and west coasts of North America, and the south-east Atlantic.

Palmoconcha walvisbaiensis (Hartmann, 1974)

Figs 35C-D, 37A-E

Loxoconcha walvisbaiensis Hartmann, 1974: 297-298, pl. 65 (figs 488-497). Boomer, 1985: 54-56,

pl. 1 (figs 14-15).

non Loxoconcha cf. L. walvisbaiensis Hartman, 1974. Frewin, 1987: 50, pl. 14 (fig. G).

Illustrated material

SAM-PQ-MEF-0562, LV, TBD 3940, 184 m

SAM-POQ-MEF-0563, RV, TBD 3940, 184 m

SAM-PQ-MEF-0564, RV, TBD 3926, 236 m

SAM-PQ-MF-0565, LV, TBD 3926, 236 m

Hamburg University Catalogue No. K30069, LV, Walvis Bay.

Material

475 valves.

Remarks

The smooth ME of the hinge, the Y-shaped anterior MS, and the position of the

fulcral point allow me to confidently re-assign Hartmann’s species to the genus Palmo-

concha.

Frewin’s (1987) record of Loxoconcha cf. L. walvisbaiensis Hartmann, 1974, is

not conspecific with Hartmann’s species, and possibly belongs in the genus Saida.

Distribution

This species was originally recorded by Hartmann (1974) from Walvis Bay

lagoon. Our study has shown it to be confined to areas north of about 26°S (Fig. 38)

and, although there are no precise data on its northern latitudinal range, Hartmann

(1974) did not record it from Mogamedes (15°S: Angola).

Modern populations of Palmoconcha walvisbaiensis occur from north of the

Walvis Ridge shelf (17,5°S) to just south of Walvis Bay (c. 24°S), where the UDL and

LDL are 15 m and 236 m, respectively (Fig. 39).

Relict populations extend 2 degrees farther south (to just north of Lideritz) but

the depth range is similar to that for the modern specimens (31-280 m).

In a high percentage of cases, this species is the only ostracod recovered from the

sample in which it occurs (10 of 30 sites), whereas in 22 out of 30 it constitutes

>50 per cent of the total ostracod assemblage. Although absent from within the main

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 55

YL MEE

Ag ee y ay Z

Y tj Yj Yi

tj ti UY g Uy Yj

Fig. 37. A-E. Palmoconcha_ walvisbaiensis (Hartmann, 1974). A-C. TBD 3926, 236 m.

A. SAM-PQ-—MF0564, RV, SEM 2428. B—C. SAM-PQ-MF0565, LV. B. Internal view, SEM 2421.

C. MS, SEM 2423. D. Paratype, Hamburg University catalogue slide No. K30069, LV, Walvis Bay,

SEM 2566. E. Paratype, Hamburg University catalogue slide No. K30069, LV, Walvis Bay, SEM 2568.

F. Palmoconcha subrhomboidea (Brady, 1880), SAM—PQ-—MF0568, LV, TBD 5254, 40 m, SEM 2965.

Scale bars = 100 microns.

56 ANNALS OF THE SOUTH AFRICAN MUSEUM

A 15 longitude 929 25

el et TE ie) a ee en eee

\ uu WR 25

20 s 2)

\ 4

\ aS

(

pe)

(>)

modern is |

latitude

oo ie

aN 0 100 200m

Fig. 38. A. Distribution of Palmoconcha walvisbaiensis (Hartmann, 1974). Squares = relict sites,

crosses = modern sites (seaward and southward extent shown by dashed line). See Fig. 7 for abbrevi-

ations. B. Latitudinal water-depth distribution of sites with P. walvisbaiensis (Hartmann, 1974).

Squares—relict sites, crosses—modern sites.

diatomaceous mud belt, this species seems relatively tolerant of oxygen-depleted

water, in which ostracod assemblages are generally sparse and of low diversity. In

addition, its southward modern limit lies in the vicinity of the maximum southerly

intrusion of warm, saline Angola Current water (e.g. Shannon 1985).

Palmoconcha? walvisridgensis sp. nov.

Figs 40B, 41B-—C

?Loxoconcha sp. aff. L. australis Brady, 1880. Bold, 1966: 170, pl. 3 (fig. 6).

Derivation of name

Named for the type locality of the species, the Walvis Ridge abutment shelf.

Holotype

length height

SAM-—POQ-MF-0566, C, TBD 3888, 154 m 0,50 mm 0,30 mm

Paratype

length height

SAM-PQ-MF-0567, C, TBD 3972, 200 m 0,50 mm 0,30 mm

Material

5 valves.

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA D1

Fig. 39. A—C. Palmoconcha subrhomboidea (Brady, 1880), TBD 5254, 40 m. A. SAM-PQ-MF0569,

RV, SEM 2968. B. SAM—PQ-MF0570, RV, SEM 2969. C. SAM—PO-MF0571, LV, SEM 2962.

D-F. Kuiperiana angulata sp. nov. D. SAM-PQ—MEFO0572, holotype, LV, TBD 2924, 158 m,

SEM 2919. E. SAM—PQ-MF0573, RV, TBD 3587, 140 m, SEM 2916. F. SAM—POQ-MF0574, RV,

TBD 3524, 475 m, SEM 2923. Scale bars = 100 microns.

58 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 40. A. Kuiperiana angulata sp. nov., SAM—PQ-—MF0576, RV,

MS, TBD 2974, 186 m. B. Palmoconcha? walvisridgensis sp. nov.,

SAM-PO-MF0566, holotype, carapace, dorsal outline, TBD 3888,

154 m. Scale bars: A = 100 microns, B = 200 microns.

Diagnosis

Species with asymmetrically rounded, ventrally directed AM outlines, straight

DM, and two strong, curved ventrolateral ribs.

Description

External features. Elongate ovate lateral outline. Broad, asymmetrically rounded

AM with wide rims, ventrally directed. PM rounded, semi-caudate, with apex dorsally

directed. DM straight, with a small step at the anterior cardinal angle. VM slightly

convex, partly obscured by lateral surface overhang. Surface strongly reticulate, with

two curved, sub-parallel ribs on ventrolateral surface. Reticulation is coarsest in sub-

central areas.

No internal features were observed in specimens that were either poorly pre-

served, or carapaces.

Remarks

Palmoconcha? walvisridgensis is probably conspecific with the taxon recorded

from the Pliocene of Gabon by Bold (1966) as Loxoconcha aff. L. australis Brady.

The specimen illustrated in Bold (1966, pl. 3 (fig. 6)) appears to be somewhat abraded

and consequently has less robust surface ornamentation than my material, but its

overall valve shape and reticulation pattern is very similar. The lectotypes of L. aus-

tralis Brady, 1880, as illustrated by Puri & Hulings (1976, pl. 18 (figs 17-18), pl. 19

(figs 1-4)) show that P.? walvisridgensis differs from Brady’s species in lacking the

strong upward sweep of the posteroventral outline, and in the outline of the postero-

dorsal margin, which is more acuminate in our species. In addition, L. australis has a

distinct posterodorsal hinge ear that gives the posterior end of the DM outline a slight

concavity.

An Indo-Pacific species that has a similar outline and ornamentation to P.? wal-

visridgensis is Loxoconcha paiki Whatley & Quanhong, 1987, from the Persian

Gulf—Malacca Straits region. This differs from my species in the outline of the antero-

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA By)

Fig. 41. A. Kuiperiana angulata sp. nov., SAM—PQ-MFO0S575, carapace, dorsal view, TBD 2924,

158 m, SEM 2920. B—C. Palmoconcha? walvisridgensis sp. nov. B. SAM-PQ- MF0566, holotype,

carapace, right view, TBD 3888, 154 m, SEM 2972. C. SAM-—POQ-MEF0567, carapace, left view,

TBD 3972, 200 m, SEM 2973. D-F. Ruggieria cytheropteroides (Brady, 1880). D. SAM-PQ- MF0S588,

carapace, dorsal view, TBD 2472, 201 m, SEM 2426. E. SAM—PQ-MF0589, LV, TBD 2975, 180 m,

SEM 2403. F. SAM—PQ-MF0590, RV, TBD 2975, 180 m, SEM 2399. Scale bars = 100 microns.

60 ANNALS OF THE SOUTH AFRICAN MUSEUM

ventral area, and in its more strongly curved muri in the reticulum of the central valve

area.

Distribution

Palmoconcha? walvisridgensis was found at three stations in the northernmost

part of the study area: on the southern part of the Walvis Ridge abutment shelf

(20,4°S) and to the north of the abutment on the narrow shelf at the southern end of

the Angola Basin, just south of the Kunene River (17,5°S) (Fig. 42). Modern valves

were recovered only from the latter area.

The depth range suggested by these sites is relatively narrow and deep

(154-200 m) with the modern population occupying the shallower depth.

These data suggest that P.? walvisridgensis is a subtropical species that inhabits

the middle to outer shelf.

Palmoconcha subrhomboidea (Brady, 1880)

Figs 37F, 39A—C

Loxoconcha subrhomboidea Brady, 1880: 121, pl. 28 (figs 4a—d). Puri & Hulings, 1976: 298-299,

pl. 18 (figs 15-16).

Loxoconcha sp. B Keeler, 1981: 143-154, pl. 8 (figs 10-11).

Loxoconcha sp. B192 Frewin, 1987: 46-47, pl. 16A-—F, text-fig. 2.11A.

15 longitude 20 25 B

R \ag @

ad *) [ i

| 7 oN 6

cpt K J P. subrhomboidea

4 |

TaN 304

4 o, 2 WR - 7

201 B Or al

SBN ?Palmoconcha walvisridgensis s

SE =

4 oo \ WB

} 20

{ ® a

o 500 0

se 0 1000 m

re = iL = K.angulata

¢L

J Bre

| aH OR

4 ay aN

30 8 \

| nk

| Palmoconcha ‘

5 subrhomboidea ce te ee

SS ao

Fig. 42. A. Distribution of Palmoconcha? walvisridgensis sp. nov. (triangles, dashed line shows

extent), Kuiperiana angulata sp. nov. (squares = relict sites, solid line south-east of Cape Peninsula

shows southern extent; crosses = modern sites), and Palmoconcha subrhomboidea (Brady, 1880) (dia-

monds, dot-dashed lines shows seaward extent). See Fig. 7 for abbreviations. B. Latitudinal water-

depth distribution of sites with P. subrhomboidea (Brady, 1880) (triangles), K. angulata sp. nov. (solid

squares), and P. ?walvisridgensis sp. nov (crosses).

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 61

Illustrated material

SAM-PQ-MF-0569, RV, TBD 5254, 40 m

SAM-PQ-MF-0570, RV, TBD 5254, 40 m

SAM-—PQ-MF-0571, LV, TBD 5254, 40 m

Material

39 valves.

Remarks

The characteristic features of this species are a strong, curved, ventrolateral ridge

and furrow, and a sharply arched dorsomedian rib. Its placement within Palmoconcha

is not certain because no unequivocal views of the MS were available.

Distribution

Brady (1880) found this species only at ‘Challenger’ Station 140 in 15-20 fm

(27-37 m) in False Bay, and in the present study I have established that it does not

extend farther north, having recorded it only from False Bay (40 m), and at two sites

west of the Hout Bay area (90-94 m) (Fig. 42A—B). Modern specimens occur in False

Bay and at 94 m off the western side of the Cape Peninsula, whereas relict valves

occur in False Bay and at 90 m off the Peninsula. Its extension on to the Agulhas

Bank, where Keeler (1981) noted it at four sites with a water depth range 65-112 m,

indicates that P. subrhomboidea is a warm-water taxon. It is not known whether any

of Keeler’s (1981) material was from modern populations.

Frewin (1987) illustrated this species from Palaeogene (?Upper Palaeocene—

Middle Eocene) sediments of the Agulhas Bank, where she recorded five valves. A

similar (but not conspecific) species (Loxoconcha sp. A3243 Frewin, 1987: 48) occurs

in Upper Eocene strata of the same area.

Genus Kuiperiana Bassiouni, 1962

This genus was erected by Bassiouni (1962) to differentiate species of Loxocon-

cha that possess a long DM and consequently a long hinge ME. Previous records of

this genus have been confined to Oligocene—Miocene strata of north-western Europe

(e.g. see Uffenorde 1981; Kempf 1986)

Kuiperiana angulata sp. nov.

Figs 39D-F, 40A, 41A

Derivation of name

Angularis—Latin = angular, reference to angular, truncated alae.

Holotype

length height

SAM-PQ-MF-0572, LV, TBD 2924, 158 m 0,48 mm 0,29 mm

62 ANNALS OF THE SOUTH AFRICAN MUSEUM

Paratypes

length height width

SAM-PQ-MF-0573, RV, TBD 3587, 140m 0,50mm 0,30 mm —

SAM-PO-MEF-0574, RV, TBD 3524,475 m 0,58mm 0,29 mm —

SAM-POQ-MEF-0575, C, TBD 2924, 158 m 0,51 mm — 0,25 mm

SAM-PO-MF-0576, RV, TBD 2974, 186 m

Material

62 valves.

Diagnosis

Reticulate species of Kuiperiana with small, posteriorly angular alae.

Description

External features. Small sized, sub-quadrate in lateral outline. AM asymmetrically

rounded, PM with a blunt caudal process. DM straight, with small, prominent cardinal

angles. There are narrow AM and PM rims. Surface overall reticulate, with short

longitudinal ribs that converge anterior of the central area. There is a small, pos-

teriorly angular ala that is ventrally directed. This enhances the convex VM outline. In

dorsal view the carapace tapers anteriorly, and has a slight median constriction.

Internal features. Hinge gongylodont, with a smooth ME and a strongly lobed

PTE in RV. MS small, consisting of four rounded/elliptical adductors as two com-

pound pairs, and a rounded eight-shaped frontal scar. Normal pore canals prominent

and widely spaced.

Remarks

Reference of this species to Kuiperiana is not unequivocal because Bassiouni’s

type (Loxoconcha wanneri Kuiper, 1918) does not possess alae. However, the overall

shape, ornamentation and hinge of K. angulata sp. nov. fit such a placement reason-

ably well. Whatley & Quanhong’s (1987) genus Alataconcha is an alate loxoconchid,

but it has a short hinge, a convex LV DM, and relatively large alae. In external view,

Kuiperiana angulata is reminiscent of Loxoconcha heronislandensis Hartmann, 1981,

which occurs widely in the south-eastern Pacific (see Whatley & Quanhong 1987), but

lacks the latter species’s prominent eye tubercle, and has more longitudinally aligned

ornamentation. None of the loxoconchids previously reported from southern Africa

can be confused with the alate K. angulata.

Distribution

This is the most widely distributed loxoconchid on the continental shelf of south-

western Africa (Fig. 42A, B), although it rarely constitutes >10 per cent total ostra-

cod assemblage.

The modern population of Kuiperiana angulata sp. nov. lies in a narrow belt

between latitudes 19,9°S (Walvis Ridge abutment shelf) and 30°S (Hondeklip Bay,

Namaqualand coast), where its UDL and LDL are 150 m and 283 m, respectively.

The main relict populations lie on the Walvis and Orange shelves, but there are

three sites off the Cape Peninsula. Although the relict UDL is similar to that of the

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 63

modern fauna (131 m off the Cape Peninsula—Fig. 42B), the LDL suggests a large

increase into deeper water compared to modern sites, and in the Walvis—Orange

sector it may increase to 1 000 m (although the latter may be an allochthonous occur-

rence).

Plotting abundance against water depth for the total assemblage suggests two

peaks: c. 300 m and 750 m. The shallower of these is probably real, but the veracity of

the latter is uncertain.

Overall distribution of loxoconchids

Only two of the species that have been reported from the continental shelf off

south-western Africa have a widespread distribution: Palmoconcha walvisbaiensis,

which is confined to areas north of 27°S, and Kuiperiana angulata, which occurs along

the whole margin from the Walvis Ridge abutment to south of the Cape Peninsula

(Fig. 36). Loxoconcha megapora (including the variety magna) also has a wide geo-

graphical range, but so far reports of it have been confined to widely spaced coastal

sites.

Two other shelf species have much more limited ranges: Palmoconcha? walvis-

ridgensis is restricted to the subtropical northernmost area in the vicinity of the Walvis

Ridge abutment, and P. subrhomboidea is restricted to the Agulhas Bank region, and

penetrates along the west coast only as far as the Cape Peninsula.

Plotting the summed and smoothed abundances of the loxoconchids shows that

north of approximately 25°S (middle of the Liideritz—Walvis Bay shelf sector) they

constitute the dominant modern ostracod taxon (Fig. 4). This is particularly so in the

shelf sector between Walvis Bay and the Walvis Ridge abutment, where P. walvis-

baiensis accounts for 98 per cent of the loxoconchid population. They are minor

components of the modern fauna on the northern Orange Banks, and off the Cape

Peninsula. Within the relict assemblages, the loxoconchids become increasingly abun-

dant north of approximately 26°S, with peaks off Walvis Bay, on the Walvis Ridge

abutment, and in the vicinity of the Kunene River (Fig. 5). They are an insignificant

component of the relict fauna on the Orange Shelf, and areas farther south.

Changes in the relict and modern distributions of these taxa show that within the

older populations two subtropical Atlantic species penetrated up to 200 km farther

south: P. walvisbaiensis—2 degrees; and P.? walvisridgensis—2,5 degrees, whereas

Kuiperiana angulata established a population off the Cape Peninsula, 400 km south of

its modern limits, but did not inhabit the intervening area off the Namaqualand coast.

In contrast, there was no corresponding extension north of the Cape Peninsula of the

range of the warm water (‘Agulhas’) species Palmoconcha subrhomboidea.

Family Trachyleberididae Sylvester-Bradley, 1948

Genus Ruggieria Keij, 1957

Ruggieria cytheropteroides (Brady, 1880)

Figs 41D—-F, 43A-B, 44

Cythere cytheropteroides Brady, 1880: 78, pl. 15 (figs Sa—d). Puri & Hulings, 1976: 272-273, pl. 9

(figs 5-8).

Bosquetina sp. Keeler, 1981: 41-43, pl. 2 (fig. 1).

Ruggieria cytheropteroides (Brady, 1880) Boomer, 1985: 19-21, pl. 1 (figs 1-3).

64 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 43. A-B. Ruggieria cytheropteroides (Brady, 1880). A. SAM—PQ-MF0591, RV, TBD 3863,

150 m, SEM 2410. B. SAM—PQ-MFO0592, LV, TBD 2975, 180 m, SEM 2405. C-F. Henryhowella

melobesioides (Brady, 1869). C. SAM—PQ-MF0475, RV, TBD 311, 184 m, SEM 2625. D. SAM-—

PQ-MF0478, LV, TBD 311, 184 m, SEM 2626. E. SAM—PQ-MF0480, LV, TBD 3561, 655 m,

SEM 2591. F. SAM—POQ-MF0481, RV, TBD 3561, 655 m, SEM 2493. Scale bars = 100 microns.

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 65

Illustrated material

SAM-—PQ-MF-0588, C, TBD 2472, 201 m

SAM-PO-MEF-0589, LV, TBD 2975, 180 m

SAM-PO-MF-0590, RV, TBD 2975, 180 m

SAM-—PQ-MF-0591, RV, TBD 3863, 150 m

SAM-PO-MF-0592, LV, TBD 2975, 180 m

Material

5 489 valves.

Remarks

Ruggieria cytheropteroides differs from Keij’s genotype (Cythere micheliniana

Bosquet, 1852—see Uffenorde 1981, pl. 6 (figs 7, 10, 12)) from the Eocene—Oligo-

cene of north-western Europe, in being slightly plumper, with a more prominent

ventrolateral overhang. Otherwise the essential generic components are well seen in

Brady’s species (e.g. 1880, pl. 15 (fig. 5a)), although the lectotypes selected by Puri &

Hulings (1976) are worn, broken and probably instars. Features of note are the promi-

nent ventrolateral keel with a sharp posterior spine, the large ocular sinus, the

denticulate hinge ME and the elongate and weakly lobate RV ME, and the MS with

four large adductors, a relatively small U-shaped anterior scar, and a prominent pit

and antero-adjacent boss in front of the dorsal adductor.

Ruggieria cytheropteroides is similar in outline and general appearance to two

species that range along the northern Indian Ocean area (Gulf of Oman to Java Sea—

Whatley & Quanhong 1988): R. darwinii (Brady, 1868) and R. indopacifica Whatley

& Quanhong, 1988. Both these species differ from R. cytheropteroides in being coar-

sely reticulate.

Ruggieria and related genera are a widely distributed and diverse group on the

continental shelf of western and south-western Africa. Three species of the genus have

been recorded from the Tertiary of west and equatorial Africa: R. tattami Reyment,

1963 (Palaeocene—Eocene of Nigeria), and R. tetraptera tetraptera (Seguenza) and

R. rotundata (Ruggieri) (Bold 1966—Mio-Pliocene of Gabon), but none of these

species seems close to R. cytheropteroides, and may not be congeneric with Keij’s

types. Dingle (1976) did not report Ruggieria from the Tertiary of the Natal continen-

tal shelf, but Frewin (1987) illustrated an elongate species with strong posteroventral

spines (Ruggieria sp. A485) from the Upper Eocene of the eastern Agulhas Bank.

Fig. 44. Muscle scars of Ruggieria cytheropteroides

(Brady, 1880), SAM—PQ—MF0591, RV, TBD 3863,

150 m, SEM 2411. Scale bar = 100 microns.

66 ANNALS OF THE SOUTH AFRICAN MUSEUM

A 145 longitude 99 25 B 15 longitude 99 25

MODERN RELICT

20 20

latitude

30 30

Fig. 45. Distribution of Ruggieria cytheropteroides (Brady, 1880). A. Modern sites. B. Relict sites. See

Fig. 7 for abbreviations.

Keen (1975) recorded eight Recent species of the genus from the continental

shelf off West Africa (although he suspected that they are in fact not congeneric with

the holotype): R. triangulata Omatsola, 1972 (20-80 m), R. beninensis Omatsola,

1972 (20-30 m), R. lekkii Omatsola, 1972 (20-30 m), R. nigeriana Omatsola, 1970

(20-30 m), R. martinssoni Omatsola, 1972 (0-110 m, 10-60 m living), R. tricostata

Omatsola, 1972 (20-30 m), R. boldi Keen, 1975 (20 m), and R. leonensis Keen, 1975

(sandy sediments, 60-110 m). The last-named is very similar in shape, ornamentation

and internal features to R. cytheropteroides. It differs in having a faint surface reticu-

lation, a weak AM ridge, and in being slightly more triangular in lateral outline over

the anterior cardinal angle. Clearly, the two species are closely related.

Babinot & Kouyoumontzakis (1986) have recorded three of the West Africa taxa

from modern sediments off the mouth of the Congo River: R. lekkii (38-44 m);

R. martinssoni (38 m); and Ruggieria aff. R. triangulata (38 m).

Distribution

Brady (1880) recorded this species from ‘Challenger’ Station 142 off the Cape of

Good Hope (300 m).

Ruggieria cytheropteroides (combined modern and relict specimens) is overall the

second most abundant ostracod taxon (after Pseudokeijella lepralioides (Brady, 1880))

on the south-western African continental shelf (22% of all specimens recorded in the

present study, 39% of specimens in the samples containing the dominant taxa).

Modern specimens extend over a latitudinal range of 15 degrees between 19° and

35°S (Figs 4, 45A, 46). Off the Cape Peninsula (Fig. 46C) the species occurs in water

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 67

A 18-2795 BB 27-32°S

500 400 600 m 200 400 m

Cc 32-36°S

200 400 600 m

Fig. 46. Abundance of Ruggieria cytheropteroides (Brady, 1880) as percentage of ostracod fauna

plotted against water depth (5-point running means). Modern populations (Gioses) enclosed by

dashed line. A. 18—27°S. B. 27-—32°S. C. 32-36°S.

depths between 94 m and either 290 m or 475 m (the latter may be allochthonous),

whereas north of 32°S (Fig. 46B) it occupies mid-outer shelf depths between 147 m

and 469 m. The species becomes less abundant north of 27°S, and occurs in water

depths between 150 m and 475 m (Fig. 46A). In the vicinity of Walvis Bay (23—25°S)

the UDL of R. cytheropteroides increases to 295 m and the species is confined to the

outer shelf area. The northernmost sample site on the Walvis Ridge shelf is a single

valve.

These distributions give overall UDL and LDL of modern specimens for the west

coast as 94-475 m, with water depths of the main population abundances for the

Walvis, Lideritz—Orange-Namaqualand, and south-western Cape areas decreasing

steadily southward from 400 m, via 200 m to 150 m.

Relict specimens of R. cytheroptercides occur over the same latitude range as the

modern populations (i.e. Walvis Ridge abutment 19°S to south-western Cape 35°S:

Fig. 45B). The overall pattern of their latitudinal abundances is also similar (Figs 4,

5), with the main concentration of high abundance samples on the Namaqualand

—Orange Shelf sector. Across-shelf profiles (Fig. 46A, B) show that in the northern

sector, the maximum population abundance lies on the outer shelf (c. 370 m), and

that it moves inshore to c. 300 m on the Orange Shelf. The abundance/water depth

68 ANNALS OF THE SOUTH AFRICAN MUSEUM

profile is more complicated off the south-western Cape (Fig. 46C) where the major

abundance peak occurs at c. 190 m, with only a minor peak at the c. 300 m.

Table 4 and Figure 47 summarize the depth range changes for the various

R. cytheropteroides populations.

Subfamily Trachyleberidinae Sylvester-Bradley, 1948

Genus Henryhowella Puri, 1957

Aspects of the status of this genus have recently been reviewed by Dingle et al.

(1990), particularly as they pertain to local species. The present publication adopts

their taxonomic strategy.

Henryhowella melobesioides (Brady, 1869)

Fig. 43C-F

Cythere melobesioides Brady, 1869: 162, pl. 12 (figs 10-11); 1880: 108, pl. 18 (figs le-g). Puri &

Hulings, 1976, pl. 25 (figs 1-2).

non Cythere melobesioides Brady, 1869. Brady, 1880, pl. 18 (figs la—d).

Cythere nodulifera Brady, 1869: 163, pl. 19 (figs 24-25).

Henryhowella sp. Keeler, 1981: 162-163, pl. 9 (fig. 14).

Henryhowella sp. Boomer, 1985, pl. 1 (figs 6-8, 18).

non Henryhowella sp. Boomer, 1985: 25-27, pl. 3 (figs 38-39).

Henryhowella melobesioides (Brady, 1869) Dingle et al., 1990: 311-318, figs 42C-F, 43A—-F, 44A—D, 47A.

Illustrated material

SAM-PO-MF-0475, RV, TBD 311, 184 m

SAM-PO-MF-0478, LV, TBD 311, 184 m

SAM-—PQ-MF-—0480, LV, TBD 3561, 655 m

SAM-PO-MF-0481, RV, TBD 3561, 655 m

TABLE 4

Water depth distribution of Ruggieria cytheropteroides.

Depth Main Shift

UDL LDL range conc.

(m) (m) (m) (m) (m)

MODERN

North 150 475 325 400

Orange—Namaqualand 147 469 32) 200

South-western Cape 94 AT5 381 150

or 290 or 200

RELICT

North 150 590 440 370 -30

Orange—Namaqualand 135 530 395 290 +90

South-western Cape 94 560 466 190 +40

and 300 =and +150

UDL — upper depth limit

LDL — lower depth limit

Main conc. — depth of the sites with greatest abundances

Shift — difference between relict and modern depths of greatest abundances

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 69

latitude

0 200 400 600 m

Fig. 47. Latitudinal water-depth distribution of sites with Ruggieria cytheropteroides

(Brady, 1880).

Material

429 valves.

Remarks

Dingle et al. (1990) could not recognize any geographical consistency in the mor-

phological variation of individuals in the populations of Henryhowella on the

continental margin off south-western Africa. Consequently, in view of the uncertainty

surrounding the taxonomy of widely reported species such as H. asperrima (Reuss,

1850), they considered all morphological variants on the continental margin off south-

western Africa could be accommodated in Brady’s species H. melobesioides (Brady,

1869), which he regarded (Brady 1880) as conspecific with the type specimens from

Mauritius (Brady 1869). I adhere to this view.

Brady (1880) recorded H. melobesioides from c. 300 m (150 fm) off the Cape

Peninsula, and his figured specimens have relatively nodose spines. Dingle er al.

(1990) remarked that the deeper-water populations tend to have more slender spines,

and that coarsely spinose individuals are representative of the continental shelf popu-

lations (compare Figs 43C, D with Fig. 43E). I regard this as a response to

environmental factors (e.g. energy of bottom water).

Distribution

Henryhowella melobesioides is widely distributed along the continental margin of

south-western Africa (Fig. 48A, B) but, because Keeler (1981) did not find it on the

eastern Agulhas Bank, the eastward limit of the species lies between 19,28°E and

23,21°E. It is essentially a cold-water, west-coast taxon around southern Africa.

Modern populations occur along the west-coast margin between 35° and 19°S. Off

the Cape Peninsula the depth range is 140-290 m, but it increases to 430—990 m at the

latitude of Saldanha Bay. Consequently, north of the Cape Canyon (33,5°S) there are

no modern populations of H. melobesioides on the continental shelf. A wide barren

70 ANNALS OF THE SOUTH AFRICAN MUSEUM

A 15 longitude 20 25

latitude °S

latitude

0 1,0 2,0 3,0 km

Fig. 48. A. Distribution of Henryhowella melobesioides (Brady, 1869). Crosses = relict sites, solid

squares = modern sites. See Fig. 7 for abbreviations. B. Latitudinal water-depth distribution of sites

with H. melobesioides (Brady, 1869). Crosses = relict sites, solid squares = modern sites.

zone between the Orange River and Lideritz separates the Namaqualand and Liide-

ritz—Walvis Bay upper slope populations (Fig. 4), where in the latter the UDL and

LDL are 725 m and 1 430 m. Clearly, the whole of the low dissolved oxygen environ-

ment of the Walvis shelf, and the suspensate rich and lower-salinity shelf adjacent to

the Orange River are environmentally unsuitable for modern populations.

100

0 2 4 km

Fig. 49. Abundance of Henryhowella melobesioides (Brady, 1869) as percentage of ostra-

cod fauna plotted against water depth (5 point running means). 1 = abundance peak on

outer shelf off south-western Cape. 2 = abundance peak off Namaqualand.

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA Wik

Relict distribution patterns are similar to those of the modern populations, with

the exception of the area off Namaqualand, where the UDL is similar to that off the

Cape Peninsula (c. 100 m). A notable similarity between the two populations is the

absence of both modern and relict faunas in the Walvis—Orange shelf area. A major

difference is in the overall greater LDL of relict populations, with specimens occurring

to a maximum of 2 916 m in the extreme south.

Dingle et al. (1990) have previously considered the distribution of H. melobe-

sioides in deep water (>950 m), and Figure 53 shows a summed population profile

across the margin into abyssal depths. The species dominates the ostracod populations

between about 750 m and 1 500 m (from the core of the salinity minimum zone to the

base of the Antarctic Intermediate Water mass). Dingle et al. (1989) have suggested

that this is an Atlantic-wide phenomenon. The minor peak at the shallow (left side) of

Figure 49 reflects an abundance of H. melobesioides on the upper slope west of the

Cape Peninsula.

Genus Pseudokeijella gen. nov.

Diagnosis

Plump, ovate genus. Ornamentation is densely reticulate, with two antero-

marginal ribs emanating from a moderate to prominent eye spot. PM is spinose

posteroventrally. There is no ventrolateral carina or tendency to develop ventrolateral

spines or ridges. Hinge is holamphidont with denticulate LV ME. MS have an anterior

structure consisting of two very small, close-lying or partially fused scars (in a com-

pressed U-shaped), and four adductors of which the second is long and curved and the

others small and rounded.

Type species. Cythere lepralioides Brady, 1880.

Derivation of name

Pseudo- plus Keijella with reference to similarity to this genus.

Remarks

This genus is erected to accommodate a species that is closely allied to the two

genera Ruggieria Keij, 1957, and Keijella Ruggieri, 1967, but which differs from both

on significant features of outline, ornamentation and MS.

The type species of Ruggieria is Cythere micheliniana Bosquet, 1852, which is car-

inate with a posteroventral spine, and bears several further longitudinal ridges. Ketj

(1957) specifically mentioned in his remarks that Ruggieria lacks ornamentation with

concentricity. This genus is represented off south-western Africa by a typical species,

Ruggieria cytheropteroides (Brady, 1880).

Keyella is based on the species Cythere hodgii Brady, 1866. The type specimen of

this species, from the eastern Mediterranean, is lost, but Doruk (1973) has illustrated

topotypic material, as well as erecting two new species of the genus from Turkey. This

is an elongate taxon with a postero-ventrolateral spine and/or swelling whose valve

surface is either smooth, or has longitudinal rows of elongate fossae (‘slots’ of Doruk

1573),

2 ANNALS OF THE SOUTH AFRICAN MUSEUM

Pseudokeijella has some features of both these genera, and on balance is closer to

Keijella (hence the name) (Fig. 51). It differs from Ruggieria by lacking the character-

istic ventrolateral carina, but the MS are similar, with the third scar small and round

and set distinctly posterior to the fourth, and the second elongate. Keijella lacks a ven-

trolateral carina, and prominent longitudinal ridges, with the result that the overall

shape and surface ornamentation are more similar to those of Pseudokeijella.

However, Pseudokeijella has an ovate outline (in contrast to the elongate sub-

quadrate outline of Keijella), is plumper, and lacks any postero-ventrolateral spine or

swelling. The MS of Keijella and Pseudokeiella differ to the extent that the adductors

of the former are more elongate, and the anterior scar of the type species has the

shape of three sides of a rectangle.

Whatley & Quanhong (1988) have reported 10 species of Keijella from the

Malacca Straits region. I suspect that at least two of these (K. japonica (Ishizaki) and

K. reticulata Whatley & Quanhong) belong to Pseudokeijella. In addition, several

species previously referred to Leguminocythereis appear close to Pseudokeijella in

outline and ornamentation (their internal features are mostly not known). Within this

category I include Leguminocythereis lokossaensis Apostolescu, 1961, Leguminocythe-

reis frescoensis Apostolescu, 1961, ?Leguminocythereis cf. L. lokossaensis Aposto-

lescu (Dingle 1976), ?Leguminocythereis sp. 1 Dingle, 1976; Leguminocythereis cf.

L. exigua (Apostolescu) (Frewin 1987); Leguminocythereis sp. 1507 Frewin, 1987.

Pseudokeijella lepralioides (Brady, 1880)

Figs SOA-F, 51E-F, 55E-F

Cythere lepralioides Brady, 1880: 94, pl. 19 (figs Sa—d). Puri & Hulings, 1976: 280-281, pl. 12

(figs 10-11).

Ruggieria lepralioides (Brady) Keeler, 1981: 173-175, pl. 10 (figs 1-3).

Leguminocythereis? sp. Boomer, 1985: 47-49, pl. 1 (figs 4-5).

Leguminocythereis sp. 1507 Frewin, 1987: 44—45, pl. 1ISA—D.

Illustrated material

SAM-PO-MEF-0557, LV, TBD 6824, 90 m

SAM-PO-MEF-0558, RV, TBD 6824, 90 m

SAM-PO-MEF-0559, RV, TBD 6836, 80 m

SAM-PO-ME-0560, LV, TBD 6836, 80 m

SAM-PQ-MEF-0561, carapace, TBD 6847, 94 m

Material

8 181 valves.

Remarks

The lectotype of Brady’s (1880) species selected by Puri & Hulings (1976) is prob-

ably a penultimate instar, which has signs of physical wear. Our material shows that

the ventral parts of the AM and PM are spinose, and that the eye spot is moderately

well-developed, with two ribs running from it sub-parallel to the AM. The hinge is

holamphidont with, in the RV, a prominent tooth at the posterior part of the ATE, an

ovate smooth PTE, and a straight, denticulate ME. There is a deep internal ocular

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 73

Fig. 50. A-F. Pseudokeijella lepralioides (Brady, 1880). A. SAM-PQ-MF0561, carapace, dorsal

view, TBD 6847, 94 m, SEM 2389. B. SAM—PQ-MF0560, LV, MS, TBD 6836, 80 m, SEM 2381.

C. SAM—PQ-MF0559, RV, hinge, TBD 6836, 80 m, SEM 2387-88. D. SAM-PQ-—MF0560, LV, hinge,

TBD 6836, 80 m, SEM 2382-83. E-F. TBD 6824, 90 m. E. SAM—PQ-MF0557, LV, SEM 2372.

F. SAM—PQ-MF0558, RV, SEM 2377. Scale bars = 100 microns.

74 ANNALS OF THE SOUTH AFRICAN MUSEUM

A B

jC) $e

“e

C D

Sake.

E F

Fig. 51. Comparison of outline and MS of Pseudokeijella gen. nov. with

Ruggieria Keij, 1957, and Keijella Ruggieri, 1967. A-B. Ruggieria micheli-

niana (Bosquet, 1852), type species of Ruggieria Keij, 1957. A. Specimen

from Lower Miocene of Northern Germany (after Uffenorde 1981, pl. 6

(fig. 7)). B. Specimen from Lower Miocene of France (after Keij 1957, pl.

15 (fig. 5)). C-D. Keijella hodgii (Brady, 1866), type species of Keijella

Ruggieri, 1967. C. Specimen from Upper Miocene of Turkey (after Doruk

1973, pl. 1:9:54 (1)). D. Specimen from Mio—Pliocene of San Marino (after

Doruk 1973, pl. 1:9:56 (3)). E-F. Pseudokeijella lepralioides (Brady,

1880). E. SAM-—PQ-MF0558, RV, TBD 6824, 90 m. F. SAM-PO-

MF0559, RV, TBD 6836, 80m. Scale bars. A, C, E = 200 microns;

D, F = 100 microns; B unknown.

sinus anterior to the anterior hinge elements. The MS pattern consists of four dissimi-

larly sized adductors, two small anterior spots that lie very close, or are partly fused to

a flattened U-shape, and a prominent fulcral point. There is no noticeable MS depres-

sion, and consequently no STC on the outer lateral surface. In dorsal view,

P. lepralioides is elliptical.

Distribution

Brady (1880) reported this species from two localities off south-western Africa:

Station 140 in False Bay (30-40 m), and Station 142, south-west of the Cape of Good

Hope (300 m).

Pseudokeijella lepralioides (combined modern and relict specimens) is overall the

most abundant ostracod taxon on the south-western African continental shelf (32% of

all specimens recorded in the present study, 39% of specimens in the samples con-

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA WD

15 longitude 20 25

+ \\WR

20 WF

g ne

= aa \

masa OR

mien x

ace

oe x

+ mR

+ 4 + \

CP mee

Fig. 52. Distribution of Pseudokeijella lepralioides (Brady, 1880). Crosses = relict

sites, solid squares = modern sites. See Fig. 7 for abbreviations.

taining the dominant taxa of this paper), and Keeler (1981) recorded it as the most

abundant species on the eastern Agulhas Bank (average 10% in 8 samples).

Modern specimens are concentrated in two areas (Figs 4, 52). On the Orange

Shelf directly west of the Orange River (28—29°S), where UDL and LDL lie between

126m and 183m water depth, and between Cape Columbine (33°S) and Cape

Agulhas where the species occurs between 40 m and 155 m. Two isolated occurrences

off the Namaqualand coast in 88 m and 205 m.

The modern populations are most abundant on the Orange Shelf at c. 160 m, and

off the south-western Cape at c. 100 m.

Relict specimens of P. lepralioides occur on the Walvis Ridge shelf (19°S) and the

eastern Agulhas Bank (Figs 5, 52). The greatest abundance of relict specimens lies on

76 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 5

Water depth distribution of Pseudokeijella lepralioides.

Depth Shift

UDL LDL range

(m) (m) — (m) (m)

MODERN

Orange 126 183 Si

Namaqualand 88 205 117

South-western Cape 40 155 LIS)

RELICT

Walvis 150 300 150 =

Orange 100 240 140 83

Namaqualand 88 310 222 105

South-western Cape 40 220 180 65

UDL — upper depth limit

LDL — lower depth limit

Shift — difference between relict and modern depth ranges

the Orange Shelf and, in contrast to the modern populations, the species is relatively

poorly represented off the south-western Cape, although here relict specimens of

P. lepralioides are an important secondary element. There is a southward decrease in

the UDL from 150 m on the Walvis Shelf to 88 m off Namaqualand, and 40 m off the

Cape Peninsula, and a corresponding, but less marked, decrease in LDL from

c. 300 m on the Orange Shelf and areas farther north, to 220 m in the south.

Across-shelf abundances decrease in the northern area (18—32°S), with major

changes at 160 m and 200 m, but in the south (32—36°S), there is a general increase to

about 200 m, beyond which values decline.

Table 5 and Figure 53 summarize the depth range changes for the various

P. lepralioides populations.

9S S

latitude

250 500m

Fig. 53. Latitudinal water-depth distribution of sites with Pseudokeijella lepralioides

(Brady, 1880).

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA Wi)

Family Xestoleberididae Sars, 1928

Genus Xestoleberis Sars, 1866

Fourteen species of this genus have been recorded around southern and south-

western Africa. Nine of these occur in the present study area: Xestoleberis africana

Brady, 1880—west coast continental shelf to Agulhas Bank; Xestoleberis hartmanni

sp. nov.—continental shelf west of the Cape Peninsula to Cape Agulhas; Xestoleberis

ramosa Miller, 1908—coastal sites from Liideritz to Simonstown harbour (False

Bay); Xestoleberis capensis Miller, 1908—coastal sites from Simonstown harbour to

Knysna Lagoon; Xestoleberis crenulata Klie, 1940—Lideritz Bay; Xestoleberis ferax

Klie, 1940— Lideritz Bay; Xestoleberis baja Klie, 1940—Lideritz Bay; Xestoleberis

humilis Klie, 1940—Lideritz Bay; and Xestoleberis aff. X. rotunda Hartmann, 1964

(Hartmann 1974)—coastal sites Cacuaco (Mozambique) to Liideritz.

Xestoleberis africana Brady, 1880

Figs 54A-E, 56A-B

Xestoleberis africana Brady, 1880: 126, pl. 30 (figs 4a—c). Puri & MHulings 1976, 299, pl. 19

(figs 15-16).

?Xestoleberis sp. B Keeler, 1981: 182-183, pl. 10 (figs 14-15).

Xestoleberis spp. Boomer, 1985: 60-61, pl. 3 (figs 52-53).

Illustrated material

SAM-—PO-MF-0583, RV, TBD 6847, 94 m

SAM-PQ-MF-0584, LV, TBD 6847, 94 m

SAM-PQ-MF-—0585, LV, TBD 6847, 94 m

SAM-—PO-MF-0586, RV, TBD 6847, 94 m

SAM-PQ-MF-0587, C, TBD, 6847, 94 m

Material

500 valves.

Remarks

This distinctively shaped, rather globular, thick-shelled species was re-illustrated

by Puri & Hulings (1976). My SEM photographs show that the ME of the hinge in the

RV and LV is locellate and denticulate, respectively, and not smooth as reported in

the description of the lectotype. The small ‘Xestoleberis’ spot is well illustrated in

Figure 54D.

Distribution

Brady (1880) originally recorded this species only from ‘Challenger’ Station 140

in False Bay (15-20 fm (27-37 m)). The present study shows it to be the most widely

distributed species of the genus on the continental shelf off south-western Africa

(Fig. 57A).

Modern populations of X. africana are restricted to a narrow depth range off the

Cape Peninsula (33,96—34,09°S; 80-95 m) and in False Bay (40 m) (Fig. 57B).

Relict specimens occur over a latitudinal range 22°S (Walvis Bay) to the eastern

Agulhas Bank, and fall into three population groups. Two sites occur in the north

78 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 54. A-E. Xestoleberis africana Brady, 1880, TBD 6847, 94 m. A. SAM—PQ-—MF0583, RV, SEM

2657. B. SAM-—PO-MF0584, LV, SEM 2658. C. SAM-—PQ-MF0585, LV, SEM 2663.

D. SAM-PQ-MF0586, RV, SEM 2652. E. SAM—PQ-—MF0587, carapace, dorsal view, SEM 2661.

F. Xestoleberis hartmanni sp. nov. SAM—PQ-—MF0579, carapace, dorsal view, TBD 6825, 160 m, SEM

2677. Scale bars = 100 microns.

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 79

(vicinity of Walvis Bay), a cluster of sites lies on the Orange—Namaqualand shelf

between the Orange and Olifants rivers, and a third centre extends from the Cape

Peninsula to the eastern Agulhas Bank. The upper depth limits of the sites north of

34°S all lie deeper than 173 m, whereas in the south relict specimens occur as shallow

as 40 m in False Bay. With the exception of one isolated deep site, the lower depth

limits along the whole shelf lie between 283 m and 290 m (Fig. 57C). The deep site (in

545 m) is relatively isolated from the main south-western Cape populations and con-

tains a single, probably allochthonous valve.

Xestoleberis hartmanni sp. nov.

Figs 54F, 55A—D, 56G-H, S, 58

Derivation of name

The species is named for Professor G. Hartmann (University of Hamburg) for his

important contribution to the study of modern marine ostracods around southern

Africa.

Holotype

length height

SAM-—PO-MEF-0578, LV, TBD 6825, 160 m 0,50 mm 0,38 mm

Paratypes

length height width

SAM-—PQ-ME-0579, C, TBD 6825, 160 m 0,51 mm — 0,30 mm

SAM-PQ-ME-0580, RV, TBD 6825, 160 m 0,49mm 0,30 mm —

SAM-PQ-ME-0581, LV, TBD 6825, 160 m 051mm 0,34 mm —

SAM-PQ-MEF-0582, RV, TBD 6825, 160 m 0,45mm 0,29:mm —

Material

20 valves.

Diagnosis

Species with an ‘angular’ aspect resulting from a strongly and asymmetrically

arched DM, with the addition of a truncated posterodorsal margin in the RV. RV and

LV hinge ME are locellate and denticulate, respectively.

Description

External features. LV and RV differ considerably in lateral outline. In both valves

the narrow AM is asymmetrically rounded, ventrally directed and somewhat

extended. In LV the PM is broadly rounded, with a continuous sweep over the

posterodorsal area. In RV the PM is truncated, rounded ventrally, but angular across

the posterodorsal area. The LV DM is strongly arched, with the rounded, highest

point just behind mid-length. This contrasts with the RV DM, which is straight to the

rounded highest point (lying just anterior of mid-length), whence the DM sweeps

downward to the AM. In both valves the VM is slightly convex. In dorsal view the

carapace is lemon-shaped: distinctly acuminate anteriorly, and more rounded, but

80 ANNALS OF THE SOUTH AFRICAN MUSEUM

apap ihe:

Fig. 55. A-D. Xestoleberis hartmanni sp. nov., TBD 6825, 160 m. A. SAM—PQ-MF0578, holotype,

LV, SEM 2673. B. SAM—PQ-MF0580, RV, SEM 2674. C. SAM-PO-MFO0581, LV, SEM 2678.

D. SAM-PO-MF0582, RV, SEM 2681. E-F. Pseudokeijella lepralioides (Brady, 1880), TBD 6836, 80 m.

E. SAM—PQ-MF0559, RV, SEM 2385. F. SAM—PQ-MF0560, LV, SEM 2380.

Scale bars = 100 microns.

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 81

A K L

ferax

africana

Cc D M N

baja

ramosa

E F O Pp

capensis

> = / | |

hartmanni

capensis hartmanni

crenulata ramosa

g. 56. Outlines of Xestoleberis species from southern Africa. A-B. X. africana (Brady, 1880),

TED 6847, 94 m. A. SAM—POQ-MF0574, LV. B. SAM—POQ-MF0573, RV. C-D, Q. X. ramosa

Miller, Simonstown harbour, traced from ‘Miller UMS WAN), C IY. D. RV. Q. Carapace, dorsal

view. E- F, R. X. capensis Miller, Simonstown harbour, traced from Miiller (1908: 127). E. LV.

F. RV. R. Carapace, dorsal view. G—H, S. X. hartmanni sp. nov., TBD 6825, 160 m. G.

SAM-PQ—-MEF0578, holotype, LV. H. SAM-PQ- MF0580, RV. S. SAM-PO- MF0579, carapace,

dorsal view. I-J. X. crenulata Klie, Liideritz Bay, traced from Klie (1940, figs 44-45). I. Dyae RV:

K-L. X. ferax Klie, Liideritz Bay, traced from Klie (1940, figs 51-52). K. LV. L. RV. M-N. X. baja

Klie, Lideritz Bay, traced from Klie (1940, figs 57-58). M. LV. N. RV. O-P. X. humilis Klie, Liide-

ritz Bay, traced from Khie (1940, figs 61-62). O. LV. P. RV. Scale bars = 200 microns; other scales

not known.

humilis

somewhat drawn out posteriorly. Valve surface is smooth, with numerous, distinct,

but small normal pore openings.

Internal features. Marginal areas are relatively narrow, but in the material avail-

able details were not well preserved. The hinge is antimerodont, with relatively short

terminal elements, with a locellate ME in the RV. No unambiguous views of the MS

were obtained because of generally poor preservation, but the adductors consist of

four elongate scars, with a small U- or V-shaped anterior scar and a further small ven-

trally adjacent scar (Fig. 58). The ‘Xestoleberis’ spot is indistinct, dorsally situated and

small. There is no well-developed eye socket.

Remarks

None of the other locally occurring species of Xestoleberis possesses a comparably

‘angular’ outline to X. hartmanni (Fig. 56). Although the outline of the RV of

X. ramosa is similar, the DM outline of the LV of Miller’s species is broadly

82 ANNALS OF THE SOUTH AFRICAN MUSEUM

A 15 longitude 20 25 B

33,5

a, ‘ Hout Bay

o #/ oa

‘ is False Bay

NO 4

“

C. Point

34,0

34,5

ory

latitude

°S

latitude

) 0,5 1,0km

Fig. 57. A. Distribution of Xestoleberis hartmanni sp. nov. (squares) and X. africana Brady, 1880

(crosses). See Fig. 7 for abbreviations. B. Distribution of X. hartmanni sp. nov. (squares) and_X. afri-

cana Brady, 1880 (crosses) off the south-western Cape. C. Latitudinal water-depth distribution of sites

with X. hartmanni sp. nov. (crosses) and X. africana Brady, 1880 (squares).

rounded, and in dorsal view is more elliptical, with rounded extremities. The other

continental shelf species, X. africana, has an overall rounded and inflated appearance

and cannot be confused with X. hartmanni.

Distribution

Xestoleberis hartmanni is limited to the waters off the south-western Cape. One

site only contains modern specimens (15 m in Hout Bay).

“an &

Fig. 58. Xestoleberis hartmanni sp. nov., SAM—

PO-MF0581, LV, MS, TBD 6825, 160 m.

Scale bar = 100 microns.

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 83

Relict populations occur at three sites off the Cape Peninsula (42-160 m), and at

two further isolated, much deeper locations: south-west of Saldanha (990 m), and

south-west of Cape Point (545 m) (Fig. 57B). The species reaches a maximum abun-

dance of 6 per cent total ostracod population in 160 m off the Cape Peninsula.

7 eee

D. exilis

?P. walvisridgensis ae

Earetiberisis [cae aera

hee

P. walvisbaiensis ee Ew |

Eat

a eae

H. melobesioides [art aa ere

B. k. knysnaensis os ee is ee ea ae ee

kK. angulata Sa ae

R. cytheropteroides

P. lepralioides

P. lacrimata

A. (A.) levetzovi

A. (A.) flabellicostata

X. africana

ieee ee Ee

N. boomeri

A. (A.) keeleri

X. hartmanni

P. subrhomboidea

C. dromedaria

Walvis Walvis Luderitz Orange Saldanha Cape

Ridge Bay River Peninsula

Fig. 59. Modern and relict latitudinal ranges of dominant ostracod taxa on the continental margin off

south-western Africa. Thick lines = modern range, thin lines = relict range.

ANNALS OF THE SOUTH AFRICAN MUSEUM

en ee EEE nnn UE

Saploisaqojaul

Dyjamoycuuay]

sapioyvsda]

Dyjallayopnasd

sisuapuskuy

pluosuag

sapiosajdosayjao

DUAISSnY

LOITAa

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9 H1dVL

QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 85

SUMMARY

Figures 4, 5 and 59 and Table 6 summarize the distribution of the dominant ostra-

cod taxa on the continental margin of south-western Africa.

On a regional scale there is dominance by two groups, with an area of overlap in

the Walvis Bay—Orange River zone: loxoconchids—Cytherella—Bensonia form a north-

ern assemblage, and R. cytheropteroides and P. lepralioides dominate in the south.

This regional pattern occurs in both relict and modern assemblages. Henryhowella is

locally dominant in both northern and southern sectors.

A comprehensive discussion of the distribution of the ostracod taxa on the conti-

nental shelf off south-western Africa (both dominant and minor forms) will be given

in Dingle (in press) and Dingle & Girandeau (in press).

ACKNOWLEDGEMENTS

The samples on which this study is based were collected while the author was

Director of the Marine Geoscience Unit, University of Cape Town. Fellow scientists

and crew of the University’s then-research vessel “Thomas B. Davie’ are thanked for

their dedication and endurance afloat. The following agencies funded sea-time:

University of Cape Town, Geological Survey, SANCOR, and FRD. Some of the lab-

oratory work was undertaken while the author was on sabbatical leave as Visiting

Professor at University College London, and I gratefully acknowledge facilities pro-

vided by Professors M. Audley-Charles and A. R. Lord. Professor G. Hartmann

(University of Hamburg) is thanked for the loan of paratypes of Palmoconcha walvis-

baiensis (Hartmann, 1974). My colleague, Professor R. C. Whatley (University

College Aberystwyth) is thanked for advice on taxonomy, and the manuscript has

been improved through the constructive criticism of the following referees: Professors

Whatley and A. R. Lord, and Dr H. J. Oertli (Pau, France).

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QUATERNARY OSTRACODS FROM SOUTH-WESTERN AFRICA 89

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Espanola de Micropaleontologia 20 (1): 5-37.

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6. SYSTEMATIC papers must conform to the International 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°5S1’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. Du Toit 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

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

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FiscHER, P. H., DuvaL, 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. 1960b. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bulletin of

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ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 102 °#£Band

October 1992 Oktober

Part 2 Deel

TAXONOMIC REVISION OF

AFRICAN CYPRIDINI. PART II.

DESCRIPTION OF RAMOTHA GEN. NOV.

(CRUSTACEA, OSTRACODA)

K. MARTENS

Cape Town Kaapstad

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D1619

TAXONOMIC REVISION OF AFRICAN CYPRIDINI

PART II. DESCRIPTION OF RAMOTHA GEN. NOV.

(CRUSTACEA, OSTRACODA)

KOEN MARTENS

Koninklijk Belgisch Instituut voor Natuurwetenschappen, Zoetwaterbiologie

(Royal Belgian Institute of Natural Sciences, Freshwater Biology),

Vautierstraat 29, B—1040 Brussels, Belgium

(With 17 figures and 1 table)

[MS accepted 31 July 1991|

ABSTRACT

Ramotha gen. nov. is described to accommodate a number of African species, previously assigned

to Cypris, Eucypris and Strandesia. It belongs to the Cypridini and in this tribe is characterized by a T1

with a divided penultimate segment, by a RV with an inwardly displaced frontal selvage and by a LV

without a frontal selvage but with a large inner list. The type species of the new genus is R. hirta (Sars)

from the Western Cape. Ramotha curtisae sp. nov. is described from bisexual populations, originating

from Namibia and Zimbabwe. Ten other species are also transferred to this genus and are redescribed.

Eucypris bouilloni Kiss is furthermore placed in the synonymy of R. crassa (Klie). The males of

R. hirta (Sars), R. producta (Sars) and of R. trichota (G. W. Miller), thus far unknown, are here

described. Representatives of this new genus are found mainly in temporary bodies of water in East

and southern Africa and all species have restricted geographical distributions.

CONTENTS

PAGE

JFMTROGMICTOM o.oo. 8.8 58 bee O Bis Ou OneteCcEs CHEN aera NUM an st ts Anat Se a a 91

Wicakemict lec Genie GOSH tat retie reek, ele eh atl nny canes Meee eee nee Scnls Sue anai aller al due ave ai aie 92

RaxOnOmiCiG eSCHPHONS 4) aon. mo aeeiie rine tease cemiem eae init oun erties ald ane eB 95

IC VELORGH EIS PE CICS pea eR a a sc AN PE Ne ete waht che’ sv sieit ce oars 126

DISCUSSIONMINE eee YR tee a eae Mets ead Ue ou Gl ones Ne Ol esas 7)

haxonomyzandsZOOLCOORAPNY ae 2 saci ee oe Sela ce Acc na needs aoe eee kD,

OOO I oe ie Brcero cha eae ca catgol Gi heey Onn ns lee agar eC CONE oe 129

PXCKMOMICEO CTR CIES terra tay eras MNase UE OME a5 ret oye ele eleia's 0.5 129

[SENCTRETIGES oe. bra: aieitord tem Sher Ga seea 1S ByOee e o reeri Une eae ga ee aA 130

INTRODUCTION

Sars (1895) described Cypris corpulenta and Cypris trigona from a swamp near

Knysna (South Africa, Cape Province). Later, Sars (1924) transferred these and the

species meanwhile described by Miiller (1908, 1914) to the genus Eucypris, and added

three more taxa to the list. Related taxa were reported from East African water

bodies by Lowndes (1931), Klie (1939, 1944), Lindroth (1957) and Loffler (1968).

Upon re-investigation of the types of Strandesia crassa Klie, it was shown that this

species does not belong in the Cypricercinae, but rather has characters shared by the

above taxa.

Ann. S. Afr. Mus. 102 (2), 1992: 91-130, 17 figs, 1 table.

92 ANNALS OF THE SOUTH AFRICAN MUSEUM

All these species are united by a number of features, which seem to place them

between the genera Cypris and Eucypris, as interpreted in a classical sense, hence the

confusion in the older literature. However, they do not belong in Cypris s.s., because

they possess an inwardly displaced selvage in the RV only (in both valves in Cypris)

and have the penultimate segment of the T1 divided (fused in Cypris). For the latter

reason, Sars (1924) removed these taxa to Eucypris. It is here shown that they do not

belong to this genus either, as they lack a ‘c’-seta on the Mx2, which is typical of all

Eucypridini (Martens 1989).

The genus Ramotha gen. nov. is here described to comprise all of the above

species, as well as one new species, and is assigned to the Cypridini, because the hemi-

penis (in species where males are known), displays the ‘8’-shaped coils of the inner

(post labyrinth) spermiductus, which is typical of this tribe. A re-assessment of the

subfamily Cypridinae and its nominate tribe is presented in the first part of this

revision (Martens 1990). Apart from Ramotha gen. nov., the Cypridini to date also

comprise the genera Cypris, Pseudocypris and Globocypris.

The phyletic lineage analysed in the present paper constitutes a good example

of the present chaos and confusion in African ostracod taxonomy and illustrates the

difficulty of identifying African ostracods with the literature as it stands. The pres-

ent genus has 12 nominal representatives (of which one is a synonym), originally

described in no less than three different genera and subfamilies!

MATERIAL & METHODS

List of museums and collections

AM Albany Museum, Grahamstown, RSA.

GW Ernst-Moritz-Arndt-Universitat, Greifswald, Germany.

KBIN- Koninklijk Belgisch Instituut voor Natuurwetenschappen, Brussels,

Belgium.

MRAC Koninklijk Museum voor Midden-Afrika, Tervuren, Belgium.

OSLO Zoological Museum, Oslo, Norway.

UPPS Uppsala Universitet, Zoologiska Muset, Uppsala, Sweden.

SAM _ South African Museum, Cape Town, RSA.

SMN State Museum of Namibia, Windhoek, Namibia.

ZIZM_ Zoologisches Institut und Zoologisches Museum, Hamburg, Germany.

Abbreviations used in text and figures

Ai = antennula; A2 = antenna; be = bursa copulatrix in hemipenis; Cp = cara-

pace; db = dorsal branch of furcal attachment; di = distal end of unbranched furcal

attachment; H = height of valves; il = inner list; im = inner margin; L = length of

valves; lc = line of concrescence; Is = lateral shield of hemipenis; LV = left valve; Md

= mandibula; ms = medial shield of hemipenis; Mx1 = maxillula; Mx2 = maxilla; pr

= proximal end of furcal attachment; R = Rome organ; RV = right valve; sl =

selvage; Tl = first thoracopod; T2 = second thoracopod; vb = ventral branch of

furcal attachment; vm = valve margin; W = width of valves.

Chaetotaxy of the limbs follows the model proposed by Broodbakker & Danielo-

pol (1982), and revised for the A2 by Martens (1987).

REVISION OF AFRICAN CYPRIDINI 93

TAXONOMIC DESCRIPTIONS

Class OSTRACODA Latreille, 1806

Subclass Popocopa G. W. Miiller, 1894

Order PODOCOPIDA Sars, 1866

Family Cyprididae Baird, 1845

Subfamily Cypridinae Baird, 1845

Tribe Cypridini Baird, 1845

Genus Ramotha gen. nov.

Derivation of name

The new genus is named after Ramoth, last remaining golden queen dragon at

Benden Weyr on the planet Pern. This is, of course, a tribute to her creator, Miss

Anne McCaffrey. Gender of the new genus is feminine.

Type species. Eucypris hirta Sars, 1924.

Diagnosis

A cypridinid genus, with mostly large (2-3 mm) and globular species. RV with

frontal and caudal selvage inwardly displaced over a large distance, but with anterior

valve margin ventrally not with lip-like extension. LV without inwardly displaced

selvage, but with a large inner list, situated in the centre of a wide anterior calcified

inner lamella and on the posterior inner margin; ventrally with a conspicuous outer

list. T1 with penultimate segment divided and with seta d: 2—3 times as long as seta do.

Additional generic features

Caudal valve margin of the RV crenulated. Al with Rome organ small. A2 with

natatory setae extending well beyond tips of claws; in males with a sexual dimorphic

chaetotaxy of apical claws and setae, showing a supplementary reduction of claw Gi.

Gamma seta on Md palp long, relatively slender and hirsute. Mx1 palp with terminal

segment elongated. Mx2 without ‘c’-seta. T2 with distal segments pincer-shaped.

Furca and furcal attachment well developed.

Other species

Ramotha capensis (G. W. Miller, 1908), R. corpulenta (Sars, 1895), R. crassa

(Klie) (syn.: R. bouilloni (Kiss) syn. nov.), R. curtisae sp. nov., R. kenyensis (Lind-

roth), R. montana (Lindroth), R. producta (Sars, 1924), R. purcelli (Sars, 1924),

R. trichota (G. W. Miller, 1908), and R. trigona (Sars, 1924).

Taxonomic position

The genus belongs to the Cypridini, because of the presence of the ‘8’-shaped

loops of the inner spermiductus in the hemipenis. It differs from Cypris s.s. and from

Pseudocypris because it has the penultimate segment of the T1 divided; it furthermore

lacks an inwardly displaced selvage on the LV, whereas the selvage on the RV has a

different shape. Pseudocypris furthermore has a marginal selvage on both valves, and

94 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 1. Original illustrations of some South African species of Ramotha (all redrawn after Sars 1924—

Annals of the South African Museum). A-B. R. producta (L = 2,40 mm). C-D. R. corpulenta

(L = 2,10 mm). E-F. R. hirta (L = 1,90 mm). G-H. R. trigona (L = 1,75 mm). I-J. R. trichota

(L = 3,30 mm). K-L. R. capensis (L = 1,80 mm). M-N. R. purcelli (L = 2,30 mm). A, C, E, G, I,

K, M: Cps in left lateral view. B, D, F, H, J, L, N: Cps in dorsal view.

REVISION OF AFRICAN CYPRIDINI 95

most species of this genus display large lateral ala on the valves. Globocypris also has

a large frontal selvage on the LV.

Ramotha hirta (Sars, 1924)

Figs 1E-F, 14H-L, 16H (parthenogenetic populations)

Figs 2-4, SA—H (bisexual populations)

Cypris corpulenta G. W. Miller, 1908: 150-151, figs 1-6 (non Sars, 1895).

Eucypris hirta Sars, 1924: 110-111, pl. 3 (figs 5-6).

non Eucypris hirta Lowndes, 1931: 1292 (= R. crassa (Klie)).

non Eucypris hirta Klie, 1944: 17-18 (= R. montana (Lindroth)).

Ramotha elephantina Martens, 1990: 159 (nomen nudum).

Type locality

Pond on Green Point Common, near Cape Town (South Africa).

Type material

SAM-A11116: c. 20 good 2 from the above locality (tube labelled ‘Cape of

Good Hope’); these specimens are here considered the syntypes of this species. Lecto-

type (here designated): a 9, selected from the above syntypes, with soft parts

dissected in glycerine on a sealed slide and with valves stored dry (SAM-—A40043); all

other syntypes become paralectotypes.

Other material investigated

Parthenogenetic populations. SAM-—A11115, 1 dissected 9 from Cape of Good

Hope. SAM-A11119, 1 9 from Cape of Good Hope. SAM-—A11283, 2 9, labelled

‘SAM EEF’. SAM-A11284, 2 9 from Cape Flats. SAM-A11285, 1 9, no locality

given. (All identified by G. O. Sars.)

Bisexual populations. 18 9 and 2 C raised from dried mud, collected from small

pools in granite on Olifantsberg near Leopoldsville (approx. 32°12’S 18°25'B),

Western Cape Province, Republic of South Africa. Mud collected by Dr J. A. Day

(original collections ‘02’, ‘C4’ and ‘SWT2’) between June 1980 and July 1981. Deposi-

tion: 1 dissected o' (SAM-—A40038), 1 dissected 92 (SAM—A40039), 1 O’ and 1 9

(both dissected) (KBIN OC1489-1490), 1 C’ and 2 9 stored dry (after use for SEM

illustration—MRAC.56779-56780), the remaining specimens in toto in_ spirit

(KBIN OC1491).

Diagnosis (parthenogenetic populations)

An easily recognizable species with valves densely set with long hairs. Cp in

dorsal view (Fig. 14L) with anterior edge beak-like and asymmetrically pointed to the

right side. Valves in lateral view (Fig. 14H—J) relatively high and short. Mx2 palp in

some specimens with an additional lateral seta (Fig. 16H). Furcal attachment distally

branched.

Measurements (in wm, mean + S.D., n = 5)

L = 1 472 + 40; H = 931 + 24; W = 896 + 27; W/L = 0,59-0,63.

96 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 2. Ramotha hirta (Sars), C’, (SAM-—A40038). A. A2. B. Mx1, showing part of chaetotaxy. C. Al.

D. T2. E. Md palp. F. furcal attachment. Scale: 156 wm for A, C, D, F; 81 wm for B, E.

REVISION OF AFRICAN CYPRIDINI 97

Diagnosis (bisexual populations)

Medium-sized species, with ventral margin (including inner list and selvage) con-

spicuously produced in the first third (especially in the 9). Cp in dorsal view with

greatest width (c. one-half of the length) situated in the first third of the length,

anterior tip concavely and asymmetrically produced and posterior margin bluntly

pointed. Furcal attachment in both sexes without dorsal branch. Prehensile palps

asymmetrical, the right one being the largest. Hemipenis with lateral shield protruding

towards the ventral side.

Measurements (in wm, mean + S.D.)

@: L=1 724; H = 1 034; W = 1 000 (m = 1).

©: L=1755 + 104; H = 1 083 + 66; W = 1 069 + 35; W/L = 0,61-0,66 (n = 5).

Additional description of C

RV (Fig. 5B) with anterior margin more broadly rounded than posterior one;

dorsal margin weakly curving and sloping towards the caudal side, merging with that

margin without an obvious angle; towards the frontal side weakly indented at about

one-third from the front; ventral margin nearly straight, apart from the conspicuous

curve of the selvage and valve margin in the anterior third. Calcified part of the inner

lamella relatively narrow, on both anterior and posterior sides with traces of a blunt

inner list; valve margin on both posterior and anterior sides set with a series of short

ridges (Fig. 5G). Central muscle scars with mandibular scars large and with scar

pattern of adductor muscles conforming to that of the subfamily.

LV (Fig. 5A) with anterior margin more broadly rounded than posterior one;

dorsal margin weakly curving and passing into the caudal margin with a blunt angle,

no anterior indentation; ventral margin somewhat more sinuous than in the RV and

with matching curves of valve margin and inner list. Caudal calcified inner lamella

somewhat wider than in the RV and with a large inner list positioned near the inner

margin. No ridges on the valve margin.

Cp in ventral view with RV reaching beyond LV anteriorly and with LV overlap-

ping RV on the ventral side. Greatest width situated in the anterior third of the Cp.

Surface densely pitted and set with short setae.

Al (Fig. 2C) seven-segmented and typical of the subfamily. Terminal segment

c. 1,5 times as long as its greatest width, all natatory setae long.

A2 (Fig. 2A) with natatory setae reaching well beyond tip of terminal claws;

aesthetasc Y short. Apical chaetotaxy (Fig. 3A) with typical sexual dimorphic charac-

ters (see Martens 1987), but with claw Gi: short; seta of y3 longer than the aesthetasc.

Md with coxa elongated (Fig. 3H). Md palp (Fig. 2E) four-segmented. First

segment with large respiratory plate (exopodite—not shown in the figure) and a group

of four apical setae: one long and smooth, one short, narrow and smooth (alpha-seta),

the latter flanked by one large ‘s’-seta, set with a double row of setulae, and by one

short ‘s’-seta. Second segment with two groups of apical setae—an internal group con-

sisting of a short and hirsute beta-seta, three long and smooth and one long and

barbulated setae; external group consisting of two long and one shorter setae. Penulti-

mate segment with six subapical external setae (three long and three shorter) and one

Narrow and hirsute gamma-seta of intermediate length, as well as three subequal

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REVISION OF AFRICAN CYPRIDINI 99

Fig. 4. Ramotha hirta (Sars). A. O’, right Mx2 (respiratory plate not shown) (SAM-—A40038). B. CO’,

left prehensile palp (SAM—A40038). C. 2, Mx2 (only part of chaetotaxy shown) (KBIN OC1490).

D. 9, palp of other Mx2 of same specimen as in C (KBIN OC1490). E. 0’, hemipenis, detail of part

of inner anatomy (KBIN OC1489). F. ©’, hemipenis (KBIN OC1489). G. 9, genital lobe

(SAM-A40039). Scale: 156 wm for C, D, F, G; 81 wm for A, B, E; 33 um for details in A and B.

ANNALS OF THE SOUTH AFRICAN MUSEUM

REVISION OF AFRICAN CYPRIDINI 101

apical setae on the internal side. Terminal segment with four claw-like and with two or

three short and slender setae, all apically inserted.

Mx1 with three endites, a two-segmented palp (Fig. 2B) and a large respiratory

plate. Third endite with two smooth apical claws and with a long and stout lateral

seta, apart from the normal apical setae. First segment of palp (Fig. 3G) with six sub-

apical and subequal setae; terminal segment elongated, 2,5—3 times as long as its basal

width, carrying six apical claw-like setae.

Mx2 with asymmetrical palps, a large endopodite with typical chaetotaxy con-

forming to the subfamily and a respiratory plate (exopodite) with six plumous rays

(five long and one short—not shown in the figures). Endopodite (Fig. 4A) elongated,

carrying two short ‘a’-setae, one larger, central ‘b’-seta and one lateral ‘d’-seta, ‘c’-seta

absent. Apical chaetotaxy consisting of c. 15 setae of different size and shape.

Right prehensile palp (Fig. 4A) with elongated basal segment, carrying two apical

sensory outgrowths; terminal segment three-dimensionally curved; when flattened on

a Slide broad, with a narrow apex and with distal margin showing a wide, blunt angle,

apically with one broad sensory organ.

Left prehensile palp (Fig. 4B) with basal segment similar to that of the right palp;

terminal segment narrower, with a proximal blunt tooth, this segment gradually nar-

rowing towards the tip, the latter bearing a single sensory organ.

T1 (Fig. 3C) with seta d: more than three times as long as do; all segments short,

wide and hairy; penultimate segment divided; apical claw stout and only in its distal

half set with a double row of spines. Second segment with one apical seta. Seg-

ment 3A with one subapical seta; segment 3B with one large and one minute apical

setae. Fourth segment with one lateral seta and, apart from the apical claw, with one

other apical claw-like seta.

T2 (Fig. 2D) a cleaning limb with an apical pincer and without further special

features.

Furca (Fig. 3F) with ramus narrow and curved, carrying two claws and two setae

and with ventral margin serrated with minute setulae. Furcal ramus (Fig. 2F) in this

species consisting of a single ramus, not distally split into a dorsal and a ventral

branch.

Rake-like organs (Fig. 3D—-E) T-shaped, but with apical part swollen, carrying

few blunt teeth.

Hemipenis (Fig. 4F) with a broadly rounded medial shield, asymmetrically

expanded towards the ventral side and a pointed, protruding lateral shield, the latter

boot-like and with the tip pointing in ventral direction. Internal anatomy with the

normal labyrinth, consisting of the elongated parts ‘a’ and ‘c’ and the rounded hinge-

Fig. 5 (see opposite). A-H. Ramotha hirta (Sars). A. O', LV, internal view (KBIN OC1489). B. oO’,

RV internal view (KBIN OC1489). C. 9, LV, internal view (KBIN OC1490). D. 9, RV, internal

view (KBIN OC1490). E. 2, Cp, ventral view, detail of anterior edge (MRAC.56779). F. 2, Cp,

ventral view (stereo-pair, MRAC.56779). G. CO’, tilted RV, internal view, detail of posterior margin

(KBIN OC1489). H. 9, Cp, dorsal view (MRAC.56779). I-O. Ramotha purcelli (Sars), paralecto-

types. I. 9, LV, internal view (SAM-A40046). J. 2, RV, internal view (SAM-A40046). K. 9, Cp,

dorsal view (SAM-—A40048). L. 2, Cp, ventral view (SAM-A40048). M. 9, Cp, dorsal view, detail of

anterior edge (SAM-—A40048). N. Q, tilted RV, internal view, detail of posterior edge

(SAM-—A40046). O. 9, RV, internal view, detail of central muscle scars (SAM-—A40046).

Scale: 1 563 wm for A-D, F, H; 1 320 um for I-L.

102 ANNALS OF THE SOUTH AFRICAN MUSEUM

joint ‘b’, followed by the 3—5 ‘8’-shaped coils of the inner spermiductus, situated dis-

tally from the labyrinth, the sclerotized semi-circular loop and the various hollow

trabeculae leading to the bursa copulatrix (Fig. 4E).

Four pairs of testical tubes present.

Additional description of @

All valve features (Fig. 5C—D) basically as in the male, but valves generally

slightly larger; ventral outgrowth even more pronounced than in the male and caudal

inner list in the LV situated more towards the interior.

Cp in dorsal and ventral view (Fig. SE—H) with greatest width situated in the

anterior third; RV frontally reaching beyond LV.

A2 (Fig. 3B) basically as in the male, apart from the normal sexual dimorphism

in the apical chaetotaxy.

Mx2 (Fig. 4C—D) with palp undivided, carrying the normal unequal three apical

setae, but also 1-2 supplementary lateral setae.

T1 somewhat plumper and heavier than in the male.

Genital lobe (Fig. 4G) undivided and without specific characteristics.

Ovaria on both sides curved upwards.

Discussion

The shape and the hairy aspect of the valves allow an easy identification of this

species within the genus. Nevertheless, confusion exists in the older literature.

G. W. Miller (1908) erroneously reported specimens of this species as Cypris cor-

pulenta (see p. 105). Lowndes (1931) and Klie (1944) both reported this taxon from

East African inland waters. The material of the latter author was available for re-

examination (ZIZM, Hamburg) and, although heavily decalcified, could still be iden-

tified as belonging to R. montana (Lindroth) (see p. 113).

Lowndes (1931) mentioned that he had only a few specimens in bad condition

and therefore no attempt was made to obtain his material. Relying on the locality

from which his material originated (Elgon), it is very likely that he was actually

dealing with R. crassa, reported from this area by Klie (1939) and Kiss (1959), and his

specimens are here tentatively referred to the latter species.

Remarks

It is noteworthy that the masticatory processes of both mandibular coxae are

always situated in the space created by the ventral bulges in both valves. Undoubt-

edly, these ventral bulges are adaptations to the species’ feeding strategy and

behaviour, which unfortunately remain unknown to date.

Ramotha capensis (G. W. Miller, 1908)

Figs 1K-L, 6E-J, 7A-F

Cypris capensis G. W. Miller, 1908: 153-154, figs 1-6; 1914: 70.

Eucypris capensis Sars, 1924: 112, pl. 3 (figs 9-10).

Type locality

Zeekoevlei, Plumstead, near Simonstown (South Africa).

REVISION OF AFRICAN CYPRIDINI 103

Type material

GW II 24627(4): c. 80 9 in spirit in a tube, labelled ‘Cypris capensis, Gaus Expe-

dition (paratypus)’. Lectotype (here designated): a 9, with soft parts dissected in

glycerine on a sealed slide and with valves stored dry (no. GW.24627b, c), selected

from the above syntypes; all other syntypes become paralectotypes.

Other material investigated

GW II 25131(70): c. 50 Q in spirit in a tube, labelled ‘Siid-Afrika, Deutsche Siid-

polar Exp.’ (det. G. W. Miiller). SAM-—A11288: 16 Q in spirit in a tube, labelled

‘SAM EE’ (det. G. O. Sars) (used for descriptions: SAM—A40051-40052). SAM-—

A11927: various 9 in spirit in a tube labelled ‘Valkenberg vlei, coll. 15 July 1922 by

Stephensen’ (det. D. H. Eccles—these specimens smaller). KBIN OC1492 (G501/18):

1 Q in spirit from Wiesdrif, Western Cape Province (approx. 34°40'12”S 19°54'47"E)

(coll. Dr J. King, 11 May 1989) (det. K. Martens).

Diagnosis

In lateral view (Fig. 7A—B), rather elongated and with a sinuous ventral margin.

In dorsal view (Fig. 7E—F) with greatest width (c. two-thirds of length) situated in the

middle, with both anterior and posterior edges with strong beak-like extension.

Ventral outer list on LV strong (Fig. 7C—D); anterior, ventral and posterior valve

margins of RV set with internal ridges. Valve surface densely pitted; ‘poren-warzen’

present.

Mx2 palp in some specimens with an additional lateral seta (Fig. 61); furcal

attachment distally branched (Fig. 6F).

Measurements (in wm, mean + S.D., n = 5)

L=1791 + 25;H =1014 + 19; W = 1179 + 26; W/L = 0,65-0,68.

Discussion

This species is easily recognizable and can be distinguished from its congeners by

the shape of the valves in lateral view, but especially by the morphology of the cara-

pace in dorsal and ventral views. Ramotha capensis appears to be restricted to the

Western Cape Province.

Ramotha corpulenta (Sars, 1895)

Figs 1C-D, 6A-D, 7G—J

Cypris corpulenta Sars, 1895: 30-32, pl. 5 (figs 2a—c).

Eucypris corpulenta Sars, 1924: 110, pl. 3 (figs 3-4).

Type locality

Swamp near Knysna, east of Cape of Good Hope (South Africa).

Type material

SAM-A11114: c. 20 9 raised from dried mud, collected from the type locality.

Lectotype (here designated): a 9, selected from the above syntypes, with soft parts

104 ANNALS OF THE SOUTH AFRICAN MUSEUM

: ea na

—=+-_—> I

Fig. 6. A-D. Ramotha corpulenta (Sars), 2, (SAM-—A40049). A. Furcal attachment. B. T1. C. Mx2.

D. Furca. E-J. Ramotha capensis (G. W. Miller), 9, (SAM—A40051). E. Furca. F. Furcal attach-

ment. G. Distal segment of T2. H. T1. I. Mx2 palp. J. Genital lobe. Scale: 156 wm for A-F, H—J;

81 um for G.

REVISION OF AFRICAN CYPRIDINI 105

dissected in glycerine on a sealed slide and with valves stored dry (SAM-—A40049); all

other syntypes become paralectotypes.

Diagnosis

Large and semi-globular species, with dorsum highly arched in lateral view

(Fig. 7G—H) and with ventral margin only weakly sinuous. RV caudally produced,

with few ridges on the caudal valve margin only. Cp in ventral view (Fig. 7J) with

anterior end beak-like, but less so than in R. capensis; posterior edge convex; greatest

width about two-thirds of length and situated at about one-third from the front. Valve

surface pitted, but less so and with shallower pits than in R. capensis; “‘poren-warzen’

present. Ventral outer list on LV strongly produced. Mx2 palp in some specimens with

an additional lateral seta (Fig. 6C); furcal attachment distally branched (Fig. 6A).

Measurements (in wm, mean + S.D., n = 5)

L = 1 924 + 109; H = 1 141 + 55; W = 1 162 + 86; W/L = 0,58-0,62.

Discussion

Sars’ (1924) illustrations of this species were somewhat inaccurate, as he failed to

show the caudally produced RV and illustrated the carapace too globular and wide.

G. W. Miller (1908) identified some specimens from Zeekoevlei as Cypris corpu-

lenta and provided illustrations. Relying on the shape and the size of the carapace

(1,9-2,0 mm), we can unequivocally refer these specimens to Ramotha hirta (rede-

scribed above). This species was at that stage still unknown, which makes Miller’s

error understandable. Ramotha hirta is furthermore a typical Western Cape species,

whereas R. corpulenta is only known from the vicinity of Knysna (Southern Cape

region).

Ramotha crassa (Klie, 1939)

Fig. 8A-G

?Eucypris hirta Lowndes, 1931: 1292 (non Sars, 1924).

Strandesia crassa Klie, 1939: 132-134, figs 44-47.

Eucypris bouilloni Kiss, 1959: 5-7, fig. 2 (1-10) syn. nov.

Type locality

Small lake near camp 4 of Elgon; small lake of Cladera near Elgon (Kenya).

Type material

Ramotha crassa: ZIZM 871: c. 20 Q in glycerine in a tube, labelled ‘Strandesia

crassa n. sp., Omo Exp. Calderasee des Elgon 7 Januar 1933 coll. Chappuis’. These

specimens are generally recognizable, but all carapaces are completely decalcified

(hence no SEM micrographs could be provided) and crushed. Lectotype (here desig-

nated): a 9, selected from the above syntypes, with soft parts dissected in glycerine on

a sealed slide and with valves stored dry (ZIZM/CR.871a/1); all other syntypes

become paralectotypes.

Ramotha bouilloni: MRAC 47352-47359: 5 decalcitied 9 in a 1ube, labelled ‘Mt.

Elgon, 2e lac, alt. 3 780 m, coll. Dr J. Bouillon—14.12.1953’, belonging to at least

ANNALS OF THE SOUTH AFRICAN MUSEUM

MG{{

REVISION OF AFRICAN CYPRIDINI 107

two different genera. As R. bouilloni is here synonymized with R. crassa, no lectotype

is here designated for this nominal species.

Diagnosis

Valves short and high, greatest height situated at about one-third from the front,

dorsal margin straight, sloping towards the caudal side and passing into the caudal

margin with a blunt angle on the LV (Fig. 8A—B). In dorsal view (Fig. 8C), frontal

edge pointed, with LV reaching beyond RV, greatest width (c. four-sevenths of

length) situated just in front of the middle. The ‘a’-setae on Mx2 sometimes fused at

their base (Fig. 8F—G).

Measurements

Klie (1939: 134) gave the following measurements: L = 1,6 mm; H = 0,96 mm;

W = 0,92 mm. New measurements (in wm, mean + S.D., n = 5): L = 1528 + 36;

= 917 + 19;

Discussion

This species is closely related to R. montana, but is much smaller and has rela-

tively higher valves. It was originally described from Mount Elgon (Klie 1939). From

the same region, Kiss (1959) described two other species: Eucypris montelgoni and

E. bouilloni. The type material of both nominal taxa is curated by the MRAC (Tervu-

ren); in both cases, it consists of very few, damaged and decalcified specimens.

The tube labelled ‘E. bouilloni’ contained 5 Q, three of which belong to a species

of Eucypris s.s. (no selvage in RV, no inner list in LV, ‘c’-seta on Mx2), possibly

E. virens; the remaining two specimens belong to R. bouilloni. Upon comparison with

the type material of R. crassa, it was decided that the species were synonymous. Their

similarity was evident from the figures of both nominal taxa. We here formally syn-

onymize R. bouilloni with R. crassa.

A more difficult case is the position of Eucypris montelgoni. The tube thus

labelled (MRAC 47350) contained one crushed 9, similar in appearance to R. crassa

and of approximately the same size, but definitely not referable to what was described

as E. montelgoni. Kiss (1959: 7-8) indeed cited the length of E. bouilloni as 1,7 mm,

but only 0,7 mm for E. montelgoni. We cannot consider the latter to be the larvae of

R. bouilloni (hence R. crassa), because the A2 was illustrated with 5+1 natatory setae

(A-1 larvae—the 8th instar—of Cyprididae have only five natatory setae, A-2 larvae

have four setae, etc.—see below). We are therefore, at present, unable to place

FE. montelgoni conveniently and this will remain impossible until new material of this

Species becomes available.

Fig. 7 (see opposite). A-F. Ramotha capensis (G. W. Miiller), 2. A. LV, internal view (GW.24627b,

c). B. RV, internal view (GW.24627b, c). C. Cp, ventral view (GW.24627d). D. Cp, ventral view,

detail anterior (GW.24627d). E. Cp, dorsal view (GW.24627d). F. Cp, dorsal view, detail of surface

structure (GW.24627d). G-J. Ramotha corpulenta (Sars), 9. G. LV, internal view (SAM-—A40050).

H. RV, internal view (SAM—A40050). I. RV, internal view, detail of central muscle scars

(SAM-—A40050). J. Cp, ventral view (KM.1167—lost). K-M. Ramotha producta (Sars), 2 (8th larval

instar). K. LV, internal view (KBIN OC1499). L. Cp, ventral view (KBIN OC1500). M. RV, internal

view (KBIN OC1499). Scale: 1316 wm for A-C, E, G, H, J; 980 wm for K—M; 714 um for D;

373 wm for I; 82 wm for F.

108 ANNALS OF THE SOUTH AFRICAN MUSEUM

Nias)

Boies

Fig. 8. A-G. Ramotha crassa (Klie), 9 (A—C redrawn after Klie (1939), D-G original from lectotype

specimen no: ZIZM/CR.871a/1). A. LV, external view. B. RV, external view. C. Cp, dorsal view.

D. T1. E. Genital lobe. F. Mx2, showing part of chaetotaxy. G. Mx2, detail of fused ‘a’-setae.

H. Ramotha montana (Lindroth), 9 (UPPS.228(g.1-2)), T1. I. Ramotha kenyensis (Lindroth), 9

(UPPS.601(n.1—-2)), T1. Scale: c. 840 wm for C; c. 670 wm for A, B; 156 wm for D-F, H, I; 33 wm

for G.

REVISION OF AFRICAN CYPRIDINI 109

Ramotha curtisae sp. nov.

Figs 9, 10A—M, 11A—H

Derivation of name

This species is named after Miss Barbara Curtis (Windhoek), curator of fresh-

water invertebrates at the State Museum, Windhoek, Namibia, who has generously

supplied me with Namibian ostracods over the past years.

Type locality

A small farm dam at Joyec (plot 198), Gobabis district, Namibia (approx.

NS SDSS) 8/2):

Type material

SMN 51250: c. 20 good & and Q, collected from the above locality by B. A.

Curtis on 24 June 1986. Holotype: C’, stored in toto in spirit (SMN 51513).

Allotype: 9, with soft parts dissected and with valves stored dry (SMN 51514).

Paratypes: 2 CO dissected and stored as the allotype (KBIN OC1493 and SMN 51515),

2 @ stored dry in toto (after use for SEM—SMN 51516); the bulk of the paratypes

remains in spirit, stored under the same number as the original sample. Deposition:

one dissected C is lodged in the collection of the KBIN (Brussels), the other para-

types, as well as the holotype and the allotype, are returned to the SMN (Windhoek,

Namibia).

Other material investigated

SMN 50921: 1 LV + soft parts of a 2, collected on 26 February 1986 by B. A.

Curtis from a farm dam at Biesiepan (plot 971) at the border of the Grootfontein and

the Otjiwarongo districts (approx. co-ordinates: 20°S 17°E). Accompanying ostracod

fauna: Heterocypris ovularis.

10 000s of good CG and Q in 3 samples from Shopi Pan in Hwange Nature Park,

Zimbabwe (approx. 18°S 24°E), collected on 4 September 1948 (AM/SED.91), 5 Sep-

tember 1948 (AM/SED.98) and 11 September 1948 (AM/SED.36) by Mr and Mrs

Omer-Cooper. Accompanying ostracod fauna in these samples: Pseudocypris circula-

ris, Cypricercus sp. nov., Parastenocypris junodi and Afrocypris barnardi.

Diagnosis

Valves (Fig. 1OA-F) highly arched and sub-triangular, in lateral view with

anterior margin more broadly rounded than posterior one, the latter rather pointed.

Cp in dorsal view with both anterior and posterior edge convexly rounded and with

greatest width situated in the middle. LV with anterior inner list crenulated.

Hemipenis with lateral shield boot-like (Fig. 9E—F), but with tip less elongated

than in R. hirta; medial shield far more pronounced than in the latter species. Prehen-

sile palps asymmetrical; right palp (Fig. 9C) with penultimate segment with a dorsal

angle of c. 90° (this angle greater in the type species).

110 ANNALS OF THE SOUTH AFRICAN MUSEUM

——

—

————$—

= =>

SSS

= emai Yo

Fig. 9. Ramotha curtisae gen. et sp. nov. A. CO, left prehensile palp (KBIN OC1493). B. O’, left pre-

hensile palp, detail of sensory organs on both segments (KBIN OC1493). C. ©’, right prehensile palp

(KBIN OC1493). D. 9, Mx2 palp (SMN 51514). E. Oo’, hemipenis (KBIN OC1493). F. o’, hemi-

penis, detail of part of inner anatomy (KBIN OC1493). G. 9, T1 (SMN 51514). H. co, T1

(KBIN OC1493). I. 2, genital lobe (SMN 51514). J. 9, furca, detail of apical part (SMN 51514).

K. 9, furca and furcal attachment (SMN 51514). Scale: 323 wm for K; 156 wm for E, G, H; 81 wm for

A, ©. Dy EL: 33) amitor B:

REVISION OF AFRICAN CYPRIDINI 111

Remarks

Specimens from the Hwange Nature Park (Figs 10G—M, 11A-—H) are somewhat

smaller than those from Namibia and have a significantly larger H/L ratio (see meas-

urements below). The right prehensile palp has a terminal segment that is somewhat

narrower and more elongated than in the Namibian specimens, whereas the left palp

has a more elongated penultimate segment. These differences, however, seem to fall

in the range of a reasonable variability and no taxonomic value is as yet attached to

them.

Measurements (in wm, mean + S.D., n = 5)

Namibia (SMN 51250). &: L = 1 576 + 38; H = 1 007 + 15; W = 910 + 33; W/L

= 0,55-0,63; H/L = 0,63-0,64. 9: L = 1 645 + 63; H = 1 045 + 38; W = 957 + 43;

W/L = 0,56-0,60; H/L = 0,63-0,65.

Zimbabwe (AM/SED.36). CG: L = 1 452 + 39; H = 993 + 33; W = 907 + 20;

W/L = 0,62—0,64; H/L = 0,67-0,70. 9: L = 1 489 + 26; H = 1 034 + 21; W = 955 +

9; W/L = 0,64—-0,65; H/L = 0,68-0,70.

Discussion

Ramotha curtisae sp. nov. has a somewhat isolated position in the genus and

differs markedly from all congeners except R. trigona by its sub-triangular shape in

lateral view. It differs from the latter species by the convex anterior edge in dorsal

view (beak-like in R. trigona). There can, however, be no doubt that these two taxa

(R. trigona and R. curtisae) belong to the same phyletic lineage within the genus,

R. trigona thus being the most southern representative of this species-group.

Ramotha kenyensis (Lindroth, 1957)

Figs 81, 12A—-F, 14A—D

Eucypris kenyensis Lindroth, 1957: 53-56, figs 1-6. Loffler, 1968: 153-156.

Type locality

Shallow lake in Teleki Valley on Mount Kenya (altitude 4 230 m).

Type material

UPPS (type 601n): 31 Ad 9 (3 used for SEM, 1 dissected and 2 stored in toto—

UPPS.601(n.1-3), 1 (A-1) 9, 16 (A-2) 9, 5 (A-3) @ in spirit in a tube. Lectotype

(here designated): 9, selected from the above syntypes, with soft parts dissected in

glycerine on a sealed slide and with valves stored dry (UPPS.601(n.1—2)); all other

syntypes become paralectotypes.

Diagnosis

Valves elongated (Figs 12A—B, 14A-—B), with posterior margin produced in

lateral view, anterior margin very broadly rounded, smoothly passing into the dorsal

margin, almost without an ascending part and dorsal margin, thus not showing an

anterior blunt angle. In dorsal view (Figs 12C, 14C), anterior edge pointed, greatest

tii? ANNALS OF THE SOUTH AFRICAN MUSEUM

REVISION OF AFRICAN CYPRIDINI 113

width (c. one-half of length) situated near the middle. Soft parts without special

features.

Measurements

Lindroth (1957) (in mm): L = 1,96-2,15 (n = 4); H = 1,10-1,18 (n = 4);

W = 1,07 (n = 1). New measurements (in wm, mean + S.D., n = 5): L = 2 038 + 59;

H = 1 162 + 46; W = 1 183 + 38; W/L = 0,57-0,60.

Discussion

Ramotha kenyensis differs from its geographically closest congeners, R. montana

and R. crassa, in the shape of the anterior margins in lateral view, which are far more

broadly rounded than in the two other taxa. In this, R. kenyensis agrees well with the

South African R. purcelli, to which it is indeed closely related. Ramotha purcelli,

however, has its greatest width in dorsal view situated at about one-third from the

front (in the middle in R. kenyensis).

Ramotha montana (Lindroth, 1957)

Figs 8H, 12G-M, 14E-G

Eucypris hirta Klie, 1944: 17-18 (non Sars, 1924).

Eucypris montana Lindroth, 1957: 56-59, figs 7-13. Loffler, 1968: 153-156.

Type locality

Shallow temporary water on Mount Meru (altitude 2 250 m).

Type material

UPPS (type 228g) 92B: 7 Ad 9 (two used for SEM of which one dissected—

UPPS.228(g.1-3)), 15 (A-1) 9, 4 (A-2) 9, 2 (A-4) Q, and 4 (A-5) 9 in spirit. Lec-

totype (here designated): 2, selected from the above syntypes, with soft parts

dissected in glycerine on a sealed slide and with valves stored dry (UPPS.228(g.1-—2));

all other syntypes become paralectotypes.

Other material investigated

ZIZM 837: c. 10 9 in toto in glycerine (with Cps completely decalcified, but with

Shape recognizable) in a tube, labelled ‘Eucypris hirta G. O. Sars, Belg. Congo:

Albert Nat. Park. no. 51, 11 Marz 1935, coll. H. Damas’.

Fig. 10 (see opposite). A-M. Ramotha curtisae gen. et sp. nov. A-F. From Namibia. A. Oo’, LV,

internal view (KBIN OC1493). B. co’, RV, internal view (KBIN OC1493). C. 9, LV, internal view

(SMN 51514). D. 2, RV, internal view (SMN 51514). E. 9, Cp, ventral view (SMN 51516). F. 9,

Cp, dorsal view (SMN 51516). G-M. From Zimbabwe. G. Oo’, LV, internal view (KBIN OC1508).

H. CO, RV, internal view (KBIN OC1508). I. 0’, Cp, ventral view (AM/SED.36A). J. 9, LV, inter-

nal view (KBIN OC1509). K. 9, RV, internal view (KBIN OC1509). L. 2, Cp, ventral view

(AM/SED.36B). M. o’, Cp, dorsal view (AM/SED.36A). N-Q. Ramotha trichota (G. W. Miller).

N. 9, LV, internal view (KBIN OC1505). O. 9, RV, internal view (KBIN OC1505). P. 0’, LV,

internal view (KBIN OC1506). Q. Cc’, RV, internal view (KBIN OC1506). Scale: 1 333 wm for A-M;

1 587 wm for N-Q.

114 ANNALS OF THE SOUTH AFRICAN MUSEUM

{

Fig. 11. A-H. Ramotha curtisae gen. et sp. nov., Zimbabwe. A. CO’, hemipenis (AM/SED.36C-D).

B. CO, right prehensile palp (AM/SED.36C-D). C. CO’, left prehensile palp (KBIN OC1508). D. 9,

flattened seminal receptacle (KBIN OC1509). E. o’, furca (AM/SED.36C-D). F. ©’, furcal attach-

ment (AM/SED.36C-D). G. ©’, hemipenis, detail of inner anatomy (AM/SED.36C-D). H. Oo, T1

(AM/SED.36C-D). I-L. Ramotha trichota (G. W. Miller), & (KBIN OC1506). I. Furca. J. Right

prehensile palp. K. Left prehensile palp. L. Outline of hemipenis. Scale: 323 wm for L; 156 wm for A,

D, H-K; 81 um for B, C, E-G.

REVISION OF AFRICAN CYPRIDINI 115

Fig. 12. A—-F. Ramotha kenyensis (Lindroth), all redrawn after Lindroth (1957). A. RV, external

view. B. LV, external view. C. Cp, dorsal view. D. Furca. E. RV, internal view, detail of anterior

margin. F. LV, internal view, detail of anterior margin. G-M. Ramotha montana (Lindroth), all

redrawn after Lindroth (1957). G. RV, external view. H. LV, external view. I. Cp, dorsal view.

J. RV, internal view, detail of anterior margin. K. LV, internal view, detail of anterior margin.

L. Mx2, detail of palp. M. Furca. Scale: c. 800 wm for A—C, G-I; c. 300 wm for D-F, J—M.

116 ANNALS OF THE SOUTH AFRICAN MUSEUM

REVISION OF AFRICAN CYPRIDINI 117

Diagnosis

Valves (Figs 12G—H, 14E-G) higher than in the preceding species, with anterior

margin in lateral view less broadly rounded and more pointed, passing into the dorsal

margin after a long ascending part with a blunt angle; LV posteriorly evenly rounded.

Cp in dorsal view (Figs 121, 14L) wider than in the preceding species, but with great-

est width also situated in the middle; anterior edge pointed, but less so than in

R. producta, and posterior edge convexly rounded. Soft parts without special features.

Measurements

Lindroth (1957) Gn mm, n = 2): L = 1,94-1,98; H = 1,14-1,16; W = 1,13-1,17.

New measurements (in wm, mean + S.D.,n = 5): L= 1914 + 57; H = 1 141 + 45;

W = 1 148 + 55; W/L = 0,58-0,62.

Discussion

See under the following species.

Ramotha producta (Sars, 1924) comb. nov.

Figs 1A—B, 7K-—M, 13, 15SE-K, 16A-G

Eucypris producta Sars, 1924: 109-110, pl. 3 (figs 1-2).

Type locality

Vlei near Port Elizabeth (South Africa).

Type material

SAM-A11312: 5 9 from the above locality (labelled ‘Eucypris propinqua G. O.

Sars’—MS-name) in spirit (2 9 used for SEM, nos SAM-—A40040-1171, the former

dissected). Lectotype (here designated): the dissected 9 with soft parts dissected in

glycerine on a sealed slide and with valves stored dry (SAM-—A40040); the other syn-

types become paralectotypes.

Other material investigated

A slightly aberrant form of this species was collected from various temporary

water bodies in the vicinity of Grahamstown, Eastern Cape Province (KBIN/

IG.27549-GR/12-15); in some of these localities, bisexual populations occurred

(KBIN/IG.27549-GR/17-19). The specimens from the parthenogenetic populations

are slightly more elongated and have carapaces that are narrower and show a

Fig. 13 (see opposite). Ramotha producta (Sars). A-B. Lectotype, 2 (SAM-—A40040). A. LV, inter-

nal view. B. RV, internal view. C. Paralectotype (SAM-—A40041), 9, Cp, ventral view. D-F. Parthe-

nogenetic Y from Grahamstown. D. LV, internal view (KBIN OC1502). E. RV, internal view

(KBIN OC1502). F. Cp, ventral view (KM.1173—lost). G—-L. Males and females from a bisexual

population, Grahamstown. G. 9, LV, internal view (KBIN OC1497). H. 9, RV, internal view

(KBIN OC1497). I. co’, LV. internal view (KBIN OC1494). J.c’, RV, internal view

(KBIN OC1494). K. co’, Cp, dorsal view (KBIN OC1496). L. o’, Cp, ventral view (KBIN OC1495).

M-P. 7th larval instar 9 from a parthenogenetic population near Grahamstown. M-N. LV

(KBIN OC1501). M. Internal view. N. Anterior detail. O-P. RV (KBIN OC1501). O. Internal view.

P. Anterior detail. Scale: 1 587 wm for A-L; 781 wm for M, O; 145 wm for P; 133 wm for N.

ANNALS OF THE SOUTH AFRICAN MUSEUM

118

REVISION OF AFRICAN CYPRIDINI 119

cavely pointed posterior end. The & and some of the 9 from the bisexual populations

generally have a significantly larger height/length ratio (see measurements). Other

females from the same populations have the same shape as the parthenogenetic speci-

mens. Both types of 2 were found to contain spermatozoids, although less commonly

in the more elongated specimens. Intermediate forms, although rare, were present,

which is why no taxonomic importance is attached to these morphological differences.

All this material was collected by K. Martens (with H. Barber and F. C. de

Moor) in November 1989.

Diagnosis

Q (parthenogenetic populations). Valves (Fig. 13A—B, D-E) elongated and cau-

dally produced, anterior margin in lateral view not broadly rounded, and passing into

the dorsal margin after a relatively long ascending part, with a blunt angle; the latter

angle situated considerably more towards the front than in the preceding species. LV

posteriorly not evenly rounded, showing a weak and blunt angle. Cp (Fig. 13C, F)

wide in dorsal view, but with greatest width situated slightly anterior to the middle;

anterior edge pointed, more so than in the preceding species; lateral sides mostly

running straight and parallel to each other over about one-third of the total length.

Q (bisexual populations). As in the former, but with valves considerably higher

(Fig. 13G, H); in dorsal view, most carapaces with evenly rounded lateral sides.

those of the parthenogenetic 9. Cp in dorsal view (Fig. 13K, L) with rounded lateral

sides. Hemipenis with lateral shield broadly rounded, medial shield asymmetrically

produced towards the ventral side, bluntly pointed (Fig. 16A, D). Left prehensile palp

(Fig. 16B, E) with distal segment tapering, evenly rounded, proximal segment with

two medium-sized subapical sensory organs. Right prehensile palps (Fig. 16C, F) with

distal segment sub-triangular, with three nearly straight margins; proximal segment

with 2 larger sensory organs. A2 with normal sexual dimorphism in the apical chaeto-

taxy. T1 with distal claw somewhat longer than in the 9 (Fig. 14E). Furca with ramus

straight. Other soft parts as in the Q.

Measurements

See Table 1.

Discussion

Ramotha montana and R. producta are closely related and could indeed constitute

geographically isolated subspecies of the same species. The differences cited in the

above diagnosis of R. producta, however, seem sufficient to maintain the specific

Fig. 14 (see opposite). A-D. Ramotha kenyensis (Lindroth). A-B. Lectotype, @ (UPPS.601(n.1-2)).

A. LV, internal view. B. RV, internal view. C—D. Paralectotype, 2 (UPPS.601(n.3)). C. Cp, dorsal

view. D. Cp, ventral view. E-G. Ramotha montana (Lindroth). E. Lectotype, 9, LV, internal view

(UPPS.228(g.1-2)). F. Paralectotype, 2, Cp, ventral view (UPPS.228(g.3)). G. Lectotype, 2, RV,

internal view (UPPS.228(g.1—2)). H—L. Ramotha hirta (Sars). H. Paralectotype, 2, LV, external view

(SAM-A40044). I-J. Lectotype, 9 (SAM-A40043). I. RV, internal view. J. LV, internal view.

K. Cp, ventral view (KM.1229). L. Cp, dorsal view (KM.1228). Scale: 1 389 4m for A~G; 1 111 wm

for H-L.

120 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 1

Measurements of different populations of Ramotha producta (in pm, mean + S.D.).

Population L H Ww H/L W/L

GR/12 (n = 5)

parthenogenetic 9° 2161+38 1162+38 1193+41 0,52—0,55 0,54-0,57

GR/17 (n = 5)

parthenogenetic? 9° 2321+46 1241+44 1255+31 0,52—0,55 0,53-—0,57

bisexual 9 PN SS2337/ 1231+34 1179+23 0,56—0,59 0,54—0,57

oy 1954+54 1161+27 1092+36 0,58—0,61 0,54—0,59

Lectotype RV 2241 1276 — 0,57 —

Lectotype LV 2276 1310 _— 0,58 —

status of both taxa. Future finds of intermediate populations will reveal whether this

decision can be corroborated.

Ontogeny

As larval morphology can be most illuminating when attempting to reveal phylo-

genetic relationships, aspects of the morphology of the larval stages that were

available are presented.

Sth larval stage of 2. RV (Fig. 7M) with approximately the same shape as in the

adult, but with anatomy of the valve margin substantially different: both anteriorly

and posteriorly with selvage inwardly displaced over a short distance only and with a

long and conspicuous inner list (absent in adults).

LV (Fig. 7K) on both sides with an inner list only (as in the adults), but situated

closer to the valve margin than in the adult stage. Shape of Cp in dorsal view

(Fig. 7C) as in the adult.

Al with 2+5 segments, all segments relatively shorter and wider than in the

adult; Rome organ relatively larger; distal segment with Ya long and narrow and with

one of the setae shorter, approximately two-thirds of the length of Ya.

A2 with five long natatory setae; z: with base about twice as wide as in Z2 and Zs,

i.e. rather claw-like; G2 short and incompletely developed; terminal segment with ys

bifurcated; penultimate segment with three lateral ‘t’-setae.

Md palp with seta alpha narrower, beta subequal to the one in the adult and

gamma-seta flanked by two instead of three setae.

Mx1 palp with terminal segment c. 2,5 times as long as basal width, third endite

with two smooth claws.

Mx2 with two subequal ‘a’-setae, one ‘b’- and one ‘d’-seta; respiratory plate carry-

ing four long and one short ray; palp with three apical setae.

T1 (Fig. 15F) with penultimate segment divided; claw relatively short, but with

two rows of teeth; seta d2 = c. 1,5 times the length of seta dh.

Basal segment of T2 with three setae; apically with fourth segment somewhat

more separated from third segment than in the adult, but still forming a cleaning

pincer.

REVISION OF AFRICAN CYPRIDINI 121

'n,

= \ SS Yy

\ Pi ye

eas hs

~& \ 7 . Ny fda

\ ( SS

| pies Zn

4 S S ae

{ SS SS

NN ws ae ’

\\ <SSs ee ai

\ — Ee ae {

J f = See

RAN

AHL, ay

ANYRTA A Re

Fig. 15. A~D. Ramotha purcelli (Sars), Q?(SAM-—A40046). A. Mx2, detail of palp. B. Mx2, detail of

‘a-setae. C.T1. D. Furcal attachment. E-K. Ramotha producta (Sars). E.Q (Ad), T1

(KBIN OC1502). F. 2 (8th larval instar), T1 (KBIN OC1499). G. Q (7th larval instar), T1

(KBIN OC1501). H. 9 (7th larval instar), furcal attachment (KBIN OC1501). I. Q (7th larval instar),

furca (KBIN OC1501). J. 9 (7th larval instar), other furca of same specimen (KBIN OC1501). K. 2

(Ad), furca (KBIN OC1502). Scale: 323 wm for D, K; 156 wm for A, C, E-J; 33 wm for B.

122 ANNALS OF THE SOUTH AFRICAN MUSEUM

it HAA Ao

Fig. 16. A~G. Ramotha producta (Sars), CO’. A. Hemipenis (AM/LEN.17A). B. Left prehensile palp

(AM/LEN.17A). C. Right prehensile palp, insufficiently flattened (AM/LEN.17A). D. Hemipenis

outline (KBIN OC1494). E. Left prehensile palp (KBIN OC1494). F. Right prehensile palp

(KBIN OC1494). G. Zenker’s organ (KBIN OC1494). H. Ramotha hirta (Sars), 29 (SAM—A40043),

Mx2 palp. Scale: 156 um for A, D, G, H; 81 um for B, C, E, F.

REVISION OF AFRICAN CYPRIDINI 123

Furca with rami shorter and wider, (sub-)apically still with two claws and setae.

Furcal attachment bifurcated. Rake-like organs as in the adult.

7th larval stage of °. Shape of both valves more elongated, with greatest height

situated more towards the front; ventral sinuous indentation more caudally situated

and caudal margin more pointed than in the previous larval instar (Fig. 13M, O).

Marginal anatomy similar to the 8th larval stage, but with selvage in RV and inner list

in LV situated even closer to the valve margins (Fig. 13N, P).

Al with Rome organ still visible.

A2 with four long natatory setae; apical segment with claws GM and Gm, seta ‘g’

and aesthetasc y3 penultimate segment with two subapical z-setae, claw G2 short and

only two lateral ‘t’-setae.

Md palp normal alpha-seta, beta-setae somewhat wider and gamma-setae more

smooth.

Mx1 with second palp segment approximately three times as long as the basal

width, two claws on third endite smooth.

Mx2 with only one ‘a’-seta and one ‘b’- and ‘d’-seta; respiratory plate with three

long and one short ray; palp apically with one long, one medium-sized and one minute

seta.

T1 with setae di and d2 subequal, penultimate segment divided and apical claw set

with two rows of teeth (Fig. 15G).

T2 as in the former larval stage, but with all segments relatively wider and

shorter.

Furca with ramus short (sometimes both rami significantly different—see

Fig. 15], J). Furcal attachment (Fig. 15H) bifurcated.

Ramotha purcelli (Sars, 1924)

Figs IM-N, SI-O, 15A-—D

Eucypris purcelli Sars, 1924: 108-109, pl. 2 (figs 12-15).

Type locality

Pond at Ashton, Robertson Division (South Africa).

Type material

SAM-—A11287: syntypes, c. 100 good 9 and larvae, collected from the above

locality (SAM—A40047—A40049 used for illustrations and stored separately).

Lectotype (here designated): 9, selected from the above syntypes, with soft parts

dissected in glycerine on a sealed slide and with valves stored dry (SAM-—A40046); all

other syntypes become paralectotypes.

Diagnosis

Both valves elongate in lateral view (Fig. 51, J), with frontal margin broadly

rounded, passing into the dorsal margin almost without an angle, the latter margin

passing into the caudal margin with a blunt angle; caudal margin not evenly rounded

but showing an additional blunt angle; ventral margin weakly sinuous. Greatest height

situated slightly in front of the middle. Ventral outer list on LV weak (Fig. 5L); dorsal

Outer list on this valve prominent (Fig. 5M). Cp in dorsal view with RV anteriorly

124 ANNALS OF THE SOUTH AFRICAN MUSEUM

clearly reaching beyond LV, and with greatest width situated at about one-third from

the front (Fig. 5K). Soft parts without special features.

Measurements (in wm; mean + S.D., n = 5)

LE = 2 089 22 1295 172 58 W = IO O = S56) Oso Oa

Discussion

The differences between R. purcelli and R. producta are similar to those between

R. kenyensis and R. montana. Ramotha purcelli is thus similar to R. kenyensis, but is

sufficiently different in its morphology to merit separate specific status (see discussion

under R. kenyensis).

Ramotha trichota (G. W. Miller, 1908)

ates WEI, ONO, 1,

Cypris trichota G. W. Miller, 1908: 152-153, figs 1-5.

Eucypris trichota Sars, 1924: 108, pl. 2 (figs 1-11).

Type locality

Zeekoevlei, Plumstead, near Simonstown (South Africa).

Material investigated

GW no. 25133(7g): 1 9 (‘Siidafrika—Sidpolar Exp.’). (Note: this is probably

one of the specimens used by G. W. Miller for the original description of the species.

However, this author mentioned that he had 9 9 from Zeekoevlei. As we do not

know the whereabouts of the other specimens, a lectotype will not be designated here.

The specimen was left undissected. )

SAM-A11926: c. 20 2 (with soft parts in bad condition) from Valkenbergvlei,

collected on 15 July 1922 by E. M. Stephensen (?det. D. H. Eccles).

SAM-A11286: 5 @ and numerous valves from ‘SAM EEF’ (?) (identified as

Eucypris producta by G. O. Sars, as R. trichota by K. Martens).

KBIN OC1507 (GS01/18): 1 9 from Wiesdrif, Western Cape Province (approx.

co-ordinates: 34°40'12”S 19°54'47"E), collected by Dr J. King (UCT, Cape Town) on

11 May 1989 (det. K. Martens).

KBIN OC1505-1506 (G501/19): 19 and 1 CO from Soetendalsvlei Ditch,

Western Cape Province (34°44'42"S 19°58'33"E), collected by Dr J. King (UCT, Cape

Town) on 11 May 1989 (det. K. Martens).

Diagnosis (Q)

By far the largest of all Ramotha species, with high valves (Fig. 10N, O), greatest

height situated at about one-third from the front, with dorsal margin sloping towards

the caudal side and passing into the caudal margin without an angle; anterior margin

in lateral view more broadly rounded than posterior one. Cp in dorsal view with

anterior edge beak-like (concavely produced) and posterior edge rounded; greatest

width situated slightly anterior to the middle. Soft parts without special features.

REVISION OF AFRICAN CYPRIDINI WD)

First description of C

Valves (Fig. 10P, Q) shorter and considerably higher than in the 9, in lateral

view with a sub-triangular appearance; anatomy of the valve margin identical to that

of 2. Both prehensile palps with surprisingly small penultimate segment, in right palp

only slightly longer than the terminal segment (Fig. 11J). The latter in the right palp

furthermore with a wide base and an irregular outline. Terminal segment of left palp

(Fig. 11K) sickle-shaped and rather more elongated than in the other species of

Ramotha. Hemipenis (Fig. 11L) with an elongated lateral shield, with straight and

parallel lateral margins and a nearly straight distal margin.

Remark

The inner part of the hemipenis was not yet fully developed and sclerotized in the

only C&’ specimen available for examination.

Measurements (in wm)

Type specimen (GW.25133/7g): L = 2 862; H = 1 724; W = 1 551; W/L = 0,54.

Soetendalsvlei Ditch, 9 (KBIN OC1505): RV: L = 2 724; H = 1 758. LV: L = 2 655;

H = 1758. GC (KBIN OC1506): RV: L = 2224; H = 1483. LV: L = 2 155;

H = 1 534.

Relationships

This species can at once be distinguished from all congeners by the superior size

of the 2: almost 1 mm longer than the second largest species, R. producta.

Remarks

The MRAC (Tervuren) holds two slides (RG.38570) identified as Eucypris tri-

chota by R. Kiss, the specimen originating from Bogarama in East Africa (coll.

Marlier 1949). This specimen, however, is in such a bad condition that it became com-

pletely unidentifiable and this record is therefore disregarded here.

Ramotha trigona (Sars, 1895)

Fig. 1G—H

Cypris trigona Sars, 1895: 32-34, pl. 5 (fig. 3a—c).

Eucypris trigona Sars, 1924: 111-112, pl. 3 (figs 7-8).

Type locality

Swamp near Knysna, east of the Cape of Good Hope (South Africa).

Type material

OSLO F4339: 7 decalcified Q and several empty, decalcified valves. As permis-

sion was not obtained to dissect specimens, a lectotype has not been designated here.

Diagnosis

Valves in lateral view sub-triangular (Fig. 1G—H), with greatest height situated

towards the front; both anterior and posterior margins bluntly pointed in lateral view,

but the latter more so; ventral margin nearly straight. Cp in dorsal view with greatest

126 ANNALS OF THE SOUTH AFRICAN MUSEUM

width situated at or slightly posterior to the middle; anterior edge with concave beak-

like extension. Both valves anteriorly with a wide fused zone, posteriorly with fused

zone and flange wide.

Morphology of soft parts unknown.

Measurements

According to Sars (1924), the 9 of this species is 1,75 mm long. New measure-

ments (in wm, mean + S.D., n = 3): L= 1 649 + 105; H = 948 + 69; W = 862 + 17;

W/L = 0,50-0,53.

KEY TO THE SPECIES

LAS Kengthiot 2 more than)2: 5 mmi(We Cape)renee eee ee eee R. trichota

1B. Length of 9 less than 2,5 mm... 222.20 42. odo 5. sees eee Z

2A. Ventral margin (including selvage in RV and inner list in LV) conspicuously

produced in the first third (Fig. SC, D), especially in the 2; carapace very hairy

EW.s Cape yi esi cc hog Be tice ah Ue re OWN we Aelita at R. hirta

2B. Carapace less hairy, ventral margin not conspicuously produced in the first third

ee er rt a in iam Milano sco oc ac aseco 25 3

3A. W/L ratio >2/3, in dorsal view anterior conspicuously beak-like ............ 4

3B, W/L ratio <1/2; in‘dorsal view anterior less) produced) =>. 925-2 eee 5

4A. Valves in lateral view with a nearly evenly rounded dorsal margin, a nearly

Straight ventral margin and with anterior and posterior margins nearly evenly

rounded (W.'Cape) a cack ce Ne ba we OR cee on eer ea R. corpulenta

4B. Valves in lateral view more elongated, with curved ventral margin and with

anterior margin more broadly rounded than posterior one (W. Cape).........

ee tees ete ep ONAN MME Mea araluusld Bro W o'd Sc 0.0 0 0 R. capensis

SA: Valves sub-triangulat ici. 5 5226 Gaede ss bation os eo soe ee ee 6

3B. Valves elongated; not sub-triangulan = 7.242522. 22555525552) Gee 7

6A. Carapace in dorsal view with concavely produced beak (W. Cape) ...R. trigona

6B. Carapace in dorsal view anteriorly and posteriorly convexly rounded (Namibia

and Zimbabwe) 0.055 bi le wae A oe eae nie R. curtisae

TAL” Tee 1475 mm @East Anica) y 2 75. ne) R. crassa

7B. L=c, 2mm (East‘and South Afmica) ...07....../. 5. 527. oe 8

8A. Anterior margins of isolated valves broadly rounded, passing into the dorsal

margin almost withoutam angle 7.202.252.5432 5.456 56> 454d Oe oe 9

8B. Anterior margin of isolated valves less broadly rounded, more pointed, and

with a conspicuous angle between anterior and dorsal margin ............. 10

9A. Carapace in dorsal view with greatest width situated almost at mid-length (East

PNETACAL) So seseosslhy ie cs ailese: Se as Lar ett eI eae cee ea eee ee R. kenyensis

9B. Carapace in dorsal view with greatest width situated at about one-third from the

front:(South Africa) mest sir be oe ieoud Gobel ae eal eta ae eae R. purcelli

10A. Carapace in dorsal view with posterior margin convexly rounded (East Africa)

ee ee ee a atid lua ato oe A-oxa ba BO 6 0 R. montana

REVISION OF AFRICAN CYPRIDINI 127

10B. Carapace in dorsal view with posterior margin concavely pointed (South Africa)

5 00 019-09 SIO G, DRO LatO chs OU Doone Ect Dfo G Jn Rn Ae ie Cn nner eer ea a R. producta

Note. To distinguish between the latter four species, check all figures carefully, especially when

dealing with material from localities between East and southern African regions.

DISCUSSION

TAXONOMY AND ZOOGEOGRAPHY

Ramotha gen. nov. belongs to the tribe Cypridini in the Cypridinae, because of

the presence of the ‘8’-shaped loops of the inner spermiductus in the hemipenis. Its

affinities with the Eucypridini are thus limited. For a further discussion on the validity

of these higher taxa and their mutual affinities, see the first part of this revision

(Martens 1990). Within Ramotha gen. nov., we can distinguish different phyletic

lineages:

Ramotha trichota is quite distinct within the genus and constitutes a separate lineage.

Ramotha capensis, R. corpulenta and R. hirta constitute another lineage, characterized

by a wide carapace with a pointed, beak-like frontal edge. This group forms part

of the Cape fauna.

Ramotha trigona and R. curtisae sp. nov. form another group within the genus; both

taxa have sub-triangular valves. They occur in most of southern Africa.

The final lineage is formed by the remaining five species: the small R. crassa, the

larger R. kenyensis and R. montana (all from East Africa), and the South African

species R. purcelli and R. producta. We expect this group to have representatives

also in the intermediate territories, e.g. in Zambia, Mozambique, etc.

Nevertheless, the genus forms one closely related group, quite distinct from the

other Cypridini, as is shown by the very uniform shape of the hemipenis. The genus is

furthermore well characterized by a T1 with a divided penultimate segment (fused in

Cypris and Pseudocypris) and by the anatomy of the valve margins. The latter feature

deserves some further elaboration. The closing mechanism of the anterior part of the

valves in Ramotha is formed by a large, inwardly displaced selvage in the RV, which

locks into a conspicuous inner list in the LV. In Cypris, the closing mechanism is

formed by two inwardly displaced selvages, one on each valve. Such a system also

exists in, for example, Chlamydotheca and in nearly all Megalocypridinae. On the

other hand, a similar system to that in Ramotha was described for Trajancypris

(Martens, 1989), a genus belonging to the Eucypridini. In spite of this, Cypris and

Ramotha are much more closely related to each other than to the groups cited above.

It is thus concluded that supra-generic affinities should not be based on such func-

tionally relevant aspects of valve anatomy but rather on non-functional relicts of the

morphology, e.g. the presence or absence of a ‘c’-seta on the Mx2.

Another, superficially similar genus is the Australian Alboa De Deckker, 1981.

Thanks to the kindness of Dr De Deckker, I was able to re-investigate a number of

paratypes. It appeared that Alboa does not belong in the Cypridini, as it lacks ‘8’-

Shaped loops of the inner spermiductus in the hemipenis, apart from other features

that are different. Its original allocation to the Cyprinotinae still seems the most plaus-

ible to date.

128 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 17. Distribution of African species of Ramotha gen. nov.

REVISION OF AFRICAN CYPRIDINI 129

The distribution of Ramotha is similar to that found in the genus Sclerocypris (see

Martens 1986, 1988). There are, however, a number of differences. Firstly, Sclero-

cypris also has Indian representatives. Only African species are here assigned to

Ramotha gen. nov., but it is unlikely that this genus has an exclusively African distri-

bution and representatives of this genus can be expected in India. Secondly, there is at

least one species of Sclerocypris, S. bicornis (G. W. Miiller), which is widespread in

most of Africa, and also in the northern part of the continent. Contrary to this, all

Ramotha species have restricted geographical distributions (Fig. 17).

ECOLOGY

Thus far, species of Ramotha were mostly found in temporary habitats. They are

reported from a number of small to medium-sized, permanent water bodies (Lake

Elgol in East Africa and Zeekoevlei in South Africa), but never from the larger

ancient East African lakes.

No fewer than three species of Ramotha occurred sympatrically in Zeekoevlei at

the beginning of this century. Such coexistence of congeners, although not unique in

ostracods, is of interest. Recent sampling in Zeekoevlei (December 1989) revealed

that eutrophication has destroyed most of the original fauna of this vlei and not one

representative of Ramotha was found.

All Ramotha species were thus far collected from fresh or slightly saline waters

only. Most species are furthermore exclusively known from parthenogenetic popu-

lations. A fairly thorough survey of temporary waters in the vicinity of Grahamstown

(Eastern Cape), revealed that R. producta occurs in both parthenogenetic and bisex-

ual populations in virtually adjacent vleis. Nothing is known about the life history of

these species, other than that their complete life cycle takes three weeks or less (at

temperatures between 10° and 30°C); this is the time between the first rains and the

collection of adults.

ACKNOWLEDGEMENTS

Mrs C. Behen and Mr J. Cillis offered technical assistance with the illustrations.

Mr M. Cartois patiently sorted the collections from Namibia and the Cape Province.

Material from various museums and collections was obtained through the efforts of

the following people: Miss B. Curtis (Windhoek), Dr P. De Deckker (Canberra),

Dr F. de Moor and Miss H. Barber (Grahamstown), Miss L. Hoenson and Mrs M.

van der Merwe (Cape Town), Dr R. Jocqué (Tervuren), Mr N. Langeland (Oslo), Dr

G. Miller (Greifswald), Mr H. Petersen (Hamburg) and Dr L. Wallin (Uppsala).

Dr D. Danielopol (Mondsee) and an anonymous referee read the manuscript and sug-

gested improvements. I would furthermore like to take this opportunity to thank the

people who, by their warm hospitality, their active co-operation and enthusiastic

support made my trips to Namibia and South Africa such rewarding experiences. They

are: Miss B. Curtis (Windhoek), Dr J. A. Day, Dr B. Davies, Miss E. Louw, Miss

K. Jagoe, Mrs M. van der Merwe and Miss L. Hoenson (Cape Town), Dr F. C. de

Moor, Miss H. Barber and Miss N. Kohly (Grahamstown). To all of the above and to

the many I might have forgotten, my sincere gratitude!

130 ANNALS OF THE SOUTH AFRICAN MUSEUM

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der Wetenschappen 48 (174): 81 pp + 64 figs.

Martens, K. 1987. Homology and functional morphology of the sexual dimorphism in the antenna of

Sclerocypris Sars, 1924 (Crustacea, Ostracoda, Megalocypridinae). Bijdragen tot de Dierkunde

57 (2): 183-190.

Martens, K. 1988. Seven new species and two new subspecies of Sclerocypris Sars, 1924 from Africa,

with new records of some other Megalocypridinae (Crustacea, Ostracoda). Hydrobiologia 162

(3): 243-273.

Martens, K. 1989. On the systematic position of the Eucypris clavata-group, with a description of

Trajancypris gen. nov. (Crustacea, Ostracoda). Archiv fiir Hydrobiologie (Supplement) 83 (2):

227-251.

Martens, K. 1990. Taxonomic revision of African Cypridini. Part I. The genera Cypris O. F. Miller,

Pseudocypris Daday and Globocypris Klie (Crustacea, Ostracoda). Bulletin van het Koninklijk

Belgisch Instituut voor Natuurwetenschappen, Biologie 60: 127-172.

MULLER, G. W. 1908. Die Ostracoden der Deutschen Siidpolar-Expedition 1901-1903. In: Drygalski,

E. V. ed. Deutsche Stidpolar-Expedition 10 (zool. 2): 51-182.

MULLER, G. W. 1914. Sitisswasserostracoden der Deutschen Siidpolar-Expedition 1901-1903. In:

Drygalski, E. V. ed. Deutsche Stidpolar-Expedition 16: 67—78 (2nd edition 1921, Berlin).

Sars, G. O. 1895. On some South-African Entomostraca raised from dried mud. Skrifter Videnskabs-

Selskabets. I. Mathematisk-Naturvidenskab. Klasse 8: 3-56.

Sars, G. O. 1924. The freshwater Entomostraca of the Cape Province (Union of South Africa).

Part 2. Ostracoda. Annals of the South African Museum 20 (2): 105-193.

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

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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

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Synonymy arrangement according to chronology of bibliographic references, whereby the year is

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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

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(a) The Figures, Maps and Tables of the paper when referred to in the text

Cee ethieuiouncdepictine ©, mamacoius...: ...im ©. namacolus (Fig: 10)...

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initials or full names

e.g. Du Toit 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

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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.

K. MARTENS

TAXONOMIC REVISION OF

AFRICAN CYPRIDINI.

PART II. DESCRIPTION OF

RAMOTHA GEN. NOV.

(CRUSTACEA, OSTRACODA)

VOLUME 102 PART 3 OCTOBER 1992 ISSN 0303-2515

E SOUTH AFRICAN.

OF TBE

CAPE TOWN

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

FiscHer, P. H. 1948. Données sur la résistance et de la vitalité des mollusques. Journal de conchyliologie 88 (3): 100-140.

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

expérimentale et générale 74 (33): 627-634. ,

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

Magazine of Natural History (13) 2 (17): 309-320.

Koun, A. J. 1960b. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bulletin of

the Bingham Oceanographic Collection, Yale University 17 (4): 1-51.

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

pologische Ergebnisse einer Forschungsreise im westlichen und zentralen Stid-Afrika ausgeftihrt in den Jahren

1903-1905 4 (15). Denkschriften der medizinisch-naturwissenschaftlichen Gesellschaft zu Jena 16: 269-270.

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 102 °+#£2Band

October 1992 Oktober

Part 3 Deel

ADDITIONS AND REVISIONS TO THE

LIST OF SPECIMENS OF THE

EXTINCT BLUE ANTELOPE

(HIPPOTRAGUS LEUCOPHAEUS)

L. C. ROOKMAAKER

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

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D1672

ADDITIONS AND REVISIONS TO THE LIST OF SPECIMENS OF THE

EXTINCT BLUE ANTELOPE

(HIPPOTRAGUS LEUCOPHAEUS)

L. C. ROOKMAAKER

Doormanstraat 31, West 1, 7731 BN, Ommen, The Netherlands

(With 6 figures)

[MS accepted 29 April 1991]

ABSTRACT

Very little material of the extinct blue antelope (Hippotragus leucophaeus) is known to exist.

Until now, four mounted skins, two pairs of horns and one skull have been documented and their

history is summarized. Three further specimens are now added: a hitherto unknown pair of horns dis-

covered in Cape Town; a skull in the Zoological Museum of Amsterdam; and a now unavailable skin

brought to Holland in 1805 by J. A. Uytenhage de Mist. This specimen was donated to the museum of

the Hollandsche Maatschappij der Wetenschappen in Haarlem; it is supposedly the second blue ante-

lope skin once present in this collection.

CONTENTS

PAGE

MTUGROMUCtIOMUE ee RN rant pert eerie Fah rarve ra ceoine eat ne Ae RBG SSS oe wie eaten 131

Bishotknownispecimensiof£ the blucrantclope.. 4444-44 -0405 400.0 c0 cesses ae. 132

PNAGIIONS EO LHe HSHORSDECIMENS 5.60 eet ce eee Se cee ee dee ek eee es 135

LOLMSHIN Cape MlOWw Mir ee les ne cay neni eed ba Slavoardiow eee ht IBS

PNSKulliny Amsterdam en. ec emicei cuss Hee ie see keh led.oeae sete 136

Evidencclofaskinonce imriaarlem:..... 2. she. cs eee e sees cee eet taege es cn 136

PNCKMOWIEC SEM MUSH Mere tir set eerie ere amet ile uae atone ee sah ie 140

INCLEREMCES He eRe eee Ree ian ie ee eRe eh RNIN Gale wh dale nah waite ie 140

INTRODUCTION

The blue antelope, Hippotragus leucophaeus (Pallas, 1766), became extinct as

early as the year 1800. The only evidence regarding its existence consists of a few

short descriptions written in the seventeenth and eighteenth centuries, four mounted

specimens in museums, as well as some horns and skulls, and three drawings. The

available information was comprehensively treated by Mohr (1967) in her classic

monograph on the species. Some additional material was reported by Klein (1974,

1987).

The blue antelope was a close relative of both the sable, Hippotragus niger

(Harris, 1838) and the roan, Hippotragus equinus (Desmarest, 1804). It has been

Stated that the species was mentioned for the first time in 1719 in the book on the

Cape of Good Hope by Peter Kolb. Earlier, however, its name appeared in a list of

South African animals by Johann Schreyer (1681): ‘Auch lauffen auff den hohen

Bergen von vielerley Arten der wilden Bécke, und Ziegen, als Gems-Bécke, blaue

Bocke, bundte Bécke, Rehe-Bécke, Klippsteiger, Steinbécke, wilde Bocke’ [italics

added]. The blue antelope, with its assumed shoulder height of 1-1,2 m, was some-

131

Ann. S. Afr. Mus. 102 (3), 1992: 131-141, 6 figs.

132 ANNALS OF THE SOUTH AFRICAN MUSEUM

what smaller than the sable and the roan; its teeth too were smaller and the horn cores

more compressed on the sides. It had a faint blue to grey colour, whitish underparts, a

brown forehead and top of the muzzle, and a lighter coloured patch in front of the

eyes. The blue antelope was a grazer that lived in groups of up to 20 animals. In

historic times, it was recorded only from a small region in the south-western Cape

Province, roughly between Caledon and Swellendam. Rookmaaker (1989) gave details

of the three known drawings of the blue antelope made during the eighteenth century,

i.e. one by Robert Jacob Gordon presently in the Rijksmuseum, Amsterdam, and two

almost identical drawings connected with the work of Francois Levaillant preserved in

the Library of Parliament, Cape Town, and in the Library of the University of Leiden

(Fig. 1).

LIST OF KNOWN SPECIMENS OF THE BLUE ANTELOPE

Mohr (1967) gave a reliable and detailed list of the known specimens of the blue

antelope. She enumerated four mounted skins, two pairs of horns, and four skulls.

The history of these specimens will be summarized here, with the addition of a few

historical points discovered since 1967. In this paper, I give some additional remarks

about three specimens: a pair of horns recently discovered in the South African

Museum in Cape Town; a skull found in the Zoological Museum of Amsterdam; and a

missing mounted skin in the (former) Museum of the Dutch Society of Sciences in

Haarlem.

The identification of the skulls has been debated. Mohr’s list (1967: 44) included

two skulls in the Zoological Museum in Berlin, but she concluded that these were

incorrectly referred to Hippotragus leucophaeus. Another skull in her list had been

reported in the museum of the Royal College of Surgeons in London, but it was

destroyed in an air attack in 1941. Although it had been attributed to the blue ante-

lope by Renshaw (1921), Mohr showed that this skull also more probably belonged to

a roan. The identity of the fourth skull is still inconclusive (see no. 7 below).

Mohr (1967: 43) mentioned uncertain evidence concerning a pair of horns in the

Albany Museum in Grahamstown. This refers to a note in that museum’s annual

report for 1901, where director S. Schénland (1902: 3) stated that he “came across a

pair of horns, which I take to be identical with a pair of horns identified in one of our

old catalogues as belonging to the Blaauwbok (Hippotragus leucophaeus).’ It was

suggested that this specimen was destroyed in a fire in the early 1940s, an opinion that

is confirmed by W. H. Holleman, deputy director of the Albany Museum (in litt.

18 March 1991); he informed me that the only pair of horns in that museum which

could fit the description was recently identified as belonging to Hippotragus equinus.

We can thus confine our list to the four mounted specimens, the horns and one

skull.

1. The Zoological Museum in Stockholm has a mounted skin of a young male. Its pro-

venance is not known, except that it was in the private collection of Adolf Ulrich Gmill

(1752-1797) before its accession in the museum in 1829.

2. The Zoological Museum in Vienna preserves the mounted skin of an adult female.

Nothing is known about the history of the specimen. It was first recorded in Vienna by

Kohl (1886: 84).

LIST OF SPECIMENS OF THE EXTINCT BLUE ANTELOPE 133

—

Fig. 1. The blue antelope in a collection made by Francois Levaillant around 1780, preserved in the

library of the University of Leiden (UBL 13 in Rookmaaker 1989).

3. The Rijksmuseum van Natuurlijke Historie in Leiden has the mounted skin of an

adult male, now the lectotype of Hippotragus leucophaeus (Pallas, 1766). The speci-

men was discovered by J. C. Klockner in a shop in Amsterdam before 1776. Kléckner

mounted it and then gave (or sold) it to Jacob Cornelis Sylvius van Lennep

(1746-1776). On Van Lennep’s death, his entire collection, including the blue ante-

lope skin, was bequeathed to the Hollandsche Maatschappij der Wetenschappen

134 ANNALS OF THE SOUTH AFRICAN MUSEUM

(Dutch Society of Sciences) in Haarlem, Holland. It was listed in the Society’s cata-

logue of 1803 (Van Marum 1803: x). In 1842, the Society moved its collection from

one building to another and decided to sell a number of its mammals and exotic birds.

As detailed by Husson & Holthuis (1969: 150-151), these specimens were auctioned

on 15 April 1842 by A. Engesmet in Haarlem, including ‘a very large and clean speci-

men of Antilope Strepsicheros (the kudu), a similar one of Antilope leucophas (the

blaauwbok)’, etc. (No other antelopes are mentioned.) This blue antelope was bought

by H. Schlegel for the museum in Leiden.

4. The Muséum National d’Histoire Naturelle in Paris has the mounted skin of an

adult male. Mohr (1967: 37—40) suggested that this skin was transferred in 1795 from

the collection of Willem V in The Hague, Holland. She recognized a certain similarity

between the mounted skin and the animal shown on a drawing made in South Africa

by Robert Jacob Gordon (1743-1795). Mohr did not know that Gordon had sent a

blue antelope specimen to Holland. J. N.S. Allamand (1778) wrote about the

‘tzeiran’ (the blue antelope) without any indication that he had received one from

South Africa. However, in the next volume of this Holland edition of Buffon’s

Histoire Naturelle, Allamand (1781: 38) added a footnote to the article about the

bontebok with information concerning the ‘pasan’ [= Oryx gazella| and the ‘tzeiran’:

‘M. Gordon m/’a envoié les peaux de lun & de lautre, qu'il a tués au Cap de Bonne

Esperance’. This short passage corroborates Mohr’s suggestion that the Paris blue

antelope came from Holland, and it is likely that it was forwarded from the Cape of

Good Hope by R. J. Gordon (Rookmaaker 1989: 301).

5. The Zoological Museum in Uppsala has a pair of horns that are the only remainder

of a complete mounted skin. Although C. P. Thunberg did not carry it from the Cape

of Good Hope, he received one in 1781 through the assistance of D. F. Immelman

(Rookmaaker 1989: 161, 300). The skin was present in 1846, but it seems probable

that it was destroyed later in the nineteenth century.

6. The Natural History Museum in London has a pair of horns of unknown prove-

nance. It has been suggested that these horns might have been bought by Thomas

Pennant in Amsterdam before 1781 (cf. Mohr 1967: 43). I would also like to record

here that there is the following item in the catalogue of the auction of Bullock’s

Museum in London, 10th day’s sale, Friday, May 14, 1819: ‘89 Blue Antelope,

A. Leucophaea.’ William Bullock (fl. 1795-1840) was the owner of a large Museum of

Natural Curiosities, started in Sheffield around 1795, moved to Liverpool in 1801, to

London in 1809, and exhibited from 1812 onwards in the Egyptian Temple on Picca-

dilly (Sweet 1970). The entire collection was auctioned between 29 April and 11 June

1819. The sale was attended by representatives from the major British and continental

museums, such as Walter Adam from Edinburgh, Coenraad Jacob Temminck from

Leiden, and Heinrich Lichtenstein from Berlin. There is an annotated copy of the

sales catalogue in the Natural History Museum, London, which shows that the blue

antelope mentioned above was bought by Dr Leach for £1—1—0. William Elford Leach

(1790-1836) bought animals for the British Museum, where he was in charge of the

zoological collections from 1813 to 1822. It is debatable whether the animal sold was

in fact a blue antelope if one considers its rather low price and the fact that people like

Lichtenstein apparently showed no interest in what they would have known to be a

rare exhibit. It is possible that the specimen was in a poor state.

LIST OF SPECIMENS OF THE EXTINCT BLUE ANTELOPE 135

7. The Hunterian Museum in Glasgow preserves a complete skull with horns attached.

It is not known how the specimen came to Glasgow. Klein (1974: 110) suggested that

it was, in fact, a skull of a sable antelope, Hippotragus niger.

ADDITIONS TO THE LIST OF SPECIMENS

HORNS IN CAPE TOWN

Ozinsky (1989) reported that a pair of horns was donated to the South African

Museum in Cape Town. It came from the family collection of Mr J. Piek of Obser-

vatory in Cape Town, but the earlier history is not known. It is likely that the horns

belonged to the blue antelope. The horns are registered as catalogue number

SAM-ZM40759 (Fig. 2). The distance from tip to tip has been measured as 150 mm,

and the front length over the curvature of the right hand side is 611 mm (D. Drink-

row, Collection Manager, South African Museum, in litt. November 1990).

Fig. 2. Two views of the horns recently acquired by the South African Museum, Cape Town

(SAM-ZM40759). Length 611 mm.

136 ANNALS OF THE SOUTH AFRICAN MUSEUM

A SKULL IN AMSTERDAM

During a recent review of the collections of the Zoological Museum of Amster-

dam, Holland, Dr P. J. H. van Bree, Curator of Mammals, found a hippotragine skull

with attached horns without data (ZMA 18.623). The specimen was examined in

detail by Erdbrink (1988: 144-146, pls 10-12) and cautiously identified as ‘Hippotra-

gus cf. leucophaeus (Pallas, 1766); (?) 0, ad.’ (Figs 3-6). The caution is necessary,

because there are no other recent skulls of the blue antelope with which it could be

compared. The right horn is 505 mm long over the curvature and has 31 rings, the left

horn measures 495 mm and has 29-30 rings. The skull’s width over the orbit is

154 mm, the left lower premolar row is 44 mm, the left lower molar row 72 mm (see

Erdbrink 1988 for these and other measurements).

It is rather disconcerting that there is no information about the provenance of this

specimen. One possibility is that the skull was among other unidentified specimens

from the Haarlem Cabinet in 1866 (see below), and that the skin has since been dis-

carded, leaving the skull, which has remained unattended ever since. If, in fact, the

specimen does belong to the blue antelope, which seems likely, it would be difficult to

establish an alternative source.

EVIDENCE OF A SKIN ONCE IN HAARLEM

On 23 July 1805, Martinus van Marum (1750-1836), supervisor of the collections

of the Hollandsche Maatschappij der Wetenschappen in Haarlem, wrote a letter to

Fig. 3. Blue antelope skull in the Zoological Museum in Amsterdam, Holland (ZMA 18.623): right

lateral aspect.

LIST OF SPECIMENS OF THE EXTINCT BLUE ANTELOPE 137

J. A. de Mist acknowledging receipt of a small collection of Cape specimens: ‘I was

pleasantly surprised to-day by some items of natural history, consisting of six bottles

of snakes in spirit, a box with planks of African woods and the skin, together with the

horns, of the Blue Buck, which your Honour sent to me for the Natural History

Cabinet of our Society’ (translated by Lefebvre & De Bruyn 1976: 239).

Jacob Abraham Uytenhage de Mist (1749-1823) was one of three secretaries of

the Dutch town Kampen from 1766, and was elected a member of the National Con-

vention in 1796. In 1802, he was sent as Commissioner General to the Cape of Good

Hope. He stayed three years in the southern part of Africa. From 9 October to the

end of November 1803 he travelled northwards from the Cape into the interior,

accompanied by a large group of people. Among them was Martin Heinrich Carl

Lichtenstein (1780-1853), from 1813 director of the Zoological Museum in Berlin,

who published an account of his stay in South Africa in 1811-1812. Lichtenstein kept

the journal of De Mist’s expedition, which included only a few zoological details and

nothing about the blue antelope (as summarized by Molsbergen 1922: 167-189).

Fig. 4. Blue antelope skull in the Zoological Museum in Amsterdam, Holland (ZMA 18.623): frontal

aspect. Width over the orbit 154 mm.

138 ANNALS OF THE SOUTH AFRICAN MUSEUM

When De Mist returned to Holland, he took with him a small collection of

natural history specimens and some ethnographical material. Van Marum (1812) gave

a report on these specimens, the preparation of which was delayed because he had

hoped to receive their descriptions from Lichtenstein. De Mist’s gift consisted of

333 species of insects (put in order by Lichtenstein in South Africa), and besides it

included: ‘De Huiden van een Leeuw, van een Tyger, en van een Blaauwe Bok

(Antilope Leucophaea), alsmede de Horens van verscheidene soorten van Antilopen

en van een Rhinoceros’ (Van Marum 1812: 166). [Translated: The skins of a lion, a

leopard, and a blue antelope (Antilope Leucophaea), as well as the horns of several

species of antelopes and those of a rhinoceros. |

One wonders how much Lichtenstein knew about the specimen brought home by

De Mist. Considering his understanding of the blue antelope’s extinction in South

Africa and his personal acquaintance with De Mist, it is almost impossible that he

would not have heard about it, and may even have examined it. In his travel book,

Lichtenstein (1811: 265) said that ‘Im Jahre 1800 war noch einer geschossen, dessen

Fell jetzt in Leyden aufbewahrt wird; seitdem aber hat man Keine mehr gesehen.’ In a

paper about antelopes written just a few years later, Lichtenstein (1814: 160) clearly

described a skin of the blue antelope that he had examined personally: ‘Das Fell, nach

welchem ich beschreibe, war leider das letzte das gesehen worden ist. Man hat seit

1799, wo dieses Thier geschossen ward, keine wieder eingetroffen.’ He did not say

where he saw the animal. However, it is very tempting to assume that Lichtenstein

was talking about De Mist’s specimen of the blue antelope. Why did Lichtenstein

write, in 1811, that the animal was ‘now in Leiden’? He returned to Europe in June

1806 and he may have heard that De Mist took his specimen to Holland. Still, Leiden

does not fit the facts, because the collection was given to Haarlem in 1805 and De

Mist is not known to have lived in Leiden. Maybe further research can clarify these

statements.

Considering the above references, I suggest the following history for De Mist’s

blue antelope. The specimen probably was killed just before 1800, since which time it

Fig. 5. Blue antelope skull in the Zoological Museum in Amsterdam, Holland (ZMA 18.623): ventral

aspect with upper tooth row.

LIST OF SPECIMENS OF THE EXTINCT BLUE ANTELOPE 139

was kept in a private collection or in a shop of natural history specimens; it was given

to De Mist or he may have bought it. He took the skin to Holland arriving on 8 July

1805, after which he donated it to the society in Haarlem. From 1812 onwards, the

history of the skin becomes obscure.

We need to consider the possibility that the animal was a roan (Hippotragus

equinus) rather than a blue antelope. In the absence of any evidence, it is useless to

speculate. However, since it is likely that Lichtenstein’s description was drawn up

after De Mist’s specimen, it is almost certain that it was in fact a blue antelope and not

a roan. If this conclusion is correct, it appears that the Dutch Society of Sciences in

Haarlem possessed two mounted skins of this rare species from 1805 onwards, i.e.

specimen number 3 mentioned in the list of known specimens (see p. 133) and the one

brought by De Mist. Tuijn (1971) mentioned both animals in passing, without giving

further details. As stated above, the Haarlem society auctioned one blue antelope in

April 1842 and that one passed to the Rijksmuseum van Natuurlijke Historie in

Leiden.

It is, of course, important to establish which specimen of the blue antelope was

bought by the Leiden Museum: the one bequeathed by Van Lennep or the one

received in 1805. It is probable that Mohr (1967: 35) gave the correct clue when she

said that the specimen still in the Leiden Museum ‘hat mit dem urspringlichen Bild

bei Buffon [= Allamand] 1778—und nur mit diesem—derart viele Ahnlichkeiten,

daB sie sich gegenseitig beeinflu8t haben mussen.’ This can be safely followed and jus-

tifies the conclusion of Husson & Holthuis (1969) that the Leiden blue antelope was

one of the syntypes of Antilope leucophaea Pallas, 1766.

Surprisingly, the second Haarlem blue antelope, the one donated by De Mist, has

disappeared from the records. In fact, there is not a single clue as to what happened to

Fig. 6. Blue antelope skull in the Zoological Museum in Amsterdam, Holland (ZMA 18.623):

mandible. Left premolar row 44 mm, molar row 72 mm.

140 ANNALS OF THE SOUTH AFRICAN MUSEUM

this skin with horns; a skeleton or skull were never mentioned. One reason for this

must be that interest in the Cabinet of the Society in Haarlem dwindled after the

death of Van Marum. The history of the Society and the Cabinet, written by Bierens

de Haan (1941, 1952), shows that Van Marum’s successors, J. G. S. van Breda

(secretary 1838-1864) and E. H. von Baumhauer (secretary 1864-1885) did not care

properly for the zoological specimens of the museum. In 1866, a committee of investi-

gation reported that a large part of the collection had lost its value due to age and lack

of maintenance. Obviously, nobody was interested in keeping records about the

collection. In November 1866, the largest part of the Haarlem Cabinet was transferred

to the Museum of the Zoological Gardens in Amsterdam; much was bought by the

Amsterdam Zoo but some of the more valuable specimens were only given on loan,

like the Duchassaing & Michelotti collection of sponges (Wiedenmayer 1977: 252). An

inventory was made at that time, but it only included ‘all sorts of lower animals’ (Smit

1986: 108) and birds or mammals were not mentioned during the transactions (Bierens

de Haan 1952: 266). It is unlikely that a complete stuffed blue antelope was included.

ACKNOWLEDGEMENTS

I am grateful to Dr P. J. H. van Bree for sharing the information of his discovery

of the blue antelope skull in the Zoological Museum of Amsterdam and for general

encouragement. Dr D. P. Bosscha Erdbrink kindly helped by supplying photographs

of the skull in Amsterdam. An earlier version of this paper was read by

Dr C. Smeenk, Dr L. B. Holthuis, Dr R. G. Klein and Mr R. Rau and I am thankful

for their comments. Mr W. H. Holleman provided information about the horns in the

Albany Museum.

REFERENCES

ALLAMAND, J. N. S. 1778. Histoire naturelle, générale et particuliére servant de suite a histoire des

animaux quadrupédes. Nouvelle édition, Supplément 4. Amsterdam: J. H. Schneider.

ALLAMAND, J. N. S. 1781. Histoire naturelle, générale et particuliére servant de suite a histoire des

animaux quadrupédes. Nouvelle édition, Supplément 5. Amsterdam: J. H. Schneider.

BIERENS DE HAAN, J. A. 1941. De geschiedenis van een verdwenen Haarlemsch Museum van Natuur-

liike Historie: het Kabinet van Naturalien van de Hollandsche Maatschappij der Wetenschappen

1759-1866. Haarlem: Erven F. Bohn.

BIERENS DE Haan, J. A. 1952. De Hollandsche Maatschappij der Wetenschappen 1752-1952.

Haarlem [n.p.].

[BULLocKk, WILLIAM]. 1819 [1979]. Sale catalogue of the Bullock Museum 1819. A facsimile reprint

with manuscript prices and buyers names. London: Harmer Johnson & John Hewett.

ERDBRINK, D. P. Bosscua. 1988. Protoryx from three localities East of Maragheh, N. W. Iran. Pro-

ceedings of the Koninklijke Nederlandse Akademie van Wetenschappen (B) 91 (2):101-159.

Husson, A. M. & Hortuuis, L. B. 1969. On the type of Antilope leucophaea Pallas, 1766, preserved

in the collection of the Rijksmuseum van Natuurlijke Historie, Leiden. Zoologische Medede-

lingen, Leiden 44 (11): 147-157.

KLEIN, R. G. 1974. On the taxonomic status, distribution and ecology of the blue antelope, Hippo-

tragus leucophaeus (Paltas, 1766). Annals of the South African Museum 65 (4): 99-143.

KLEIN, R. G. 1987. The extinct blue antelope. Sagittarius, Cape Town 2 (3): 20-23.

KouL, F. F. 1886. Ueber neue und seltene Antilopen des k.k. Naturhistorischen Hofmuseums.

Annalen des k.k. Naturhistorischen Hofmuseums, Wien 1: 75-86.

LEFEBVRE, E. & DE Bruyn, J. G. 1976. Martinus van Marum, life and work 6. Leyden: Noordhoff

International.

LIST OF SPECIMENS OF THE EXTINCT BLUE ANTELOPE 141

MaruM, Martinus vAN. 1803. Naamlijst van het Naturalien-Kabinet van de Bataafse Maatschappij

der Wetenschappen te Haarlem. Natuurkundige Verhandelingen der Hollandsche Maatschappij

van Wetenschappen, Haarlem 2 (1): i—xxxii.

Marum, Martinus VAN. 1812. Naturalia en andere byzonderheden, in den jaare 1805 van de Kaap

de Goede Hoop, en deszelfs onbeschaafde volksstammen medegebragt, en aan het Naturalien-

Kabinet der Maatschappij ten geschenke gegeven, door Mr. J. A. de Mist. Natuurkundige Ver-

handelingen der Hollandsche Maatschappij van Wetenschappen, Haarlem 6 (1): 159-169.

Monr, E. 1967. Der Blaubock, Hippotragus leucophaeus (Pallas, 1766): eine Dokumentation. Mam-

malia Depicta: 1-81. Hamburg & Berlin: Paul Parey.

MotrsBERGEN, E. C. GopEE. 1922. Reizen in Zuid-Afrika in de Hollandse tijd, 2: Tochten naar het

Noorden 1686-1806. ’s Gravenhage: Martinus Nijhoff.

Ozinsky, S. 1989. Rare blue antelope horns found. Sagittarius, Cape Town 4 (3): 31.

RENSHAW, G. 1921. The blaauwbok (Hippotragus leucophaeus Pall.). Journal of the Society for the

Preservation of the Fauna Empire (n.s.) 1: 24-26.

ROOKMAAKER, L. C. 1989. The zoological exploration of Southern Africa 1650-1790. Rotterdam &

Brookfield: A. A. Balkema.

SCHONLAND, S. 1902. Report of the Director. In: Report of the Committee of the Albany Museum,

for the year 1901: 2-5. Cape Town: W. A. Richards & Son.

SCHREYER, J. 1681. Neue Ost-Indianische Reise-Beschreibung. Leipzig: J. C. Wohlfart.

Smit, P. 1986. Hendrik Engel’s alphabetical list of Dutch zoological cabinets and menageries.

Amsterdam: Rodopi.

SwEET, J. M. 1970. William Bullock’s collection and the University of Edinburgh, 1819. Annals of

Science 26 (1): 23-32.

Tuun, P. 1971. The cabinet of natural curiosities. Jn: ForBes, R. J. ed. Martinus van Marum, life

and work 3. Leyden, Noordhoff International.

WIEDENMAYER, F. 1977. A monograph of the shallow water sponges of the Western Bahamas. Basel

& Stuttgart: Birkhauser.

6. SYSTEMATIC papers must conform to the International 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

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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.

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Note punctuation in the above example:

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L. C. ROOKMAAKER

ADDITIONS AND REVISIONS TO THE

LIST OF SPECIMENS OF THE

EXTINCT BLUE ANTELOPE

(HIPPOTRAGUS LEUCOPHAEUS)

VOLUME 102 PART 4 NOVEMBER 1992 ISSN 0303-2515

OF THE SOUTH AFRICAN

yo ~ MUSEUM

CAPE TOWN

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

FiscHer, P. H. 1948. Données sur la résistance et de la vitalité des mollusques. Journal de conchyliologie 88 (3): 100-140.

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

expérimentale et générale 74 (33): 627-634.

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

Magazine of Natural History (13) 2 (17): 309-320.

Koun, A. J. 1960b. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bulletin of

the Bingham Oceanographic Collection, Yale University 17 (4): 1-51.

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

pologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika ausgefiihrt in den Jahren

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(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 102 °+#2Band

November 1992 November

Part 4 Deel

TWO NEW SPECIES OF OIL-COLLECTING

BEES OF THE GENUS REDIVIVA FROM THE

SUMMER RAINFALL REGION OF

SOUTH AFRICA

(HYMENOPTERA, APOIDEA, MELITTIDAE)

V. B. WHITEHEAD

KES REINER

Cape Town Kaapstad

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D1745

TWO NEW SPECIES OF OIL-COLLECTING BEES OF THE GENUS REDIVIVA

FROM THE SUMMER RAINFALL REGION OF SOUTH AFRICA

(HYMENOPTERA, APOIDEA, MELITTIDAE)

V. B. WHITEHEAD

Entomology Department, South African Museum, Cape Town

K. E. STEINER

National Botanic Institute, Kirstenbosch, Claremont

(With 9 figures and 4 tables)

[MS accepted 25 May 1992]

ABSTRACT

Two species of oil-collecting bees of the genus Rediviva are described from the summer rainfall

region of southern Africa. Rediviva pallidula is a high-altitude bee that closely resembles R. neliana

Cockerell, but is distinguished by its paler pubescence on the leg extremities and metasomal terga and

by differences in the shape of sternum seven and genital capsule among males. Females collect oil

from six Diascia species (Scrophulariaceae) and two species of terrestrial orchids. Rediviva saetigera is

the only oil-collecting bee in the summer rainfall area with short tarsomeres 2, 3 and 4 on the front

legs of the female, and with modified hairs for collecting oil on the front basitarsus and tibia only.

There are also distinctive long pollen-collecting hairs on the frons of females. The oil hosts of R. saeti-

gera are two species of shrubby trees, Bowkeria cymosa MacOwan and Anastrabe integerrima

E. Meyer ex Benth. (Scrophulariaceae), the former occurring on the eastern Transvaal escarpment

and the mountains of Swaziland and the latter in coastal forests from East London to northern Natal

and Mogambique.

CONTENTS

PAGE

NTNGROCMUIC HOM Me ew ry a tal einen tial Riedy N MMS esc BN eye. Lina aN Teli Utd 143

Wie thi oc Sipser ee UA eu cera ces Sie sual he se aut ole Salldaid ae 144

IDESCHIPEONSHR ree re ae Mane ne nies aan sn en eat SPR 25 aa 145

INCAIVIV OND QUIGUIGESPENOV. Ctra wo ei aides Bese eee oe aes ee 145

INCU SOMATA Ss, WONTS Ws dis Sooo aa ao tard eo ee Ge Se oto ene IS)5)

NC SMOMOCTSMONS gob on Cee SOE eS Bee oe eo ee ee RE nt er ee 164

INCICREN CE Sewer EER eS ee aac an Ono ie eee neni enc) wa wane 164

INTRODUCTION

We have been investigating the relationships between oil-collecting bees of the

genus Rediviva and their oil and nectar host plants in southern Africa (Whitehead et

143

Ann. S. Afr. Mus. 102 (4), 1992: 143-164, 9 figs, 4 tables.

144 ANNALS OF THE SOUTH AFRICAN MUSEUM

al. 1984; Whitehead & Steiner 1985; Steiner & Whitehead 1988, 1990, 1991). Rediviva

bees can be conveniently grouped according to their occurrence in the summer and

autumn rainfall regions of the eastern Cape and eastern Transvaal, and the winter

rainfall areas of the western Cape. Bees from these two regions do not overlap either

temporally or geographically.

In the winter rainfall region there are four described species—R. peringueyi

(Friese), R. intermixta (Cockerell), R. longimanus Michener, and R. emdeorum Vogel

& Michener (Michener 1981; Vogel & Michener 1985), and 10 undescribed species

(Whitehead & Steiner in prep). The summer rainfall group is also made up of four

described species, R. neliana Cockerell, R. politissima (Cockerell), R. rufocincta

(Cockerell) and R. colorata Michener (Michener 1981). Rediviva neliana and R. poli-

tissima do not differ in male genitalia nor on non-meristic characters, except colour of

pubescence. At present, these two species are considered to constitute part of a single

highly variable species, which may, with more detailed study, result in the recognition

of one or more additional species. Two other Rediviva species that are readily distin-

guishable are described below. Although detailed morphological studies have only

recently started, it would appear from data available at present that the species from

these two areas form a monophyletic group. A phylogenetic analysis of the group will

be included with the revision of the genus.

METHODS

Measurements were made with a calibrated eye-piece graticule, using a dissecting

microscope. Forelegs were removed and mounted on card and each segment meas-

ured separately (Steiner & Whitehead 1990, 1991). Wing length is the distance from

the outer edge of the tegula to the wing tip. Body length was measured from the

anterior ocellus to the tip of the abdomen and, although there was some variability

due to the position of the abdomen, this measurement was found to be a good esti-

mate of body size (Steiner & Whitehead 1990). Male genitalia and the associated

sterna were extracted with a bent needle after the bee had been relaxed. The genital

capsule with associated sterna were macerated in cold 10 per cent KOH overnight,

rinsed with water, acidified with acetic acid, rinsed again and then stored in glycerine.

All dissected parts were placed in microvials attached to the relevant insect. Mouth-

parts were removed after treating the whole head as outlined above. Both mouth-

parts and genitalia were drawn in glycerine using a drawing tube attached to a dissect-

ing microscope. When drawing the mouth-parts, pressure had to be applied to the

coverslip to flatten some of the structures.

The terms used for the various structures are those of Michener (1981) and for

the description of surface sculpture those of Eady (1968). For leg orientation we have

used the terminology of Roberts & Brooks (1987), whereby the leg is considered to be

a cylindrical lateral projection at right angles to the long axis of the body. The symbols

S and T with the appropriate numeral are used to indicate the sterna and terga of the

metasoma; S1, for example, refers to the first sternum of the metasoma.

TWO NEW SPECIES OF OIL-COLLECTING BEES 145

DESCRIPTIONS

Rediviva pallidula sp. nov.

Figs 1-5

Rediviva politissima Manning & Brothers, 1986: 108-111 (part.—Sani Flats specimens only).

Diagnosis

Integument black, females with brown pubescence on dorsal areas of mesosoma,

on rest of the body pubescence white to pale straw-coloured. Male pubescence similar

but paler. Malar space relatively long, one-quarter to one-fifth as long as wide. In

areas of overlap with R. neliana, R. pallidula females can be distinguished by the pres-

ence of white pubescence on the discs of metasomal terga two, three and four; in

R. neliana females, the pubescence on the disc of metasomal tergum four is always

black. The hairs on the leg extremities of R. pallidula are white to pale yellow in con-

trast to the dark brown to black hairs of R. neliana. Males of the two species can be

separated by the shape of sternum seven as well as differences in the genital capsule.

Type locality

Natal, Sani Pass, 2 490 m, 2929CB, collected on flowers of Diascia integerrima

Benth. (Scrophulariaceae).

Etymology

Pallidus, Latin for somewhat paler, referring to the paler colour of the pubes-

cence in contrast to that of R. neliana with which it may be confused.

Material examined

Type material. Holotype: SAM—HYMB0001, female, Natal, Sani Pass, 2 490 m,

2929CB, V.B. Whitehead, 16 Feb. 1986. Allotype: SAM-—HYMB0002, male,

Lesotho, Sani Top, 2 740 m, 2929CB, V. B. Whitehead, 15 Feb. 1986. Paratypes

(46 99, 25 0C')—Cape Province: 19, Lady Grey, Joubert’s Pass, 2 140 m,

3027CB, K. E. Steiner, 13 Jan. 1985; 1 9, Lady Grey, Joubert’s Pass, 3027CB, K. E.

Steiner, 8 Jan. 1986; 1 9, Lady Grey, Joubert’s Pass summit, 3027CB, V. B. White-

head, 8 Jan. 1984; 3 99, 1 Oo’, Ben Macdhui, 3027DB, K. E. Steiner, 12 Jan. 1984;

1 9, Naudesnek, 3028CA, 2 240 m, K. E. Steiner, 28 Jan. 1986; 1 9, Naudesnek,

3028CA, 2 370 m, K. E. Steiner, 29 Jan. 1986; 1 9, Naudesnek, 2 170 m, 3028CA,

K. E. Steiner, 31 Jan. 1986; 1 o’, Naudesnek, 2 280 m, 3028CA, K. E. Steiner,

19 Jan. 1989; 1 CO’, Naudesnek, 2 300 m, 3028CA, K. E. Steiner, 19 Jan. 1989; 1 9,

1 &, Naudesnek, 3028CA, V. B. Whitehead, 11 Jan. 1985; 1 9, 1 Oo’, Naudesnek,

3028CA, V. B. Whitehead, 12 Jan. 1985; 1 9, Naudesnek, 2 370 m, 3028CA, V. B.

Whitehead, 28 Jan. 1986; 1 9, Naudesnek, 3028CA, V. B. Whitehead, 19 Jan. 1989;

1 9,2 OC’, Naudesnek, 2 480 m, 3028CA, V. B. Whitehead, 19 Jan. 1989; Lesotho:

1 Co’, Blue Mountain Pass, 2928AC, V. B. Whitehead, 7 Jan. 1985; 1 9, 1 C’, Blue

Mountain Pass, 2928AC, V. B. Whitehead, 24 Jan. 1985; 1 9, 4 O'C’, Blue Mountain

Pass, 2928AC, V. B. Whitehead, 25 Jan. 1985; 1 9, Lebelonyane Pass, 3028AA,

K. E. Steiner, 14 Jan. 1985; 1 9, Lebelonyane Pass, 2 400 m, 3028AA, K. E. Steiner,

27 Feb. 1990; 1 9, 1 C&’, Lebelonyane Pass, 3028AA, V. B. Whitehead, 14 Jan. 1985;

146 ANNALS OF THE SOUTH AFRICAN MUSEUM

1 9, Lebelonyane Pass, 3028AA, V. B. Whitehead, 27 Feb. 1990; 499, 1G,

Mapholaneng, 2828BB, K. E. Steiner, 13 Feb. 1986; 1 oO’, Mapholaneng, 2828BB,

V. B. Whitehead, 13 Feb. 1986; 1 9, Oxbow, 2828DC, K. E. Steiner, 23 Jan 1985;

2 22, Oxbow, 2828DC, V. B. Whitehead, 23 Jan. 1985; 1 co’, Sani Pass, 2 720 m,

2929CB, K. E. Steiner, 14 Feb. 1986; 4 99, Sani Pass, 2 720 m, 2929CB, K. E.

Steiner, 15 Feb. 1986; 1 9, Sani Pass, 2 720 m, 2929CB, V. B. Whitehead, 14 Feb.

1986; 1 oO’, Sani Pass, 2 750 m, 2929CB, V. B. Whitehead, 14 Feb. 1986; 1 0’, Sani

Pass, 2 720 m, 2929CB, V. B. Whitehead, 15 Feb. 1986; Natal: 4 O'C', Sani Pass,

2 080 m, 2929CB, K. E. Steiner, 21 Jan. 1985; 1 9, Sani Pass, 2 250 m, 2929CB,

K. E. Steiner, 16 Feb. 1986; 1 9, Sani Pass, 2 290 m, 2929CB, K. E. Steiner, 16 Feb.

1986; 1 9, Sani Pass, 2 440 m, 2929CB, V. B. Whitehead, 15 Feb. 1986;1 9,2 00,

Sani Pass, 2 350m, 2929CB, V. B. Whitehead, 16 Feb. 1986; 2 99, Sani Pass,

2 490 m, V. B. Whitehead, 16 Feb. 1986.

Other material (112 99, 9 Cco')—Cape Province: 49, Carlisle’s Hoek,

3027DD; 35 99, Naudesnek, 3028CA; Lesotho: 6 99, 1c’, Blue Mountain Pass,

2928AC; 1 9, Lebelonyane Pass, 3028AA; 1 0’, Makapung Dip, 2929AC; 5 99,

Mapholaneng, 2828BB; 499, 20°C’, Moteng Pass, 2828DA; 21 9Q Sani Pass,

2929CB; Natal: 36 9, Sani Pass, 2929CD.

Description

Female

Measurement and ratios. Holotype: body 11,7 mm, forewing 10,7 mm. Other

material: means given in Table 1 for 15 specimens selected at random from six

different localities where possible, otherwise all specimens at particular locality used.

Integumental colour. Black, except apical one-fourth of mandibles and flagellar

segments 3-10 brown; wings lightly tinted with brown; stigma and costa light brown,

other veins dark brown to black; apical margins of metasomal terga T2 to T4 light

brown, apical margins of sterna S1 to S4 light brown becoming translucent.

Structure. Head wider than long, 3,8:2,9 mm (type); means of head measure-

ments from sample of the paratypes from various collection localities presented in

TABLE 1

Mean lengths (+SD) of forelegs, body, foreleg/body ratios and forewings of females

(n = 67) and males (n = 27) of R. pallidula from all localities.

Females Males

Length Range Length Range

(mm) (mm) (mm) (mm)

Foreleg 12,2+0,49 10,3-13,3 08,9+0,25 08,8-09,3

Body 10,8+0,26 09,8-12,5 09,9+0,34 09,9-10,1

Foreleg/body 01,1+0,03 00,9-01,3 00,9+0,03 00,8-01,0

Forewing 08,8+0,15 08,2-09,3 08,2+0,24 07,9-08,5

TWO NEW SPECIES OF OIL-COLLECTING BEES 147

Fig. 1. Rediviva pallidula sp. nov. Female, mouth-parts. A. Labium, anterior view. B. Distal part

of labium, posterior view. C. Fragmentum of prementum, mentum and lorum, anterior view.

D. Maxilla, outer view. E. Basistipital process of maxilla. F. Inner view of maxilla to show galeal

comb. G. Labrum.

148 ANNALS OF THE SOUTH AFRICAN MUSEUM

1mm

Fig. 2. Rediviva pallidula sp. nov. A. Ventral and B. Dorsal views of oil-collecting segments of

female fore tarsus. C. Hind tibia and basitarsus of female (hairs removed). D-—F. Dorsal view of S8

of males to show variation. D. S8 of male from Naudesnek. E. S8 of male from Sani Pass. F. S8 of

male from Blue Mountain Pass.

TWO NEW SPECIES OF OIL-COLLECTING BEES 149

Table 2; inner orbits converging above, distance between eyes greater than length of

eyes, 2,5:2,3 mm (holotype), with mean of 2,5:2,2 mm (n=33) for paratypes

(Table 2); inner distance between lateral ocelli slightly greater than ocellocular dis-

tance (0,6 mm); malar space one-quarter to one-fifth as long as wide (holotype

L: W = 0,24); mean L:W for populations from all the collection areas in Table 2.

Mouth-parts: glossa triangular, one-half as long as prementum, extending beyond

labial palps; paraglossae extending one-quarter length of glossa; labial palps nearly

reaching tip of glossa, basal segment one-third length of palp, terminal segment

slender, well sclerotized (Fig. 1A—B); ligular arms occupying basal two-thirds of pre-

mentum (Fig. 1A); premental fragmentum, mentum and lorum as in Figure 1C;

subligular process and basiglossal sclerite (Fig. 1B); cardo slightly shorter than stipes,

stipes 3,5 times as long as wide, lateral margin with scattered hairs, extremely long

posteriorly (Fig. 1D); basistipital process short (Fig. 1E); maxillary palp not extending

beyond tip of galea, six-segmented, segments longer than wide; galea with broadly

pointed apex, short hairs along apical and lateral margins (Fig. 1D), galeal comb dis-

tinct with 17 teeth (Fig. 1F); mandibles curved, strong preapical and apical tooth

(Fig. 4B); labrum triangular, twice as wide as long, with dense long hairs on margin

(Fig. 1G). Antennae with first flagellar segment about three-quarters length of scape

and longer than segments 2 plus 3, segments 4 to 9 subequal and almost as long as

broad, segment 10 twice as long as broad terminating in a shiny, oblique, slightly

convex surface. Front legs long, varying from 10,3 mm to 13,3 mm, depending on col-

lection locality (Table 1), tarsus making up about one-third of the leg length;

tarsomeres 2—4 with dense, finely branched hairs on dorsal and lateral surfaces, spatu-

late hairs on sides of segment 2 but only at apex of segments 3 and 4 and isolated on

segment 5; four terminal segments constitute oil-collecting apparatus in this species;

basitarsus bearing stout unbranched hairs (Fig. 2A—B). Hind legs with tibia nearly as

wide as basitarsus; basitarsus roughly ovoid, distal dorsal angle rounded, spurs pale

brown, finely dentate (Fig. 2C); wings (Fig. 4A) with second submarginal cell slightly

broader than long (1:0,8), receiving first recurrent vein beyond middle (20:11), third

submarginal one-and-a-half times as broad as long, receiving second recurrent vein

beyond middle (27:20), basal vein nearly three times as long as first abscissa of RS,

meeting Cu at junction of Cu—V; jugal lobe of hind wing less than half length of vanal

lobe (12:32); hamuli 11-12. Propodeal triangle small, nearly parallel-sided at apical

TABLE 2

Mean measurements (+SD) of head width (HW), head length (HL), interocular distance

(IOD), eye length (EL) and the length/width ratio of the malar space (MS-L/W) of

R. pallidula females (n = 33) and males (n = 21) from all localities.

HW(mm) HL(mm) J0D(mm) EL(mm) MS-L/W

Females 3,7+0,04 3.0+0,04 2,540.07 2.2+0,03 0,2+0,01

Range 3,4-3,7 /oei=3)A0) 2,4-2,6 DP)

Males aS LOROlk 2,8+0,11 Dekh) N22 2,0+0,89 0,2+0,02

Range 3,4-3,6 A349) 2,1-2,4 AVP 3k

150 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 3. Rediviva pallidula sp. nov. Male genitalia and terminal sterna. A. S6, ventral view. B. S7,

dorsal (left) and ventral (right) view. C. S7, side view. D. Dorsal and ventral view of S8. E. S8, side

view. F. Genital capsule, dorsal view (left) and ventral view (right). G. Genital capsule, side view.

TWO NEW SPECIES OF OIL-COLLECTING BEES 151

smm

1mm

Fig. 4. Rediviva pallidula sp. nov. Female. A. Right wings.

B. Right mandible, outer view.

two-thirds, broadening slightly at base, shiny and finely rugose; rest of propodeal

surface shiny but coarsely rugose. Apical margins of metasomal S2 to S4 concave, as is

gradulus of S2. Pygideal plate dull, faintly rugose with some coarse punctures basally.

Sculpture. Head: clypeus with basal area coarsely punctate, surface between

punctures shiny, punctures becoming finer and less dense towards apex, apical margin

practically impunctate and finely reticulate; frons coarsely punctate either side of

frontal ridge, becoming more finely punctate in para-ocular areas and towards anterior

ocellus, area between ocelli and preoccipital ridge coarsely punctate; small triangular

area between lateral ocelli and preoccipital ridge shiny and impunctate or with scat-

tered fine punctures. Mesosoma: scutum finely punctate on disc, punctures coarser

lateral to parapsidal line; median line distinct, extending one-third length of segment;

scutellum finely punctured, coarser posteriorly and laterally; metanotum finely punc-

tured; propodeum finely rugose, triangle shiny and faintly strigose. Metasoma: T1

shiny with fine scattered punctures on apical margin; T2, T3 and T4 shiny with denser

fine punctures, TS with coarse punctures, surface between punctures finely rugose; S2

to SS finely punctured, areas between punctures finely reticulate, apical margins

impunctate.

152 ANNALS OF THE SOUTH AFRICAN MUSEUM

Vestiture. Head: brush of pale yellow hairs at centre of ridge on anterior margin

of clypeus, rest of clypeus covered with strong white hairs, long unbranched pale

yellow hairs on median area with a few scattered black hairs; supraclypeal and para-

ocular areas with white, finely branched hairs, scattered black hairs on para-ocular

areas; scattered dark hairs on vertex; genal area with long white plumose hairs. Meso-

Transvaal

Cape

Province

0 100 200 km

{L-ues te

Fig. 5. Known distribution of R. pallidula (@) and R. saetigera (A). L = Lesotho, O.F.S. = Orange

Free State, S = Swaziland, T = Transkei.

TWO NEW SPECIES OF OIL-COLLECTING BEES 153

soma: margins of scutum, scutellum and metanotum densely covered with light brown

hair, interspersed with some dark hairs; hair shorter and more scattered on disc of

scutum and scutellum; finer and more branched on propodeum except triangle which

is bare; episternum and sternum covered in dense white hair; hairs on tibiae and

tarsae light brown, tarsal segments 2—5 of front legs with finely branched oil-collecting

hairs on dorsal and lateral areas, bare underneath; hairs on anterior area of middle

legs paler with a dark patch at apex of tibia; anterior face of hind tibia and basitarsus

with a finely branched mat of pale hairs through which strong pale unbranched hairs

protrude; posterior face with strong unbranched pale brown hairs only; penicillum of

light brown hair on distal margin of basitarsus. Metasoma: white apical hair bands on

S1 to S4, fimbrium on T4 black dorsally, white laterally; white decumbent hairs

basally on T2, T3 and T4, with some scattered black hairs in this area on T4; subapical

band of long erect hairs on S82 to SS.

Male

Measurement and ratios. Allotype male, body 9,8 mm, forewing 8,7 mm. Means

for various localities are given in Table 1.

Integumental colour. Black, except tip of mandibles, under-side of antennae,

posterior surface of hind tibia, leg extremities, lateral area of T1, tegulae, costal vein,

and bases of other veins piceous to dark brown.

Structure. Head: wider than long, 3,4:2,6 mm; distance between eyes 2,1 mm,

eye length 1,9 mm (allotype). Means for males in Table 2. Ocellocular distance equal

to distance between lateral ocelli; malar space one-quarter to one-fifth as long as wide,

mean L:W=0,2 mm; first flagellar segment more than two-thirds length of scape

(27:39), slightly shorter than length of flagellar segments 2 plus 3 (27:31), segment 2

shorter than 3, segments 3-10 subequal, segment 11 slightly longer, terminating in

convex, shiny oblique area. Mesosoma: forelegs not obviously lengthened, equal to or

shorter than body, FL: B =0,8-1,0 (n = 27) (Table 1); hind tibia nearly twice as wide

as basitarsus (45:24); tibial spurs pale, finely serrated, basitibial plate covered with

fine white hairs; prominent dorsal spur at distal end of fore tibia. Wing venation as in

female, jugal lobe more than one-third length of vanal lobe (29:80). Metasoma: S2 to

S5 with apical margins straight or shallowly concave at middle; S6 with disc concave,

shiny, covered with short white hairs, lateral lobes with dense tufts of white hairs,

apical margin emarginate, with short white hairs (Fig. 3A); short apical lobes of S7

with long hairs on apical and lateral margins; lateral lobes striated, translucent, ovate,

projecting upwards (Fig. 3B—C); S8 with margin of sloping tip variable (Figs 2D-F,

3D-E). Genitalia (Fig. 3F—G): gonostyli slightly shorter than penis valves, narrow

with large translucent membranous area below, apical setae straight, projecting at

right angles; volsellae large with distinct teeth on opposing surfaces of digitis and

cuspis, opening posterolaterally.

Sculpture. As in female on head and mesosoma, punctures coarser on sterna and

terga of metasoma.

Vestiture. Head: labrum with white to straw-coloured hairs on anterior margin;

longer unbranched hairs on outer and inner margins of mandibles; tufts of branched

hairs on anterior ridge of clypeus, dense long white hairs on rest of clypeus, paraocular

154 ANNALS OF THE SOUTH AFRICAN MUSEUM

areas, frons and genal area; shorter sparser dark hairs on vertex, scattered dark hairs

along inner eye margins. Mesosoma: scutum covered with finely branched straw-

coloured hairs, shorter on disc with scattered black hairs; scutellum and metanotum

with long straw-coloured branched hairs; episternal areas and three basal segments of

legs covered in long white branched hairs; hairs on under-side of tibia and tarsus of

forelegs light brown. Metasoma: dense apical white hair bands on T1 to T6, fimbrium

on 17 light brown; erect shorter and sparser white hairs basally on disc of T2 to TS,

some black hairs mixed with white on disc of T3 to T5, becoming progressively denser

towards apex of metasoma; sparser preapical hair bands on S1 and S3 to S6, S7 with

tufts of pale brown hairs on lateral lobes; long white hairs on under-side of S8 project-

ing beyond the spatulate tip.

Variations

There is some variation in intensity of coloration of the brown hairs on the

scutum, scutellum and metanotum in both sexes. The greatest variation is, however,

in the length of the forelegs of females from the various localities and is correlated

with the spur lengths of the oil-secreting Diascia species at a particular locality

(Steiner & Whitehead 1991).

Host flower records

Nectar is not produced by oil-secreting hosi plants, so bees must obtain it from

other plants. Both males and females take nectar from the same source, and males

also patrol nectar plants in search of receptive females. Although it is the females that

mainly visit the oil-producing flowers, males have occasionally been collected patrol-

ling them, presumably in search of females. Rediviva pallidula females collect oil from

six species of Diascia (Scrophulariaceae), which include D. anastrepta Hilliard &

Burtt, D. barberae Hook f., D. cordata N. E. Brown, D. fecaniensis Hilliard & Burtt,

D. integerrima Benth., and D. stricta Hilliard & Burtt. They also collect oil from the

terrestrial orchids Pterygodium cooperi Rolfe and a species of Corycium. Nectar

plants are Lobelia preslii A. DC., Wahlenbergia cuspidata v. Brehm (Campanulaceae),

Phacocapnos pruinosis (E. Mey.) Bernh. (Fumariaceae), Geranium multisectum

N. E. Brown (Geraniaceae), and Stachys sessilis Guerke (Lamiaceae).

Distribution (Fig. 5)

Rediviva pallidula is a high-altitude bee occurring from 1 830 m to 2 720 m. In

the north-eastern Cape, it has been collected on Joubert’s Pass at Lady Grey, and at

Carlisle’s Hoek, Naudesnek and Ben Macdhui in the Rhodes district and at Bottlenek

Pass, Barkly East. In Natal, the species has been found just below the escarpment at

Garden Castle State Forest and at the top of Sani Pass. It is widely distributed in

Lesotho from Lebelonyane Pass in the south, east to the top of Sani Pass, north to

Oxbow and Moteng Pass, and west to Blue Mountain Pass. Manning & Brothers

(1986) reported R. politissima (= R. neliana) visiting several Diascia species on Sani

Pass, including D. barberae on the Sani Flats in Lesotho. We have examined this

material and found that they had a mixed sample, the specimens from Sani Flats being

the new species, R. pallidula. At lower altitudes, 1 800 m to 2 400 m, this species is

sympatric with R. neliana and shares the same oil and nectar plants.

TWO NEW SPECIES OF OIL-COLLECTING BEES 155

Rediviva saetigera sp. nov.

Figs 5-9

Diagnosis

Integument black with light to dark brown pubescence, except on episternum and

sternum where pubescence is white to cream. Female with long straight unbranched

dark hairs with bent or curled tips on clypeus, frons and vertex, parted on a line with

bases of antennae. Hairs on dorsal areas of front and middle tibiae and tarsae dark

brown to black. Front legs short with no elongation of tarsomeres 2 to 4, long dense

oil-collecting hairs on basitarsus and distal area of tibia. Glossa short, circular. This is

the only Rediviva known in the summer rainfall area with short front tarsomeres and

with oil-collecting hairs on the front basitarsus and tibia only.

Type locality

Transvaal, road to Lydenburg, 11,6 km south-west of Sabie, 2530BB, collected

on flowers of Bowkeria cymosa (Scrophulariaceae).

Etymology

Saetiger, Latin for ‘bearing bristles’, which refers to the pollen-collecting hairs on

the clypeus, frons and vertex of females.

Material examined

Type material. Holotype: SAM—HYMB0003, female, 11,6 km south-west of

Sabie, 2530BB, V. B. Whitehead, 30 Jan. 1988. Allotype: SAM—HYMB0004, male,

Graskop, God’s Window, 2430DD, V.B. Whitehead, 2 Mar. 1990. Paratypes

(46 99, 13 OC')—Cape Province: 1 9, Ocean View Farm, East London, 3228CA,

K. E. Steiner, 19 Jan. 1986; Natal, Kwazulu: 1 9, 52,6 km north-west of Eshowe,

2831CA, V. B. Whitehead, 13 Jan. 1989; 2 99, Nkandla Forest, 56,9 km north-west

of Eshowe, 2831CA, V. B. Whitehead, 13 Jan. 1989; 1 9, Nkandla Forest, 66,8 km

north-west of Eshowe, 2831CA, V.B. Whitehead, 14 Jan. 1989; 1 9, road to

Nkandla, km 38, 2831CA, K. E. Steiner, 13 Jan. 1989; 3 99, Road to Nkandla,

km 38, 2831CA, K.E. Steiner, 14 Jan. 1989; 2 99, road to Nkandla, km 38,

2831CA, K. E. Steiner, 1989; Transvaal: 2 99, Barberton, 25,5 km east of Road 38

on road to Havelock, 2531CB, V. B. Whitehead, 4 Mar. 1990; 1 9, Barberton,

9,3 km east off Road 38 on road to Havelock, 2531CB, K. E. Steiner, 4 Mar. 1990;

1 9, Barberton, 19,3 km, east off Road 38 on road to Havelock, 1 270 m, 2531CD,

V. B. Whitehead, 4 Mar. 1990; 1 9, Barberton, 31,8 km east off Road 38 on road to

Havelock, 2531CB, K. E. Steiner, 4 Mar. 1990; 5 99, Graskop, God’s Window,

1710 m, 2430DD, V. B. Whitehead, 2 Mar. 1990; 1 0’, Kaapsehoop, Berlin State

Forest, 2530DA, V. B. Whitehead, 31 Jan. 1988; 2 99, Long Tom Pass, 1 690 m,

V. B. Whitehead, 14 Feb. 1987; 1 9, Long Tom Pass, 1 590 m, V. B. Whitehead,

16 Feb. 1987; 1 C’, Pilgrim’s Rest, Vaalhoek Road, 1 280 m, 2530DD, V. B. White-

head, 2 Mar. 1990; 4 99, 0,5 km west of turn off to Pilgrim’s Rest on Sabie Road,

2530DD, K.E. Steiner, 4 Mar. 1986; 7 99, 10,7 km west of Sabie, 1 350m,

2530BA, K. E. Steiner, 3 Mar. 1986; 4 99, 11,6 km west of Sabie, 1 350 m, 2530BA,

K. E. Steiner, 5 Mar. 1986; 4 99, 11,6km west of Sabie, 1500 m, 2530BA,

156 ANNALS OF THE SOUTH AFRICAN MUSEUM

1mm

Fig. 6. Rediviva saetigera sp. nov. Female mouth-parts. A. Anterior view of labium. B. Distal part

of labium, posterior view to show subligular process and basiglossal sclerite. C. Maxilla, outer view.

D. Basistipital process. E. Inner view of maxilla to show galeal comb. F-G. Right mandible, outer

and inner views.

TWO NEW SPECIES OF OIL-COLLECTING BEES S7/

K. E. Steiner, 30 Jan. 1988; 1 9, 11,7 km west of Sabie, 1 500 m, 2530BA, K. E. Steiner,

15 Feb. 1987.

Other material. (68 29)—Natal, Kwazulu: 21 99, Nkandla, 2831CA; Trans-

vaal: 5 99, Barberton, 2531CD; 7 99, Graskop, 2430DD; 2 29, Kaapsehoop,

2530DA; 5 9 Q, Pilgrim’s Rest, 2430DD; 28 99, Sabie, 2530BA.

Description

Female

Measurement and ratios. Holotype, body 10,0 mm, forewing 9,5 mm; other

material in Table 3.

Integumental colour. Black, mouth-parts, scape and first flagellar segment black

with dark brown extremities, rest of flagellum dark brown, lighter on under-side; front

tarsae, middle and hind tibiae dark brown; tegulae light brown; wings lightly tinted

with brown, stigma and veins brown except R and M+Cu which are black.

Structure. Head wider than long, holotype 3,6:3,25 mm, other material

3,6 : 3,0 mm (n= 20) (Table 4); inner orbits of eyes converging above and below,

interocular distance greater than length, holotype 3,3:2,9 mm; interior distance

between the lateral ocelli greater than ocellocular distance; malar space narrow,

length one-eighth of width (holotype), other material in Table 4; mouth-parts

TABLE 3

Mean lengths (+SD) of forelegs, body, foreleg/body ratios and forewings of females

(n = 25) and males (n = 8) of R. saetigera from the eastern Transvaal and Nkandla,

Natal

Eastern Transvaal

Females Males

Length Range Length Range

(mm) (mm) (mm) (mm)

Foreleg 08,3+0,17 07,9-08,6 07,8+0,28 07,5-08,5

Body 10,9+0.70 09,7-12,5 09,0+0,35 08,5-09,7

Foreleg/body 0,76+0,04 0,66-0,81 0,87+0,28 0,80-0,90

Forewing 09,1+0,22 08,8-09,5 08,8+0,19 08,5-09,0

Nkandla

Females

Length Range

(mm) (mm)

Foreleg 08,2+0,12 08,0-08,4

Body 10,3+0,37 10,0-11,2

Foreleg/body 0,80+0,04 0,73-0,85

Forewing 08,9+0,26 08,3-09,2

158 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 4

Mean measurements (+SD) of head width (HW), head length (HL), interocular distance

(IOD), eye length (EL), and the length/width ratio of the malar space (MS-L/W) of

R. saetigera females (n = 20) and males (n = 8) from all areas.

HW(mm) HL(mm) JOD(mm) EL(mm) MS-L/W

Females 3,6+0,08 3,0+0,09 2,5+0,06 2,2+0,07 0,11+0,02

Range 3,4-3,8 D330 D050 1, M743) 0,07-0,20

Males 3,4+0,09 2,7+0,10 2,3+0,07 2,1+0,09 0,12+0,02

Range 39-90 4,522.8) 2,2-2,4 AAV =8) 0,08-0, 15

(Fig. 6A), glossa short, circular with short apical projection, paraglossae short not

extending beyond glossal extremity; labial palps extending beyond apex of glossa, seg-

ments 2 and 3 broader at apex; ligular arms occupying middle third of prementum;

premental fragmentum, proximal part of mentum and lorum sclerotized, posterior

ligular process and basiglossal sclerite as in Figure 6B; cardo slightly shorter than

stipes, stipes 3,5 times as long as wide, posterior margin with scattered long plumose

hairs (Fig. 6C); basistipital process short (Fig. 6D); maxillary palps extending beyond

galea, first two segments subcylindrical, following three broader distally; galea with

rounded apex, with hairs along apical and posterior margins, galeal comb large with

19 teeth (Fig. 6E); mandibles (Fig. 6F—G) strongly curved, stout preapical and apical

tooth, cap of rutellum occupying about one-third of mandible; labrum narrow, about

3,5 times as wide as long, rounded laterally with dense long hairs on anterior half,

hairs shortening posteriorly (Fig. 6H); antennae with first flagellar segment less than

half length of scape (excluding pedicellus), approximately as long as segments 2 plus

3, segments 4-9 subequal and as long as broad, segment 10 one-and-a-half times

longer than broad. Front legs short, not attenuate, sum of tarsomeres 2, 3 and 4 less

than one-quarter of leg length, similar in shape to tarsomeres of middle and hind legs

(Fig. 7A—B); hind legs with basitarsus trapezoidal in outline (Fig. 7C); tibial spurs

light brown, finely dentate. Wings (Fig. 8A) with second submarginal cell one-and-a-

half times broader than long, receiving first recurrent vein beyond middle (19:10),

third submarginal cell more than twice as broad as long, receiving second recurrent

vein beyond middle (22:17), basal vein more than twice as long as first abscissa of Rs,

joining Cu slightly distal to Cu-V; jugal lobe of hind wings slightly more than half

length of vanal lobe (34:62); ten hamuli. Propodeal triangle small, shiny, devoid of

hairs, distinguishable from adjacent propodeal surface which is coarsely punctate

reticulate, small area at base of triangle finely rugose. Apical margins of S2, S3 and S4

produced to a point in midline giving the margin a biconcave appearance; gradulus on

S2 bisinuate (Fig. 7D). Pygideal plate punctate, dull.

Sculpture. Head: clypeus and frons coarsely punctate, area between punctures

less than diameter of punctures, surface area between punctures finely reticulate,

small area adjacent to lateral ocelli, shiny, impunctate or with isolated punctures,

finely reticulate towards occipital ridge. Mesosoma: large shallow punctures on scutum

TWO NEW SPECIES OF OIL-COLLECTING BEES 159

2mm

Fig. 7. Rediviva saetigera sp. nov. Female. A. Foreleg, lateral view. B. Foreleg, ventral

view. C. Hind tibia and basitarsus (hairs removed). D. Ventral view of metasoma.

160 ANNALS OF THE SOUTH AFRICAN MUSEUM

ee —S) S\

—=-

omm

Fig. 8. Rediviva saetigera sp. nov. Female. A. Right wings. B. Head, to

show long unbranched pollen-collecting hairs.

with whole surface finely granulate; punctures coalescing on scutellum becoming scaly-

reticulate; punctures finer on metanotum with areas between punctures shiny; punc-

tures On mesepisternum elongate, tending to coalesce, surface finely granulate.

Metasoma: T1 shiny, sparsely punctured, T2 to T4 punctate, areas between punctures

finely granulate, apical margins impunctate; S2 to $4 more coarsely punctured, apical

margins impunctate, shiny.

TWO NEW SPECIES OF OIL-COLLECTING BEES 161

Vestiture. Head: long, unbranched stout black hairs between bases of antennae,

on supraclypeal area and down to just before anterior clypeal margin, shortening

anteriorly, parting at bases of antennae, hairs below antennal bases point forward and

above point backward (Fig. 8B), similar hairs on occiput between lateral ocelli and in

ocellocular area point forward, ends of hairs taper finely to a sinuate or curved point;

finely branched white hairs at base of scape; hairs on mandibles and labrum reddish-

brown, on under-side of head long, white and plumose. Mesosoma: margins of scutum

and scutellum densely covered in light brown hairs mingled with sparse black hairs,

disc of both areas with fine decumbent hairs; metanotum covered with light brown

hairs, paler on margins; propodeum (except triangle), episternum and sternum

densely covered with finely branched pale brown to white hairs; fore- and middle legs

with black hairs on basitarsus, tibia and anterior part of femur, dark brown ventrally,

hairs on front legs longer on both tibia and basitarsus, where unmodified hairs have an

under-storey of finely branched oil-collecting hairs; hind legs with dark hairs on base

of tibia and apex of basitarsus, sparse black and numerous long brown hairs protrud-

ing through mat of fine brown hairs on anterior face of tibia and basitarsus, posterior

surface covered in stout brown hairs, hairs on distal margin of basitarsus parted to

form a penicillum. Metasoma: decumbent light brown hairs on apical two thirds of T2,

T3 and T4, basal hairs paler, short on disc, longer laterally, isolated erect black hairs

basally on T4; TS and T6 with long black hairs on apical margins; long pale hairs on

apical margins of S2, S3 and S4, black on SS.

Male

Measurement and ratios. Allotype, body 9,0 mm, forewing 8,7 mm; other mat-

erial in Tables 3-4.

Integumental colour. Black, except extremities of mouth-parts, under-side of

antennal flagellum, tarsae, tibial apices, tibial spurs and tegulae, which are brown;

stigma and veins brown except R and M+Cu, which are black.

Structure. Head wider than long (3,4:2,7 mm); inner orbits as in female, distance

between eyes greater than length of eye (2,3:2,1), ocellocular distance equal to inner

distance between lateral ocelli; malar space short (L:W=3:28); first flagellar

segment of antenna short, less than half length of scape, shorter than successive flagel-

lar segments. Forelegs not modified, hind tibia slightly wider than basitarsus (15:12),

tibial spurs brown, finely serrated, basitibial plate small, twice as long as wide, basal

three-quarters covered with hairs. Wings with whole surface papillate, venation, jugal

and anal lobes as in female. Propodeal triangle as in female. Apical margins of S3, S4

and S5 pointed in middle giving margin a biconcave outline, concavities deepest on

S5; S6 (Fig. 9A) shiny on disc with scattered black hairs, apical and lateral lobes

small, lateral lobes with dense black hairs; apical lobes on S7 short with sparse hairs

on distal margins, lateral lobes large, crescent-shaped in lateral view, papillate, par-

ticularly on lower part (Fig. 9B—C); tip of S8, viewed from above, shiny, oval with a

single emargination (Fig. 9D—-F). Genitalia (Fig. 9G—H) with gonostyli narrow and

keeled below, with large translucent flange on upper surface basally; volsellae distinct

with teeth on opposing surfaces, opening lateroventrally.

Sculpture. As in female.

162 ANNALS OF THE SOUTH AFRICAN MUSEUM

1mm

Fig. 9. Rediviva saetigera sp. nov. Male genitalia and associated sterna. A. S6. B. S7, dorsal view

(left), ventral view (right). C. S7, side view. D. Dorsal and ventral view of S8. E. S8, lateral view.

F. Tip of S8, dorsal view. G. Genital capsule, ventral (left) dorsal (right). H. Genital capsule, lateral

view.

TWO NEW SPECIES OF OIL-COLLECTING BEES 163

Vestiture. Head: stout brown unbranched hairs on lower part of labrum, lighter

and shorter branched hairs on upper surface; mixture of long white and black

branched hairs on clypeus, supraclypeal area and frons; long branched black hairs on

vertex towards occipital ridge. Mesosoma: as in female. Metasoma: hair bands on

apical margins of T2, T3 and T4 brown (mixture of brown and black or all black on T4

in some specimens), black on TS, light brown on T6 and 17; basal hairs on T2 pale,

on base of T3 and T4 predominantly black with some pale hairs laterally, on TS black;

hairs on apical margins of S2, S3 and S4 long, paler than on tergum, on S5 black, hairs

longer and more dense laterally. Forelegs with light brown hairs on tarsomeres 2-5,

under-side of basitarsus and distal part of tibia; hairs on dorsal surface of basitarsus,

dorsal and posterior areas of tibia black; mixture of long black and long white

plumose hairs on the ventral side of femora; middle legs similar to forelegs but with

some dark hairs on tarsi; hind legs with more black hairs on tarsi and tibiae; hairs on

ventral surface of femora, tibiae and basitarsus light brown; tibial spurs light brown.

Variations

The brown and reddish-brown hairs on females of the eastern Transvaal popu-

lations are richer in colour than those from Natal and the eastern Cape. Males from

the eastern Transvaal also have a rich reddish-brown on the mesosomal dorsum and,

although no males have been collected from the coastal regions of Kwazulu or the

eastern Cape, it is probable that they will also have the dull colour of the females.

Males collected in the eastern Transvaal also have some variation in the amount of

brown hair in the band on the posterior margin of T4, which can be mostly brown to

all black.

Host flower records

Only two host plants have been recorded for this species: one, Bowkeria cymosa,

is a shrubby tree that occurs in the cooler higher rainfall areas of the eastern Trans-

vaal, in forest edge, stream and roadside situations. The other, Anastrabe integerrima,

is a shrub to medium-sized tree that occurs primarily in warmer coastal areas; it too

grows along forest edges or streams. Female R. saetigera bees collect pollen and oil

from both host plants. Males have been found to patrol the oil-producing flowers of

B. cymosa early in the season, presumably in search of females. No bees of this

species have as yet been found visiting nectar plants.

Distribution (Fig. 5)

Rediviva saetigera is associated with B. cymosa on the eastern Transvaal escarp-

ment from Pilgrim’s Rest to Kaapsehoop, and further south from the hills of

Barberton to the Swaziland border. Large populations occur in the Eshowe—Nkandla

area of Kwazulu, and one specimen has been collected near the eastern Cape coast

between Komga and Kei Mouth. The host plant at these sites, Anastrabe integerrima,

occurs in the coastal forests of Transkei and southern Natal but, despite searching in

several locations in these areas, we have been unable to find additional localities for

R. saetigera.

164 ANNALS OF THE SOUTH AFRICAN MUSEUM

ACKNOWLEDGEMENTS

We thank Dr Nigel Fergusson and Mr George Else of the Natural History

Museum, London, for use of facilities and their assistance in locating the material

described by Cockerell, and Professor Denis Brothers, University of Natal, Pieter-

maritzburg, for the loan of material. The National Parks Board and the Natal Parks

Board are thanked for permission to work in their respective areas and, in particular,

Messrs E. Fourie of the Mountain Zebra National Park, G. Groenewald of the

Golden Gate Highlands National Park, H. Braack of the Karoo National Park, and

K. Riggien of the Department of Forestry, for assistance with transport and

accommodation. The constructive comments of two referees are much appreciated.

REFERENCES

Eapy, R. D. 1968. Some illustrations of microsculpture in Hymenoptera. Proceedings of the Royal

Entomological Society (series A) 43 (4—6): 66-72.

MANNING, J. C. & Brotuers, D. J. 1986. Floral relations of four species of Rediviva in Natal

(Hymenoptera: Apoidea: Melittidae). Journal of the Entomological Society of Southern Africa 49

(1): 107-114.

MIcHENER, C. D. 1981. Classification of the bee family Melittidae with a review of the species of

Meganomiinae. Contributions of the American Entomological Institute 18 (3): i-1i, 1-135.

Roserts, R. B. & Brooks, R. W. 1987. Agapostemoninae bees of Mesoamerica (Hymenoptera:

Halictidae). University of Kansas Science Bulletin 53 (7): 357-392.

STEINER, K. E. & WHITEHEAD, V. B. 1988. Association between oil-producing flowers and oil-

collecting bees in the Drakensberg of southern Africa. Monographs in Systematic Botany from

the Missouri Botanical Garden 25: 259-277.

STEINER, K. E. & WHITEHEAD, V. B. 1990. Pollinator adaption to oil-secreting flowers—Rediviva

and Diascia. Evolution 44 (6): 1701-1707.

STEINER K. E. & WHITEHEAD, V. B. 1991. Oil flowers and oil bees: further evidence for pollinator

adaption. Evolution 45 (6): 1493-1501.

VOGEL, S. & MICHENER, C. D. 1985. Long bee legs and oil-producing flower spurs, and a new Redi-

viva (Hymenoptera, Melittidae; Scrophulariaceae). Journal of the Kansas Entomological Society

58: 359-364.

WHITEHEAD, V. B., SCHELPE, E. A. C. L. E. & ANTHONY, N. C. 1984. The bee Rediviva longimanus

Michener (Apoiodea: Melittidae), collecting pollen and oil from Diascia longicornis (Thunb.)

Druce (Scrophulariaceae). South African Journal of Science 80 (6): 286.

WHITEHEAD, V. B. & STEINER, K. E. 1985. Oil-collecting bees in South Africa. African Wildlife 39

(4): 144-147.

6. SYSTEMATIC papers must conform to the International 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

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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

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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.

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(a) The Figures, Maps and Tables of the paper when referred to in the text

Creme tune ieune depictime Cy namacolus)...(... .1n C. namacolus (Fig. 10)).. 7

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Punctuation should be loose, omitting all not strictly necessary

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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.

V. B. WHITEHEAD

K. E. STEINER

TWO NEW SPECIES OF OIL-COLLECTING BEES

OF THE GENUS REDIVIVA FROM THE

SUMMER RAINFALL REGION OF SOUTH AFRICA

(HYMENOPTERA, APOIDEA, MELITTIDAE)

or UNE 102 PART 5 FEBRUARY 1993 ISSN 0303-2515

SONIAR

I SONTAR

OF THE SOUTH AFRICAN

~ MUSEUM

CAPE TOWN

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(a) Author’s name and year of publication given in text, e.g.:

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Examples (note capitalization and punctuation)

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

FISCHER, P. H. 1948. Données sur la résistance et de la vitalité des mollusques. Journal de conchyliologie 88 (3): 100-140.

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

expérimentale et générale 74 (33): 627-634.

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

Magazine of Natural History (13) 2 (17): 309-320.

Koun, A. J. 1960b. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bulletin of

the Bingham Oceanographic Collection, Yale University 17 (4): 1-51.

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

pologische Ergebnisse einer Forschungsreise im westlichen und zentralen Stid-Afrika ausgefiihrt in den Jahren

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(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 102 #£Band

February 1993 Februarie

Part 5 Deel

HUMAN SUBJECTS AS MUSEUM OBJECTS.

A PROJECT TO MAKE LIFE-CASTS OF “BUSHMEN’

AND ‘HOTTENTOTS’, 1907-1924

PATRICIA DAVISON

Cape Town Kaapstad

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D1816

HUMAN SUBJECTS AS MUSEUM OBJECTS.

A PROJECT TO MAKE LIFE-CASTS OF ‘BUSHMEN’

AND ‘HOTTENTOTS’, 1907-1924

PATRICIA DAVISON

African Studies & Anthropology, South African Museum, Cape Town

(With 8 figures and 1 table)

[MS accepted 18 August 1992]

ABSTRACT

In 1906 the Director of the South African Museum initiated a project to make life-casts of ‘aborigi-

nals of the Bush and Hottentot Races’, believed at the time to be near extinction. The project was thus

regarded as having anthropological importance. Between 1907 and 1924 over 60 casts were made, and

although intended primarily as a scientific collection, many of the casts were placed on exhibition as

examples of a primitive race. This paper demonstrates how museum practice reduced the people who were

cast to specimens for public viewing. In this process human subjects became museum objects.

CONTENTS

PAGE

ITNTROGINCTION, sho. ae nolo a 6 a0 1B 0 Obs Wha ORL OF ORIG: Cie ON gen en eee ee 165

BnStonicalab ack enouiin iy wae news my aN cheesy ales Che cadly as aid awd: epee ai 165

sihtercastinieaproject, LOOT 2A ee ee ee ES ee lee ea eens 168

J CGM OCU. UNE CASS 's Bs ob So caso old 58 abe UE ote Smee re Go enP Esai ae navn rep 178

DISCUSSION o'o 0168'S bbe ee dad ory a Che Oe trons OR aR neste nee a ge an 181

ANC THOVWMEURSMNCAIS iS ole olege log bie 5.5 COR Cee en ene teres et eon ee ne 182

INCHETENCE SHE MME E IE meter rear eee ier iste, eR Attia ech a le dieu tale ea) eos 182

INTRODUCTION

During the past decade, anthropologists have become critically aware of the meth-

odological and epistemological assumptions of their discipline (Fabian 1983; Clifford &

Marcus 1986; Clifford 1988; Kuper 1988). If, as is now widely acknowledged, the disci-

pline as a whole was originally premised on treating people of other cultures as objects

of study, anthropology as practised in museums provides one of the clearest examples of

this process. The casting project undertaken at the South African Museum (SAM)

between 1907 and 1924 can be regarded as a tangible manifestation of a general concep-

tual position that underpinned the emerging discipline of academic anthropology. By

adopting a historical perspective and focusing attention on a collection of life-casts of

subjects deemed to be pure ‘Bushmen’ and ‘Hottentots’, I hope to elucidate the way in

which museum practice operated to reify these people in terms of race, to objectify them

as physical specimens, and to perpetuate unproven anthropological notions regarding

racial primitiveness.

Although it could be argued that this paper further dehumanizes the people who were

cast, my intention has been to situate the casting project historically and to increase

awareness of an inherent problem in anthropological practice, past and present.

165

Ann S. Afr. Mus. 102 (5), 1993: 165-183, 8 figs, 1 table.

166 ANNALS OF THE SOUTH AFRICAN MUSEUM

HISTORICAL BACKGROUND

Since its inception in 1825, the South African Museum has been associated with an

anthropological interest in the people widely known as ‘Bushmen’ and ‘Hottentots’.

(Although San and Khoikhoi are the preferred current terms, the use of the former terms

in this paper reflects historical usage.) Dr Andrew Smith, founder of the SAM, published

an early paper on the origin and history of the Bushmen and Hottentots (Smith 1830), and

on his 1834—1836 expedition to the interior he collected a range of ethnographic artefacts,

including ‘9 Sets of Bosjesman Bows and Arrows’ (Lye 1975). In the Museum accession

registers that date back to 1855, when the Museum was reconstituted under the Director-

ship of Edgar Layard, Khoikhoi and San artefacts appear intermittently in the Miscella-

neous Collections, categorized under various sub-headings such as ‘Work of Uncivilized

Races’, ‘Native implements’ or ‘Work of native races’. An early interest in displaying

modelled figures is suggested in the diary of Lieutenant T. Duthie, who recorded in 1832

that he and Dr Murray of the South African Institution had gone to the Museum ‘to see

the [modelled] Hottentot woman of Smith’s’ (Kirby 1965: 116). Although no further

reference to this exhibit has been found, it attests an early concern for a subject that was

later to be pursued more scientifically at the Museum.

As an army surgeon, Andrew Smith had been trained in anatomy in Edinburgh,

graduating in 1819, a few years after Robert Knox, who later became a vociferous advo-

cate of polygenism. During the first half of the nineteenth century, the debate between

polygenists and monogenists was the most contentious issue within anthropological dis-

course (Harris 1969). Polygenists, who generally also opposed the abolition of slavery,

were emboldened by the taxonomic work of Linnaeus in their attempts to differentiate the

human species (Haddon 1934). Although there is no evidence to suggest that Smith was

a polygenist (Kirby 1965), he was doubtless a participant in the controversial debates on

racial and cultural evolution. It is of interest that he presented the cranium of a ‘notorious

Bushman murderer’ to the Anatomical Museum of the Royal Army Medical Corps in

Chatham where, after returning to England, he served as Principal Medical Officer from

1837 to 1845 (Morris 1987: 14). In being concerned with questions of racial origins and

classification, Smith was a man of his time. He was a meticulous scientist and recorder

of ethnographic data but he did not dispute the widely held nineteenth-century belief that

both Hottentots and Bushmen were among the low orders of humanity.

General explanations for this conviction included the influence of climatic factors, a

presumed low position of Bushmen and Hottentots in the Great Chain of Being, and the

possibility of their being biologically different from other races. The latter polygenist

view gave rise to a scientific discourse on race that attempted to establish racial distinc-

tiveness on morphological criteria. Among these the cranial index (length : breadth ratio

of the skull) was believed to be of taxonomic significance in classifying racial types, and

was also thought to be linked to intellectual capacity (Stocking 1987). Phrenologists used

skull dimensions as evidence to support theories on unequal intellectual capability among

different races, and there was a widespread popular belief in the correlation of race and

physiognomic features with moral disposition. The amassing of morphological data on

people of different races was considered an essential prerequisite to solving current

anthropological problems, both before and after the publication of Darwin’s work on the

origin of species (Duckworth 1904).

HUMAN SUBJECTS AS MUSEUM OBJECTS 167

During the nineteenth and early twentieth centuries, hundreds of skulls of people

believed to be Bushmen found their way into European collections and were used in

morphological studies.

These osteological remains were considered to be part of the native fauna of distant

lands and no natural history collection could be considered complete unless it con-

tained a representative quantity of human skulls (Morris 1987: 12).

That skulls of Bushmen were regarded as faunal collections is telling, if not surpris-

ing, considering that an evolutionary paradigm taken from natural history had a strong

influence on the emerging ethnological sciences. In 1847, when ethnology was included

in the British Association for the Advancement of Science, it was grouped with “Zoology

and Botany’ (Altick 1978). Historical time-depth and the principles of stratigraphy, dem-

onstrated scientifically in the earth sciences, were increasingly drawn on to explain

human variation. In order to treat people as specimens, as objects of study, it was neces-

sary to distance them in concept both temporally and spatially. Accordingly, Bushmen and

other ‘primitive’ people could be treated as living fossils—specimens to be appropri-

ated for the advancement of science.

From 1859 onwards Darwin’s theory of evolution provided, by analogy, a range of

biological explanations for human cultural differences. The language of biology was

transferred as a metaphor to the discourse of the social sciences. Social evolutionists,

however, misused Darwin’s theory of natural selection in the promotion of selective

breeding or eugenics, and they also misinterpreted his concept of time by linking it to the

idea of cultural progress (Ingold 1986). Throughout the nineteenth and early twentieth

centuries, racial typology and the relationship between race and culture remained subjects

of intense public interest and academic investigation. As the authority of the scientific

disciplines of biology, natural history and ethnology grew, the ‘primitive’ or ‘lower’ races

were increasingly regarded as anthropological specimens.

By the mid-nineteenth century, the Enlightenment idea of the ‘noble savage’ had

been eclipsed by confidence in European superiority, a conviction that was confirmed by

successive exhibitions in London of aboriginal inhabitants of Africa and America. One

particular exhibition that had a notable impact on the popular image of the Bushmen was

held in 1847 at the Egyptian Hall in Piccadilly. To give an aura of scientific respectability

to the venture, it was preceded by a lecture delivered at the Exeter Hall by the anatomist,

Robert Knox. The lecture was publicized as addressing, among other things, ‘the great

question of race’ (Altick 1978: 280). Appealing to a more popular audience, the exhibi-

tion poster advertised: ‘Bosjesmans . . . The most singular specimens of that decreasing

race of human beings . . . that from their wild habits could never before be induced to

Visit a place of civilization.” The Times described the Bushmen as ‘little above the

monkey tribe, and scarcely better than the mere brutes of the field’ (cited in Altick 1978:

281). Not only were the Bushmen objects of curiosity, they were also a source of gratifi-

cation for the Victorian public whose self-esteem was enhanced relative to the spectacle

they observed so avidly.

In the context of southern African settler society, a distorted stereotype of the indige-

nous population served a more insidious purpose than simply affirming the superiority of

the settlers. It justified the violence and dispossession inflicted on the Khoikhoi and San

people (Guenther 1980). Throughout southern Africa, the resistance of the aboriginal

168 ANNALS OF THE SOUTH AFRICAN MUSEUM

population was eventually quelled and, by the second half of the nineteenth century, their

numbers had been greatly reduced. Those who survived in the Cape Colony were no

longer living as hunter-gatherers but as labourers and squatters on farms. By this time

they had ceased to be a threat to the settlers, but remnant groups were becoming of

increasing interest to philologists and ethnologists as living examples of a low order of

the human species.

Scientists working in South Africa aligned themselves with a cosmopolitan scientific

fraternity and were strongly influenced by the ideas of overseas specialists. This was

apparent in 1905 when the British and South African Associations for the Advancement

of Science held a joint meeting in South Africa. A. C. Haddon, president of the Anthro-

pological section, concluded his address by stressing the importance of investigating the

Bushmen and Hottentots, who represented ‘very primitive varieties of mankind’, and who

were ‘rapidly diminishing’ in number (Haddon 1906: 525). At the same meeting Professor

F. von Luschan, Director of the Museum fiir V6lkerkunde in Berlin, recommended that

casts from the living subject should be made of the few extant full-blooded aboriginals

of the Bush and Hottentot races.

THE CASTING PROJECT, 1907-1924

With the impetus of international scientific interest in the question of racial origins,

Dr Louis Péringuey, who had become Director of the South African Museum in 1906,

initiated a project that was aimed at making an accurate physical record of members of

the few remaining groups of ‘pure-bred’ Bushmen and Hottentots.

Hoping to enlist the official support of the Cape Government in locating suitable

subjects, Péringuey wrote to the Under Colonial Secretary of the Cape as follows in 1907:

Modelling from life aboriginals of the Bush and Hottentot Races

Sir,

Owing to the rapid disappearance by reasons which I need not mention here, of the

pure specimens of the Hottentot and Bushman races the Trustees of the Museum are

endeavouring to obtain models from the living flesh which would enable the exact

physical reproduction of the survivors of these nearly extinguished races.

I have of late endeavoured to locate pure types of both sexes, but not as successfully

as I would have expected. Moreover some of these live at distances so great that the

expenditure of travelling to such parts greatly militates against the successful result

of my endeavours.

But the Cape Government would greatly assist the Trustees of the Museum in secur-

ing the last vestiges of these people,

1. by asking the Civil Commissioners to inquire of their respective jurisdictions as

to the presence there of true Bushmen and Bushwomen, Nama Hottentots and

Korannas and to report to you.

2. by obtaining similar informations from the Superintendent of Prisons, or jails in

the country.

3. by authorizing us in the case of men and women in jail to have the casts and

necessary photographs and measurements taken by experts. The process is not a long

one; it is very simple and absolutely painless.

HUMAN SUBJECTS AS MUSEUM OBJECTS 169

The importance of securing these physical reproductions, while we still can do it, is

so great that I doubt not that you will grant my respectful request for the information

and instructions mentioned herein.

(SAM letterbook, 31 July 1907)

The project received the support of the Colonial Office, and assistance was duly

requested from the Secretary for Native Affairs, as well as Convict Stations and Magis-

trates in the northern districts of the Colony and in the Bechuanaland Protectorate. In most

cases those responding to Péringuey’s request showed willingness to assist but, not sur-

prisingly, expressed some difficulty in locating ‘pure aboriginals’. The reply of the

Superintendent of the Convict Station in Kimberley to the circular sent out by the Under

Colonial Secretary exemplifies this uncertainty:

Sir,

. .. | have the honour to state that we have a number of Bushmen, Hottentots, and

Koranna convicts stationed here. Amongst them are fairly good specimens of their

race, but I might point out that it is most difficult nowadays to find a pure-blooded

specimen.

. . might I ask whether the Director of the S. A. Museum is not in a position to

suggest someone who could confer with me at this Station any Sunday morning

between 9.15 and 10.15 a.m. It could then be decided whether the types we have are

pure enough for their models to be taken....

(My emphasis; SAM letterbook, 19 September 1907)

The reference to ‘a pure-blooded specimen’ draws attention to the fact that the

project was premised on the notion of racial purity, and furthermore that, for scientific

purposes, people could be reduced or dehumanized to objects of study, to ‘specimens of

their race’. Péringuey (1911) was explicit about the most notable physical characteristics

of ‘pure bred’ members of the Bush race—women would have steatopygia and elongated

labia minora, and men would have semi-erect penises. He noted that he had been

informed by ‘old Colonists’ that ‘the purity of the race was denoted by the angle at which

the penis stood normally’ (Péringuey 1911: 201). In his instructions to the Museum

modeller (see p. 175), Péringuey also stressed the importance of skin colour and facial

features. Subjects to be cast were measured according to predetermined morphological

criteria, and photographed from a number of different angles to show their height as well

as the above features.

Considering that it is a general practice for convicts to be de-personalized and made

to conform to disciplinary procedures, it can be assumed that suitable ‘specimens’ in

convict stations were not given the right to refuse being photographed naked, measured

in minute detail and cast ‘from the living flesh’. However, it would seem that those who

were not in prison required some persuasion. Dorothea Bleek commented in a letter

written to Péringuey from Prieska: ‘It is exceedingly difficult to get photos of the natives

without clothes on. Perhaps your modeller will be more fortunate than we were’ (SAM

correspondence, 6 August 1911). In practice the project depended on the unequal rela-

tions between the people who were cast and those who captured their likeness in plaster

and on film. The reduction of the human subject to specimen for observation is clearly

revealed in photographs that were taken at the time (Fig. 1A, B).

170 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 1. Photographic studies of an anonymous Khoikhoi woman, Caledon, c. 1912. A. Clothed in

everyday attire. B. Unclothed for scrutiny by Museum scientists. (South African Museum Photographic

Collection.)

Although very little is known about the interaction between Museum modeller and

the people who were cast, there is some evidence of reticence on the part of proposed

subjects. An example of an unwilling subject is recorded in a letter from Péringuey to the

Secretary for Native Affairs in the Transvaal (SAM letterbook, 14 September 1907). A

Bushwoman, whose physical attributes had been examined previously by members of the

Anthropological Committee of the Association for the Advancement of Science, refused

to be cast and would not be persuaded to reconsider her decision. In the face of this

disappointment, Péringuey was hoping to enlist official support in persuading the woman

to submit to being measured and photographed ‘under different aspects’ so that an accu-

rate model of her could be made. There is no record of this having been accomplished.

The woman’s refusal to comply seems especially poignant considering that Péringuey had

relied on the power of officialdom to further his project. I am not suggesting that physical

coercion was used but that bureaucratic power would have been exerted. Elsewhere

Péringuey states explicitly that Bushmen were likely to acquiesce to being modelled ‘if

HUMAN SUBJECTS AS MUSEUM OBJECTS 171

Fig. 2. James Drury (left) working on the casts in the Museum studio. Dr Péringuey is seated in the centre.

(South African Museum Photographic Collection.)

the request comes from someone in authority’ (Péringuey to Colonial Office, SAM

letterbook, 25 March 1908). Repeated appeals through the Colonial Secretary

for the co-operation of Magistrates, Constables, Chiefs and Missionaries were justi-

_ fied in terms of the perceived urgency and scientific importance of the casting project.

Thus the authority of science was linked to that of the state to give the project official

status and credibility.

Between 1907 and 1924 the Museum modeller, James Drury, photographed, meas-

ured and made field-moulds of ‘thoroughbred’ San and Khoikhoi wherever suitable sub-

jects were to be found. Plaster casts from these moulds were made and painted in the

Museum studio (Fig. 2). Drury cast people living in Prieska, Carnarvon and other villages

in the Northern Cape; Grootfontein and Sandfontein in South West Africa; Kanye in the

Bechuanaland Protectorate; and the Lake Chrissie area of the eastern Transvaal. He also

cast convicts in Cape Town, Kimberley, Windhoek and Gaborone, if they conformed to

the assumed ‘pure’ physical type. During this time Drury made body-casts of 68 people

(including 14 convicts) whom he also photographed and measured in anatomical detail.

The registered casts are listed in Table 1. In addition to the figures that Drury cast, he

made field-moulds of at least another 20 people. These moulds remained unused for some

60 years. In the 1980s casts were eventually made from the moulds (SAM—AP6186 to

SAM-AP6205).

On his expedition to Kanye in 1908, Drury was given specific instructions by

Péringuey. The memorandum is cited in full as it gives insight into the casting project as

a whole:

172

ANNALS OF THE SOUTH AFRICAN MUSEUM

For Drury—Memorandum about the Modelling

I would like to have first, a group of five or six, men, women and children photo-

graphed in the position they naturally assume, either in sitting down, or as if they

were on the march: the man carrying his few arms and chattels; the woman carrying

what they generally carry, the youngsters probably carry nothing.

But apart from these two groups, and I think that Mr Harvey will be from his

knowledge of statuary quite able to make them assume positions that will not make

the models appear too stiff, you may have to take single people somewhat like the

figures we have. Try also to place them in such a position that would not prove too

fatiguing, in order to avoid also stiffness in the reproduction.

Do not chose [sic] the two decrepit specimens. But I would far prefer however to

have those with all the wrinkles of the body, especially the belly, than to have them

as well fed as our previous specimens.

Pay special attention to the hairs [sic] in your note of the specimens, of the colour

or expression of the eye, of the shape of the ear, and above all copy the colour of the

skin, and verify your slab a couple of days after you have painted it in order to

make quite sure of the genuine colour.

Of course photographs of the full face, the quarter and side views will be taken of

each. If you run short either send for more at Mafeking, or reduce the three views

of the face to two.

You must not forget however that we are not likely for some time to come,to have

such an opportunity and any photograph, provided it is a good one will prove of

great value to us.

Men are of course desirable; women still more so. You will be very careful to take

all their peculiarities, including the ‘apron’. A special moulding of the same to be

added to the statue is very much wanted. You will endeavour to find out more or

less the age of the young ones. Could you take a woman with her little one on the

back, wraps and all, it would indeed look very natural.

But to resume avoid any stiffness in the attitude you will take the models in.

If for reasons unavoidable you were leaving Kanye without finishing the models in

hand, it is understood that Harvey will remain a few days longer. You would then

take down with you such parts as have already been taken, Harvey bringing down

the rest.

As to the remuneration to the Chief you are authorized to give him from 5 to

10 pounds provided he gives you all facility for taking the casts. You would however

not make him the present until you have ascertained from some person in authority

or the Missionary whether you should give him the full sum or the other.

As to the Bush people you will probably get at the stores the shirts and petticoats

suitable for them. The knives you have. If need be you might get more on the spot.

As to the Graphophone. Read carefully the instructions. If you have no time to attend

to that part of the undertaking, ask the Missionary there to be kind enough to do so,

while you are proceeding with the modelling. If need be I could send more cylinders.

HUMAN SUBJECTS AS MUSEUM OBJECTS aS

Endeavour to buy the garments of these Bush people in order to clothe the

reproductions with if you can, provided their garments or arms are not man-

chester or [? Birmingham] goods.

If any native curio other than Bush was procurable you may buy if not dear.

You will let me know of your arrival and how you are getting on. It may be that I

take a run up that way, but I am afraid that Finances will not permit.

I should not like you to lose some useful opportunities for the sake of a few shillings,

but I am compelled to recommend a careful handling of the fifty pounds you take to

cover expenses.

I am enclosing all correspondences with the Resident Commissioner’s office. In case

of need you should apply to him.

(Péringuey to Drury, SAM letterbook, 1908: 718-720)

This memorandum clearly conveys some of the assumptions and emphases of the

project. The stress on accurate recording of physical attributes, notably exact skin colour

(skin pigmentation remains one of the most manifest signifiers of race), facial features

and genital characteristics underlines the fact that questions about race motivated the

project. Indeed the project was regarded as adding evidence to the debate on the classifi-

cation of the racial stocks of Africa which, as Dubow (1989: 5) has commented, was

accompanied by ‘hairsplitting distinctions with respect to sub-races and the relationship

between races’. The frequent reference to ‘specimens’ confirms that the human subjects

referred to by Péringuey had been objectivized conceptually before they were literally

objectivized as casts. Indeed, I contend that this was a necessary precondition for the

project.

Genital features were considered to be of special interest as they were differentiating

characters in taxonomic classification (Gilman 1986). Despite the fact that polygenism

was easily disproved by ample evidence of inter-racial breeding and that support for

polygenist arguments declined in the second half of the nineteenth century, interest in the

genital features of Bush and Hottentot people persisted. Female genital morphology was

of particular interest and, in accordance with Péringuey’s instructions, Drury made special

mouldings of female genitalia (SAM—AP6205 to SAM—AP6209). During the nineteenth

century, both popular and academic attention had been drawn to the physical charac-

teristics of Hottentot women, following the London exhibition in 1810 of Saartjie Baart-

man (‘The Hottentot Venus’), and the dissection of her genitalia by Cuvier, after her death

in Paris in 1815 (Kirby 1954; Altick 1978; Gould 1982). In the early decades of the

twentieth century, the significance of the so-called ‘tablier Hottentot’ was still the subject

of speculation, and continued to attract scientific interest (Péringuey 1911; Drury &

Drennan 1926).

The memorandum also reveals an emphasis on ‘traditional’ artefacts that was char-

acteristic of museum collecting-practice at the time, and has continued to the present.

Although purchased shirts, petticoats and knives were suggested as gifts for the people

who were cast, Drury was explicitly instructed not to acquire garments of trade-cloth for

the Museum. No interest was shown in documenting the actual material culture of the

people to be cast. If garments were to be acquired for the Museum, they should be of the

type that would allow casts of ‘pure’ racial types to be dressed in specimens of true

174 ANNALS OF THE SOUTH AFRICAN MUSEUM

‘traditional’ attire. Both of these notions were idealized constructs and yet, ironically,

Péringuey was concerned that the casts should appear natural. This was a quest for

simulated naturalness to be achieved by completely artificial means.

That people having features assumed in advance to be typically Bushman were

selectively cast is evident from much of Péringuey’s correspondence. For example, when

asked by the Resident Magistrate in Kanye which kind of Bushmen from Ngamiland he

wished to cast, since they varied in size, being ‘diminutive, medium or large’, Péringuey

replied, “By nature the real Bushman is small . . . It would thus seem that if the Masarwa

Bush are large, one can assume that they are not of pure blood, and the diminutive kind

would therefore be preferable’ (SAM letterbook, 20 June 1915). Thus, if the casts showed

features presumed to be ‘typically Bushman’, it was because those people who were cast

had been pre-selected to show such features—the results of the project simply provided a

tautologous reinforcement of an existing idea. Those people who did not conform to a

preconceived stereotype were excluded from the project or regarded as ‘half-breeds’.

For museum purposes the casts were registered as specimens in the physical anthro-

pology collections (see Table 1), together with the human skeletal collections. Although

some personal details of Drury’s subjects were kept on record, the plaster-casts were

essentially museum objects, ‘specimens’ identified primarily by registration number, and

by those details recorded in the accession register—racial type, sex and locality. It is

instructive to consider briefly some of the information that was omitted from the acces-

sion records. Perhaps the most telling omission is the subject’s personal name. For exam-

ple, in 1911 at Prieska, three members of the Bosman family—Stuurman, Piet and Klein

Piet—were cast, as were Anna Zwartbooi, Willem Toonies and Willem Jooi, among

others, but from the accession records it is not possible to identify the casts of these

people. Although in many cases Drury noted the names of the individuals whom he cast,

they were never included in the accession register, which lists the casts only as numbered

specimens. The omission of personal names is a clear illustration of the dehumanizing

nature of the project.

The emphasis on physical type also precluded any serious attention being given to

social context. In most cases living conditions, social setting and material culture were

not recorded for museum purposes, but the records on some of Drury’s subjects who were

cast in prison give an indication of their clothing and other personal effects. The listed

possessions of 17-year-old Augeniet Booysen (SAM—AP3897, identified as a ‘half-

breed’), cast in 1908 at the House of Correction in Cape Town, where she was serving

six months hard-labour for stock-theft, include a skirt, blouse, two petticoats, chemise,

three handkerchiefs, bodice, lace collar, shoes and a ring. As she is represented in the

Museum, however, Augeniet is without any personalizing cultural attributes. This accords

with the scientific interest in the casts primarily as examples of a physical type. Ethno-

graphic details of dress and local setting were not relevant to the project and were

deliberately overlooked in the way the casts were presented to the public, even though

the Museum had access to photographs showing the socio-cultural environment of

Drury’s subjects.

Photographs taken by Dorothea Bleek at Prieska in 1911 show aspects of the living

conditions that pertained when Drury cast a remnant group of /Xam Bushmen (Figs 3-5).

Although these people were then living in a semi-permanent squatter camp on the out-

skirts of Prieska village, their racial ‘purity’ had been affirmed on ‘philological grounds’,

HUMAN SUBJECTS AS MUSEUM OBJECTS LIS

Fig. 3. Group of /Xam Bushmen, some of whom were cast by Drury, Prieska, 1911. (Photograph: D. Bleek,

South African Museum Photographic Collection.)

Fig. 4. /Xam family group outside their home. The two adult women were Dorothea Bleek’s informants

and were cast by Drury, Prieska, 1911. (Photograph: D. Bleek, South African Museum Photographic

Collection.)

ANNALS OF THE SOUTH AFRICAN MUSEUM

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Body casts made by James Drury, 1907-1924, registered with the Physical Anthropology collections of the South African Museum.

_ eee

SAM

—————— eee

NO. SEX GROUP

AP3391 6 Sarwa

AP3392 6 Sarwa

AP3393o Sarwa

AP3394 6 Sarwa

AP3395 ¢ River Bushman

AP3396 @ Sarwa

AP3397 2. Sarwa

AP3398 'Kung

AP3399 9 'Kung

AP3400 6 'Kung

AP3401 ¢ "Bushman"

AP3402 3 ‘Bushman’

AP3403o Nama

AP3404 ‘Bushman’

AP3405 3 "Bushman'

AP3406 «6 "Bushman'

AP3407 3 "Bushman'

AP3408 ¢ "Bushman"

AP3409 ¢ "Bushman’

AP3410 9¢ "Bushman’

AP3411 ¢ "Bushman"

AP3412 9¢ "Bushman'

AP3413 9 "Bushman'

AP3414 9 "Bushman"

AP3415 3 Damara/Topnaar

AP3416 3 ‘Hottentot'

AP3417 9 "Hottentot'

AP3878 9° Topnaar

AP3879 9 /Xam

AP3880 36 /Xam

AP3881 3 /Xam

AP3882 6 /Xam

AP3883 89? /Xam

AP3884 2 /Xam

AP3885 c-) /Xam

v v v Vv Vv y ad

AP3886 2 /Xam

AP3887 9° /Xam

AP3888 3 /Xam

AP3889 =o /Xam

AP3890 «¢ /Xam

AP3891 3 /Xam

AP3892 6

AP3893o Nama

AP3894 /Xam

AP3895 2 /Xam

AP3896 9 /Xam mother, Cape coloured

father

AP3897 9 "Hottentot' mother, Xhosa

father

AP3898 9 ? Hottentot or Bushman

mother, Hottentot father

AP3899 6 Korana

AP3900 «3 Kgalagadi

AP3901 2 Kgalagadi

AP3902 Kgalagadi

AP3903 ¢ "Makua‘

AP3904 9 "Hottentot'

AP3905 6 Nama

AP3906 «¢ Bondelswart

AP3907 3 Korana

AP3908 3 Cape Hottentot

AP3909 3 Griqua

AP3910 3 Topnaar

AP3911 3 Kgalagadi

AP3912 2 /Xam

AP3913 2 Auen

AP3914 2 Auen

AP3915 ¢ Naron

AP3916 @¢ Naron

AP4608 92 Heikom

AP4609 ¢ Auen

AP4610 ¢ ? Bush

TABLE 1

DESCRIPTION

Boy with bow and arrow

Sitting

Looking at spoor

Pointing to spoor

Standing

Standing, holding baby

Sitting

Old woman standing

Sitting with son

Sitting with mother

Bust only

Standing

Holding a hare

Lifting arms

Holding a stick

Dancing

Sitting, pounding a bone

Boy standing, arms folded

Carrying a pot

Sitting

Sitting, stirring food

Bending forward

Standing

Standing, with stick

Half-reclining

Standing, right arm raised

Sitting

Walking, stick over shoulder

Bow in hand

Drawing bow

Sitting cross-legged

With digging-stick

Sitting, digging

Standing with hands on hips

v v v v >

Standing, resting on stick

Pounding snuff

Bending forward

Squatting as if using fire-sticks

Standing, arm out-stretched

Crouching

Crouching, arm out-stretched

Boy sitting

Boy playing musical instrument (gorah)

Reclining

Standing, right arm on hip

Standing, arms extended

Standing

Sitting, one knee raised

Standing, carrying pot on head

Kneeling, using pestle and mortar

Standing, body cicatrized

Head only

Standing

Head only

Standing

Boy standing, arm outstretched

Standing as if in trance

Standing

Sitting, with outstretched legs

Sitting on rock

Sitting as if making ostrich egg-shell beads

Sitting as if drilling bore in beads

Standing, clapping

Dancing

Standing, clapping

Sitting as if making string

Walking

LOCALITY

Kanye, Botswana

Gaborone, Botswana

Gaborone

Kanye

Nuragas

Otjito

Grootfontein

Lake Chrissie, Transvaal

Lake Chrissie

Possession Is.

Prieska, Cape

Prieska

Camavon

Prieska

~ 7 v A

Prieska

Carnarvon

Vosburg

Upington (cast at Tokai

reformatory)

Prieska

Victoria West

Britstown (cast in House of

Correction, Cape Town)

Britstown (cast in House of

Correction, Cape Town)

Cast at Kimberley prison

Kanye

Cast in Johannesburg

Bethany (cast in Windhoek prison)

Windhoek prison

Bethany (cast in Windhoek prison)

De Beers Compound, Kimberley

Walvis Bay

Gaborone prison

Carnarvon

Sandfontein

Grootfontein

Sandfontein

De Beers Compound, Kimberley

9LI

WAASAW NVOIed¥ HLAOS JHL JO STVNNV

SLOSfg0 WhgsaW sy SLograns NVWNH

LLI

178 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 5. Three young men at the /Xam encampment, Prieska, 1911. (Photograph: D. Bleek, South African

Museum Photographic Collection.)

supported by the presence of “physical characters that could not be ignored’ (SAM

Annual Report for 1911: 22). Social and cultural information relating to particular indi-

viduals, such as old Guiman Toonies who, in his youth, had been a hunter of gemsbok,

eland and kudu, was not considered as important as his physical attributes. Later in 1911,

when the casts were put on exhibition at the South African Museum, these physical

characters became the primary focus of public interest.

EXHIBITING THE CASTS

The exhibited figures were not intended to evoke a presence of the social beings who

had been cast at a particular time and place, but instead were presented as generalized

examples of a racial type. Separated from their social and historical context, the people who

were cast were literally objectivized and reduced to scientific specimens. People, who in

actual life were living as farm labourers or domestic servants, having survived a long period

of conflict with the Cape Government, became no more than examples of a physical type

(Fig. 6). Janikie Achterdam (Fig. 7), an informant of Dorothea Bleek, was one of the people

from Prieska who was cast by Drury. In sharp contrast to the wealth of cultural knowl-

edge of /Xam life that the Bleek records reveal, in the Museum the cast of Janikie was

displayed as a numbered specimen, bereft of all cultural and social context (Fig. 8).

The main label in the exhibition (Fig. 6) of a group of casts taken at Prieska in 1911

read as follows:

CAPE BUSHMEN: The Bushmen of the Cape appear to have been the purest-

blooded representatives of the Bushman stock, much purer than those of the Kalahari

HUMAN SUBJECTS AS MUSEUM OBJECTS 179

Fig. 6. The casts on display in the South African Museum, c. 1915.

(South African Museum Photographic Collection.)

and other more northerly districts. They are now practically extinct. They were light

in colour and of small or medium height; the prominent posterior development

(steatopygy) of the women was a characteristic feature of the race.

To anthropologists the Bushmen are one of the most interesting races in the world.

There are strong grounds for suspecting that they are of the same stock as the remote

Upper Palaeolithic period. This cannot yet be definitely asserted but recent discoveries

in North and East Africa have tended to strengthen the probability considerably.

(A key to the figures followed, giving approximate age of the subject and locality in

which the cast was made.)

It is noteworthy that, while drawing attention in this label to purity of stock, in the

Same year Péringuey published the results of Dr F. C. Shrubsall’s craniological analysis

of some 162 skulls, which showed no discernible difference between San and Khoikhoi

(Péringuey 1911). He stated ‘I have given up, now, distinguishing between a so-called

Bushman or a so-called Hottentot’ (Péringuey 1911: 197). These findings were

180 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 7. Janikie Achterdam, Prieska, 1911. (Photograph: D. Bleek,

South African Museum Photographic Collection.)

Vy Vd ie é ili

Fig. 8. Cast of Janikie Achterdam (SAM—AP3895) on display in the South African Museum, c. 1912.

(South African Museum Photographic Collection.)

HUMAN SUBJECTS AS MUSEUM OBJECTS 18]

communicated to a wider public in 1913 in a series of articles by Péringuey in the Cape

Argus. Furthermore, while stating in the Museum label that the Cape Bushmen as a

physical type were practically extinct, he commented in the newspaper that a surprising

number of people speaking a Bushman language were still to be found in the Northern

Cape. Considering that the purity of the /Xam had been attested in part on linguistic

criteria, this must have been confusing evidence for Péringuey. Even in his own terms,

the Museum label was contradictory to the known evidence.

The prominence accorded to the casts of /Xam and other Khoisan people was quite

out of proportion to any proven anthropological importance. According to Péringuey’s

own assessment in 1918, “The results of the examination of the large and representative

[collection of] material of the ‘San’ Race accumulated at such great cost and difficulties

by this Museum has not, I regret to say, materialized much’ (SAM Annual Report for

1918). No later academic research contradicted this point of view, but the exhibition of

casts none the less remained unchanged until 1932, when the casts were given even

greater prominence by being placed in the centre of a new Ethnology Gallery (Davison

1991).

DISCUSSION

In retrospect, the casting project, undertaken between 1907 and 1924, provides a

clear example of the way in which museum practice reduced people of another culture to

objects of study. Through the project, San and Khoikhoi people were classified and reified

as racial Others, a stereotype that was perpetuated in the way the casts were labelled and

exhibited in the Museum. For decades after Péringuey’s death in 1924 countless visitors,

including thousands of school-children each year, viewed exhibits that gave credibility to

a flawed anthropological notion of racial typology.

Although it was not only in South Africa that an evolutionary paradigm remained

the most prevalent model in ethnographic museums long after its rejection by academic

anthropology (Coombes 1988), in the socio-political context of South Africa this would

have reinforced and added credibility to a pervasive ideology of essential racial difference

and inequality. Moreover, the power of this ideology would have been made all the more

effective through being endorsed by an institution that claimed a position of scientific

neutrality. That the presentation of the casts was perceived as being value-free is con-

firmed in a press report on Drury and his modelling skills (Cape Times, 7 February 1925):

The value of the plaster casts lies in their absolute impartiality, their pure, unadul-

terated ‘objectivity’. They are the Bushmen themselves without the gloss of ‘inter-

pretation’ or extraneous adornment . . . every shade of facial expression is caught.

Every expression, indeed, that the mind projects through the physical organism is

recorded. —

However, as shown above in Péringuey’s memorandum to Drury, the casting project

was not impartial, nor was the presentation of the casts neutral, even if they appeared so

life-like and natural that viewers tended not to question the implicit assumptions under-

lying their exhibition. It was less politically contentious to represent ‘the Bushmen’ as an

endangered anthropological type, and therefore of scientific interest, than to confront

social issues of colonial dispossession, racial conflict and integration. The emphasis was

182 ANNALS OF THE SOUTH AFRICAN MUSEUM

on an idealized, hypothetical, ‘traditional’ past. No attempt was made to represent the

actual living conditions of the remnant San communities, among whom Drury found his

subjects. It was the ‘pure’ physical type beneath the European dresses, jackets and trou-

sers that was deemed to be of scientific interest. In pursuing this interest, the people who

were cast were distanced and denied the sensibilities of the observer.

Ironically, however, if viewers had looked closely at the casts on display, they could

have seen in Drury’s accurate rendering of the skin pigmentation of his subjects pale and

darker tones that correspond to the clothed parts of the body and those that were exposed

to the sun. This minutely observed detail, fulfilling Péringuey’s instructions to a fault,

bears subtle testimony to the ideas that informed the casting project. Abstracted from the

social and cultural context of their human subjects, the casts on display reveal Péringuey’s

scientific interests, Drury’s casting expertise and the unequal power relations inherent in

the execution of the project. In brief, the casts are authentic artefacts of scientific attitudes

and museum practice in the early twentieth century.

ACKNOWLEDGEMENTS

This paper benefited greatly from discussion with my colleague, Gerald Klinghardt.

I would like to express my thanks to him. I am also grateful to Professor Martin Hall and

Dr Alan Morris for their comments on an earlier version of the paper, and to June Hosford

and Aubrey Byron for photographic work.

REFERENCES

AtTick, R. D. 1978. The shows of London. Cambridge, Mass.: Belknap Press of Harvard University

Press.

ANNUAL REPORTS OF THE SOUTH AFRICAN MUSEUM. 1855-1941. Cape Town: Government Printer.

CLIFFORD, J. 1988. The predicament of culture. Cambridge: Harvard University Press.

CLIFFORD, J. & Marcus, G. E. (eds) 1986. Writing culture. The poetics and politics of ethnography.

Berkeley: University of California Press.

Coomses, A. 1988. Museums and the foundation of national and cultural identities. The Oxford Art

Journal 11 (2): 57-68.

Davison, P. 1991. Material culture, context and meaning. A critical investigation of museum practice,

with particular reference to the South African Museum. Unpublished Doctoral dissertation, Univer-

sity of Cape Town.

Drury, J. & DRENNAN, M. R. 1926. The pudendal parts of the South African Bush race. Medical

Journal of South Africa 22: 113-117.

Dusow, S. 1989. The idea of race in early 20th century South Africa: some preliminary thought.

Unpublished paper, Africa Seminar, University of Cape Town.

DuckwortH, W. L. H. 1904. Morphology and anthropology. A handbook for students. Cambridge:

Cambridge University Press.

FABIAN, J. 1983. Time and the Other. How anthropology makes its object. New York: Columbia

University Press.

GILMAN, S. 1986. Black bodies, white bodies: toward an iconography of female sexuality in late nine-

teenth century art, medicine and literature. Jn: Gates, H. L. (ed.) Race, writing and difference:

223-261. Chicago: University of Chicago Press.

GouLp, S. J. 1982. The Hottentot Venus. Natural History 10: 22-27.

GUENTHER, M. G. 1980. From ‘brutal savages’ to ‘harmless people’. Paideuma 26: 123-140.

Happon, A. C. 1906. [Presidential address to Section H.—Anthropology.] Report of the Seventy-fifth

meeting of the British Association for the Advancement of Science, South Africa, August and Sep-

tember 1905: 511-527.

Happon, A. C. 1934. History of anthropology. London: Watts & Co.

Harris, M. 1969. The rise of anthropological theory. London: Routledge and Kegan Paul.

182

HUMAN SUBJECTS AS MUSEUM OBJECTS 183

INGoLD, T. 1986. Evolution and social life. Cambridge: Cambridge University Press.

Kirsy, P. 1954. The Hottentot Venus of the Musee de l’Homme, Paris. South African Journal of

Science 50 (12): 319-322.

Kirsy, P. R. 1965. Sir Andrew Smith, M.D., K.C.B. His life, letters and works. Cape Town: Balkema.

Kuper, A. 1988. The invention of primitive society. Transformations of an illusion. London: Routledge.

Lye, W. F. (ed.) 1975. Andrew Smith’s Journal of his expedition into the interior of South Africa,

1834-36. Cape Town: Balkema.

Morris, A. 1987. The reflection of the collector: San and Khoi skeletons in museum collections. South

African Archaeological Bulletin 42: 12-22.

PERINGUEY, L. 1911. The Stone Ages of South Africa as represented in the collection of the South

African Museum. Annals of the South African Museum 8: 1-218.

SmitH, A. 1830. Observations relative to the origin and history of the Bushmen. The South African

Quarterly Journal 1: 171-189.

STOCKING, G. W. 1987. Victorian Anthropology. New York: The Free Press.

6. SYSTEMATIC papers must conform to the International 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

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An author’s name when cited must follow the name of the taxon without intervening punctuation

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Synonymy arrangement should be according to chronology of names, i.e. all published scientific

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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.

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Note punctuation in the above example:

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In describing new species, one specimen must be designated as the holotype; other specimens

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Note standard form of writing South African Museum registration numbers and date.

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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.

PATRICIA DAVISON

HUMAN SUBJECTS AS MUSEUM OBJECTS.

A PROJECT TO MAKE LIFE-CASTS OF “‘BUSHMEN’ —

AND ‘HOTTENTOTS’, 1907-1924

XVOLUME 102 PART 6 JANUARY 1993 ISSN 0303-2515

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ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

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CRETACEOUS FAUNAS FROM ZULULAND

AND NATAL, SOUTH AFRICA.

THE HETEROMORPH AMMONITE GENUS

EUBACULITES SPATH, 1926

HERBERT CHRISTIAN KLINGER

WILLIAM JAMES KENNEDY

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D1781

CRETACEOUS FAUNAS FROM ZULULAND AND NATAL,

SOUTH AFRICA.

THE HETEROMORPH AMMONITE GENUS EUBACULITES SPATH, 1926

HERBERT CHRISTIAN KLINGER

Department of Invertebrate Palaeontology, South African Museum, Cape Town

WILLIAM JAMES KENNEDY

Geological Collections, University Museum, Oxford

(With 57 figures)

[MS accepted 27 May 1991]

ABSTRACT

The taxonomy of the heteromorph ammonite genus Eubaculites Spath, 1926 (family Baculitidae),

is reviewed and the five most important species, Eubaculites carinatus (Morton, 1834), E. labyrinthicus

(Morton, 1834), E. vagina (Forbes, 1846), E. simplex (Kossmat, 1895) and E. latecarinatus

(Brunnschweiler, 1966), are described.

CONTENTS

PAGE

IAOGICTOR a. ¢:0:.0,6 0 ere. Wis. UESed-0 SLGUNS GIS Uae ace sae NOUN at Oa ee a 185

PocationmolespecliMensimrmr teeter Hs tha a, mee Nake meee Sle gk See ales 186

FHVENG) NOCRUISS 5g Sia 08.0 ot ceo Ce eRa Ie ve Ie ey A a at ae 186

DITA ODS. CG OSCUTNEMS, Sac. os oss Cee ne ee eee 186

SHENK RC RIMONO Vere ere NE Meneses UE SN hd es aaa els whe eae 186

Sy SUei Avice Alac ONUOlOCN Emu ay ck ce etenls SIMONSEN Nt co Rare wa: Gee leet: 186

Genisee i DaGulitesEs Alle ODOp ee ee we nie Silk Pied c maid cli blo Siseile So ade ere 187

BubaculinesMabyninthicuss (MOKOn S54) aes eee os eae 194

LMbacuiitesmy aoinan (hOnDeS S40) wean ote oe ee oe eae 203

Buvaculiiesncaninatisn(Nontons i834) ane ese soe es oe os oe ee 218

Eubaculites latecarinatus (Brunnschweiler, 1966) ..................... 238

PUDAGUINESE SUMP UNOSSIMNats 895) tye Seni aan ae as cee cae Day

PACKMOW COS CINCIS HE eae une Penn eRe Aide a eUe een LE us donee a Maa bee 260

NCTC CI CES MMR PRM R IT er i Meng el) erat ies eto ou 4, Sa MARIS Gigale « FeUeN 261

INTRODUCTION

The genus Eubaculites Spath, 1926, is a prominent member of the ammonite fauna

of the Zululand Maastrichtian, and is the dominant ammonite in terms of both biomass

and numbers of individuals at certain levels in both the Lower and Upper Maastrichtian

in parts of Zululand, Chile, Western Australia, and the U. S. Gulf Coast region. Although

unknown in Antarctica, the U. S. Western Interior, North and West Africa, Greenland,

Asia, Japan, and the Middle East, its otherwise wide distribution shows it to have been

one of the most important Maastrichtian heteromorphs. The five best known species,

185

Ann. S. Afr. Mus. 102 (6), 1993: 185-264, 57 figs.

186 ANNALS OF THE SOUTH AFRICAN MUSEUM

Eubaculites carinatus (Morton, 1834), Eubaculites labyrinthicus (Morton, 1834), Eubacu-

lites vagina (Forbes, 1846), Eubaculites simplex (Kossmat, 1895), and Eubaculites late-

carinatus (Brunnschweiler, 1966), are described and discussed below.

LOCATION OF SPECIMENS

The following abbreviations are used to indicate the location of specimens mentioned

in the text:

AMNH American Museum of Natural History, New York

ANSP Academy of Natural Sciences, Philadelphia

BMNH Natural History Museum, London

GSI Geological Survey of India, Calcutta

OUM Oxford University Museum

SAM South African Museum, Cape Town

SAS South African Geological Survey, Pretoria.

FIELD LOCALITIES

Details of field localities are given by Kennedy & Klinger (1975); further descrip-

tions of these localities are deposited in the Department of Palaeontology, Natural History

Museum, London, Geological Survey of South Africa, Pretoria, and Department of Inver-

tebrate Palaeontology, South African Museum, Cape Town.

DIMENSIONS OF SPECIMENS

All dimensions are given in millimetres. Wb = whorl breadth; Wh = whorl height.

The term ‘rib index’ means the number of ribs present in a distance equal to the whorl

height at the middle of the interval measured. Taper index is:

Larger whorl height-smaller whorl height

: x 100

Distance between measurements

SUTURE TERMINOLOGY

The suture terminology of Wedekind (1916), reviewed by Kullmann & Wiedmann

(1970), is followed here: I = internal lobe, U = umbilical lobe, L = lateral lobe, E =

external lobe.

SYSTEMATIC PALAEONTOLOGY

Phylum MOLLUSCA Cuvier, 1797

Class CEPHALOPODA Cuvier, 1797

Order AMMONOIDEA Zittel, 1884

Suborder ANCYLOCERATINA Wiedmann, 1966

Superfamily TURRILITACEAE Gill, 1871

CRETACEOUS FAUNAS FROM SOUTH AFRICA 187

Family Baculitidae Gill, 1871

Genus Eubaculites Spath, 1926

(= Giralites Brunnschweiler, 1966; Cardabites Brunnschweiler, 1966;

Eubaculiceras Brunnschweiler, 1966)

Type species. Baculites vagina Forbes var. ootacodensis Stoliczka, 1866: 199, pl. 90

(fig. 14), by the original designation of Spath (1926: 80) (= Baculites labyrinthicus

Morton, 1834: 44, pl. 13 (fig. 10)).

Diagnosis

Curved or straight baculitids with pyriform whorl section in adult stage; length may

exceed | m. Dorsum flat to rounded, venter fastigiate or with a tabulate siphonal keel, the

edges of which are predominantly sharp but subrounded in some. Siphonal keel smooth,

ribbed or crenulated. In juveniles, the whorl section may be ovoid, lacking a tabulate or

fastigiate venter. Venter generally changes from fastigiate to keeled, with subrounded to

tabulate top during ontogeny; in some it remains fastigiate throughout. Development of

flank ornament variable; growth lines only, crescentic ribs, or up to two distinct rows of

tubercles. Ribs may extend to dorsolateral or ventrolateral region. Aperture apparently

simple, with ventral rostrum and sinuous lateral indentation. Suture with plump, finely

incised saddles and lobes and phylloid elements in some saddles.

Discussion

The genus Eubaculites was introduced in characteristically brief fashion by Spath

(1926: 80) as: ‘Eubaculites gen. nov. for the carinate forms of the group of E. vagina

(Forbes) and E. otacodensis Stoliczka sp. (Kossmat, “Unters. Siidind. Kreideform. Beitr.

Pal. Osterr.-Ung., vol. ix, 1895, p. 157, pl. xix, figs 15a, b genotype).’ Wright (1957:

L218) was the first to provide a diagnosis of Eubaculites: ‘Section pear-shaped, with flat

venter; ribs normally faint towards venter but on inner part form prominent long curved

bullae; row of lower lateral tubercles may be present. Suture with plump, minutely frilled

elements.’

Matsumoto (1959) described a limited Eubaculites fauna from California, and essen-

tially followed Wright (1957) in his interpretation of the genus; however, he used the term

‘tabulate ventral keel’ instead of ‘flat venter’, because he believed Eubaculites could be

derived from keeled baculitid species such as Baculites occidentalis.

Subsequent descriptions of eubaculitid faunas from various regions, e.g. Western

Australia (Brunnschweiler 1966), Zululand, South Africa (Klinger 1976), Argentina

(Riccardi 1974), Chile (Hiinicken & Covacevich 1975; Stinnesbeck 1986), the Nether-

lands (Kennedy 1986c), Austria (Kennedy & Summesberger 1986), and southern France

(Kennedy 1986a), have differed considerably in their interpretation of the genus—to the

extent that Brunnschweiler placed it in a separate subfamily, Eubaculitinae, in which he

recognized four genera.

It is necessary to look at the genus in terms of the type species. Kennedy (1986c:

193) pointed out that Spath’s ‘genotype’ (type species) is a species and not an

individual. Eubaculites ootacodensis is thus the type species of Eubaculites. The

lectotype of E. ootacodensis by subsequent designation of Kennedy (1986c: 193) is the

original of Stoliczka (1866, pl. 90 (fig. 14)) from the Maastrichtian Arrialoor Group of

188 ANNALS OF THE SOUTH AFRICAN MUSEUM

South India, whereas (as shown below) E. ootacodensis is a synonym of Eubaculites

labyrinthicus (Morton, 1834).

Because Spath (1926) did not provide a diagnosis of Eubaculites, it is possible

to decide on the limit of his generic concept only on the basis of species he referred

to it. That he included Eubaculites vagina of Forbes (1846a), the type material of which

was available to him for study, shows that the distinctive feature of this species—

a broad tabulate venter and bituberculate ribbed flanks—were encompassed. That

he cited plate 19 (fig 15) of Kossmat (1895) as a reference to the species, rather than

the original figure, is unfortunate, as this figure is of an incomplete specimen, lacking

the critical features of the venter (Fig. 7F—H). It is, however, a topotype of the species,

likely to be conspecific with E. ootacodensis of Stoliczka (= Eubaculites labyrinthicus

(Morton, 1834)) and thus characterized by binodose, ribbed flanks and a fastigiate venter.

In describing the Maastrichtian faunas from Western Australia, Spath (1940: 49)

referred Kossmat’s Baculites vagina var. simplex (1895: 156(60), pl. 19(5) (figs 13a—b,

14a-—cb)) to Eubaculites, but nevertheless separated it from Eubaculites vagina *

because it is a passage-form between Eubaculites and Baculites s.s’—thus including

forms with both tabulate and fastigiate venters in Eubaculites. Later (1953: 20), he seemed

to regard possession of a tabulate venter as an essential feature: ‘The true Eubaculites

vagina (Forbes) which I discussed recently (1940, p. 48) has a perfectly tabulate, not a

sharpened, venter already at 10 mm (long diameter) . . . so that Baculites cazadorianus is

neither a Eubaculites nor a variety of E. vagina.’ In the same article (1953: 46), he also

referred to Baculites vagina var. simplex Kossmat as E.(?) simplex. Both Wright (1957:

L218) and Matsumoto (1959: 166) emphasized the tabulate shape of the venter of

Eubaculites. Klinger (1976: 90) noted that Stoliczka’s figure of E. ootacodensis showed

a fastigiate venter, although his description may have been based in part on specimens

with tabulate venters (“. . . the siphuncle often lies nearer to one edge than to the other;

this, however, is not constant in all specimens’—Stoliczka 1866: 199).

Brunnschweiler (1966) described (amongst other heteromorphs) a eubaculitid assem-

blage of about 200 specimens from the Miria Marl of Western Australia—until then the

most extensive collection of the genus described. This included forms with tabulate and

fastigiate venter, ribbed and smooth forms, and inflated and compressed forms. These he

referred to a new subfamily Eubaculitinae and four genera. This subfamily was distin-

guished from Baculitinae (Brunnschweiler 1966: 24) for those forms ‘with a ventral keel

which appears in very early growth stages. The keel is either acute or tabulate, never

rounded.’

Brunnschweiler’s criteria for recognition of the four genera he referred to the sub-

family Eubaculitinae were as follows:

Genus Eubaculites, type species E. ootacodensis (Stoliczka). This genus was restricted to

those costate forms that are truly similar to the type species in cross-section and shape of

the keel.

Genus Giralites Brunnschweiler, 1966, type species Giralites latecarinatus Brunn-

schweiler, 1966. The genus included non-costate species in which the Wh : Wb ratio is

less than 1,8: 1.

Genus Eubaculiceras, type species Eubaculiceras compressum Brunnschweiler, 1966.

This included ribbed species with a very compressed whorl section and a Wh : Wb ratio

Ol a lnonmimonre:

CRETACEOUS FAUNAS FROM SOUTH AFRICA 189

Genus Cardabites Brunnschweiler, 1966, type species Cardabites tabulatus Brunn-

schweiler, 1966. This is a non-costate genus with a very compressed whorl section with

a Wh: Wb ratio of 2: | or more.

Klinger (1976: 83-84) regarded all these as synonyms of Eubaculites and, for strati-

graphic purposes, recognized four species groups based primarily on lateral ornament and

secondarily on compression of the whorl section, although admitting that the divisions

were arbitrary and that the groups were connected by transitions:

1. Eubaculites gr. ex. vagina (Forbes) with pyriform whorl shape, Wh : Wb less than

2 : 1, ornament consists of tubercles at mid-flank and at dorsolateral edge.

2. Eubaculites gr. ex. ootacodensis (Stoliczka). Whorl section as in E. vagina; lateral

ornament consisting of strong crescentic ribs. Venter tabulate or fastigiate.

3. Eubaculites gr. ex. latecarinatus (Brunnschweiler). Whorl section as above, but flanks

devoid of all ornament.

4. Eubaculites gr. ex. compressum (Brunnschweiler). Whorl section strongly compressed;

Wh : Wb, 2 : 1 or more. Lateral ornament, if present, very weak. Venter fastigiate or

tabulate.

Kennedy (1986a: 1015; 1986c: 194) and Kennedy & Summesberger (1986: 197)

accepted Klinger’s (1976) species groups with some modifications, but considered the

shape of the ventral keel to be of major significance. Kennedy (1986a: 1015, 1986c: 194)

recognized three species groups:

1. Species with fastigiate venter and flattened dorsum, ribbed or smooth, e.g. Baculites

ootacodensis Stoliczka, 1866, Baculites vagina var. simplex Kossmat, 1895, Baculites

rioturbioensis Htinicken, 1965, Eubaculiceras fastigiatum Brunnschweiler, 1966, Carda-

bites scimitar Brunnschweiler, 1966, and Baculites argentinicus Weaver, 1927—the last

named a nomen dubium and possibly a corroded fragment of B. rioturbioensis.

_ 2. Species with tabulate venter, flattened dorsum, dorsolateral and dorsal tubercles, some

individuals smooth, e.g. Baculites vagina Forbes, 1846, and Baculites ornatus d’Orbigny,

1847.

3. Species with tabulate venter and flattened dorsum generally ornamented by flank

ribs, some individuals may be smooth, e.g. Baculites lyelli d’ Orbigny, 1847, Eubaculites

kossmati Brunnschweiler, 1966, Eubaculites multicostatus Brunnschweiler, 1966,

Giralites latecarinatus Brunnschweiler, 1966, Giralites quadrisulcatus Brunnschweiler,

1966, Eubaculiceras compressum Brunnschweiler, 1966, and Cardabites tabulatus

Brunnschweiler, 1966.

Stinnesbeck (1986) discussed Eubaculites lyelli from Quiriquina, Chile, on the basis

of several hundred specimens; he concluded that transitions to E. latecarinatus, E. com-

pressum and E. ootacodensis occur in his material (of E. lyelli) but these cannot, as yet,

be used for stratigraphic purposes. As far as we can ascertain, no typical examples of

E. latecarinatus, E. compressum or E. ootacodensis occur at Quiriquina.

A subsequent unpublished revision of North American Eubaculites by Kennedy &

Cobban (in preparation) revealed that the prior name for Baculites lyelli of d’ Orbigny

(1847) was Baculites carinatus (Morton, 1834), and that for Baculites vagina var. oota-

codensis of Stoliczka (1866) was Baculites labyrinthicus of Morton (1834), as described

below.

190 ANNALS OF THE SOUTH AFRICAN MUSEUM

Henderson et al. (1992) revised the Eubaculites of the Miria Formation and under-

lying phosphate horizon at the top of the Korojong Calcarenite in the Carnarvon Basin,

Western Australia. They recognized Eubaculites latecarinatus (with Giralites quadrisul-

catus Brunnschweiler (1966) and Eubaculites ambindensis of Collignon (1971) as syno-

nyms) as the oldest species, present only in the phosphatic nodule bed at the top of the

Korojong Calcarenite. The overlying Miria Formation yielded two species: Eubaculites

carinatus (of which Eubaculites ootacodensis of Brunnschweiler (1966, non Kossmat),

E. vagina of Brunnschweiler (1966, non Forbes), and E. kossmati and E. multicostatus are

synonyms) and E. simplex (of which Eubaculiceras compressum, E. fastigiatum,

Cardabites tabulatus and C. scimitar are synonyms). They also showed E. simplex to

include individuals with both tabulate and fastigiate venter through ontogeny, as well as

those with an initially fastigiate venter that develop a tabulate venter subsequently.

Baculites occidentalis Meek, 1862 (see e.g. Usher 1952: 98, pl. 28 (fig. 1), pl. 31

(fig. 19), text-fig. 4; Matsumoto 1959: 150, pl. 35 (figs 2a—d, 3a—d), pl. 36 (fig. la—d),

pl. 41 (fig. la—d), pl. 42 (fig. la—c, 2a—c), text-figs 64, 65a—b, 66, 67-71; Obata & Mat-

sumoto in Matsumoto & Obata 1963: 82, pl. 23 (fig. 3), pl. 25 (fig. 1), pl. 26 (figs 1-3),

pl. 27 (figs 2-5, 8), text-figs 172-186), has an indistinct siphonal keel with rounded

edges, and was regarded by Matsumoto (1959: 154) as an intermediate stage between

Baculites and Eubaculites. We here tentatively regard it as an early form of Eubaculites.

Baculites regina Obata & Matsumoto (in Matsumoto & Obata 1963: 85, pl. 22

(figs 3-6), pl. 23 (figs 1-2), pl. 24 (figs 1-5), pl. 25 (figs 3-5), pl. 27 (figs 1, 6-7, 9),

text-figs 191-196, 200-214) from the Campanian of Honshu, was compared to Eubacu-

lites by Obata & Matsumoto (in Matsumoto & Obata 1963: 90), because of its flattened

dorsum, subangular dorsal edges and differentiation of lateral ornament. However, it lacks

a tabulate siphonal keel and is better regarded as a lateral offshoot of Eubaculites occi-

dentalis rather than in the main line of descent to Eubaculites, according to Obata &

Matsumoto (in Matsumoto & Obata 1963: 91).

Baculites lomaensis Anderson (1958: 191, pl. 48 (figs 5—6)) was regarded as being

related to Eubaculites vagina by Anderson, but it appears that this opinion was based

mainly on stratigraphic grounds. Baculites lomaensis lacks a tabulate siphonal keel and

has a suture line similar to early true Baculites.

We here recognize the following species of Eubaculites, synonyms of the most

important which are given below: Eubaculites carinatus (Morton, 1834), Eubaculites

labyrinthicus (Morton, 1834), Eubaculites vagina (Forbes, 1846), Eubaculites latecari-

natus (Brunnschweiler, 1966), Eubaculites simplex (Kossmat, 1895), and Eubaculites

occidentalis (Meek, 1862). Baculites binodosus Noetling, 1897, and Baculites riotur-

bioensis, Hiinicken, 1965, may be further valid species.

Occurrence

Where well dated, Eubaculites is typically Maastrichtian, ranging throughout most

of the stage; Eubaculites occidentalis may appear in the Upper Campanian (for the basis

of a Campanian date, see Ward 1978a, 1978b). The evolutionary origin of the genus is

not clear; it probably lies in the group of Baculites chicoensis Trask, 1856 (see Matsu-

moto 1959: 145, pl. 36 (fig. 2), pl. 37 (fig. 1), text-figs 59a—-d, 60a—b, 6la—b, 62a-b,

63a—b), from the Lower Campanian of California and British Columbia, but similarities

in ornament between E. carinatus and E. labyrinthicus, and Baculites sp. (nov.?) cf.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 19]

B. aquilaensis Reeside (in Collignon, 1970: 81, pl. 639 (fig. 8, 8a); see Fig. 57) from the

Upper Campanian of Madagascar and Baculites vanhoepeni from the Campanian of

Zululand, are striking. However, this probably is due to homoeomorphous develop-

ment only. Unfortunately, the phylogeny of the genus is difficult to elucidate, mainly

because of the very poor level of inter-regional correlation of the Maastrichtian at the

present time, this difficulty also extending to the definition of the base of the stage

(Birkelund et al. 1984) and division into substages. The present state of knowledge of

Eubaculites in its main areas of occurrence is discussed below.

South Africa. Details of the distribution of Eubaculites in Zululand were given by

Klinger (1976). These are emended here on the basis of new collections and the taxo-

nomic revision given below. Specimens previously referred to E. ootacodensis are

E. carinatus, aS are some specimens previously referred to E. vagina.

Stratigraphic distribution is as follows: Eubaculites labyrinthicus—Maastrichtian II;

Eubaculites carinatus—Maastrichtian a, Maastrichtian I-III; Eubaculites latecarinatus—

Maastrichtian ?a, Maastrichtian I—?II; Eubaculites simplex—Maastrichtian I-II]; Eubacu-

lites vagina (?)—Maastrichtian I.

Eubaculites carinatus is most abundant in Maastrichtian III, but occurs as early as

Maastrichtian a (ex Campanian IV of Kennedy & Klinger 1975). Eubaculites latecari-

natus is very abundant in Maastrichtian I and possibly persists to Maastrichtian II. Off-

shore data (Klinger et ai. 1980) suggested it may already occur as early as Maastrichtian a.

Eubaculites simplex is known from Maastrichtian I-II. Typical binodose adult E. vagina

are not known from Zululand—possible E. vagina (herein interpreted as E. labyrinthicus)

are known from Maastrichtian I.

Mozambique. Material described by Crick (1924) is poorly preserved, but is here

identified as follows: Baculites sheringomensis = Eubaculites carinatus; Baculites vagina

var. ootacodensis = Eubaculites carinatus; and Baculites vagina vat. simplex = Eubacu-

lites carinatus.

Chile. The Quiriquina Formation has yielded only one species, E. carinatus, which

is also the most common ammonite there (Fig. 37) (cf. Htinicken & Covacevich 1975;

Stinnesbeck 1986). Stinnesbeck (1986: 207) claimed that transitions to E. latecarinatus,

E. compressum and E. ootacodensis are present in his collections but stated that these do

not, as yet, have any stratigraphic significance (E. ornatus d’Orbigny, 1847, is most

probably a specimen of E. vagina from Pondicherry, India). Stinnesbeck (1986) dated

the Zone of Eubaculites lyelli (= E. carinatus) as upper Lower Maastrichtian to Upper

Maastrichtian.

Argentina. Eubaculites carinatus is relatively common in the Neuquén Basin of

northern Patagonia. Riccardi (1974) dated it as Lower—Middle Maastrichtian.

Weaver (1927) described Baculites argentinicus from the Cerro Huantraico in

Neuquén Province. This is based on two fragmentary specimens (see Olsson 1944: 105

(263)) and has generally been interpreted as a nomen dubium (cf. Riccardi 1974: 397;

Kennedy 1986a: 1015; 1986c: 194). Camacho (1967) recorded this species from Paso del

Sapo on the Chubut River, and regarded it as a Eubaculites. It belongs to the group of

E. simplex in having a fastigiate venter. Camacho (1968: 328, pl. 3 (figs 5a—b, 6)) recorded a

baculitid with a fastigiate venter, and a true Eubaculites with tabulate venter from this area.

Subsequent work by M. Uliana (in Riccardi 1974: 396) has yielded Baculites rioturbioensis.

192 ANNALS OF THE SOUTH AFRICAN MUSEUM

Baculites argentinicus and B. rioturbioensis appear to be synonyms, and are contempo-

raries of Eubaculites.

Baculites rioturbioensis was described from the ‘Yacimiento Rio Turbio’, Santa

Cruz Province in the Austral Basin of Patagonia, near the boundary with Argentina

and Chile. Hiinicken (1965: 63) dated it as uppermost Campanian to basal Maas-

trichtian. Baculites rioturbioensis has a flat dorsum, fastigiate venter, and broad

undulating lateral ribs; it belongs to the group of E. simplex. A ribbed specimen of

Eubaculites was described from a borehole in Santa Cruz Province by Garcia &

Camacho (1965: 72, pl. 1 (fig. 4)).

There may thus be an older Eubaculites carinatus—Baculites rioturbioensis assem-

blage and a younger one with E. carinatus alone.

Western Australia. Based on the work of Henderson & McNamara (1985) and

Henderson et al. (1992), the following sequence can be recognized in the Carnarvon

Basin. The oldest assemblage with Eubaculites comes from the nodule bed at the top of

the Korojong Calcarenite (12—20 cm), with Eubaculites latecarinatus, Nostoceras (N.)

attenuatus Brunnschweiler, 1966, N. (N.) fischeri Brunnschweiler, 1966, and Gun-

narites kalika (Stoliczka, 1865). The lower part (75-150 cm thick) of the overlying

Miria Formation yields Eubaculites simplex and rare non-heteromorphs, including

Phyllopachyceras forbesianum (d’Orbigny, 1850), Pachydiscus jacquoti australis

Henderson & McNamara, 1985, Gunnarites kalika, Neophylloceras surya (Forbes,

1846) and Gaudryceras kayei (Forbes, 1846). The upper part of the Miria Formation

(45-50 cm) yields abundant Eubaculites carinatus, common Diplomoceras cylin-

dracum (Defrance, 1816) and Glyptoxoceras rugatum (Forbes, 1846), rare Baculites

lechitides Brunnschweiler, 1966, and 22 species of non-heteromorphs described by

Henderson & McNamara (1985). The succeeding Palaeocene Boongarooda Greensand

yields reworked Eubaculites carinatus.

South India. The oldest occurrence of Eubaculites in South India is that of Eubacu-

lites labyrinthicus (as E. vagina var. ootacodensis) and E. simplex in the white sandstone

of the Arrialoor Group north of Ootacod, near Arrialoor, which also yielded Pachydiscus

tweenianus (Stoliczka, 1865) and Anapachydiscus arrialoorensis (Stoliczka, 1865), and is

said to yield Hauericeras gardeni (Baily, 1955). (Pachydiscus preegertoni Collignon,

1952, was also illustrated from an unspecified horizon near Arrialoor as Ammonites eger-

tonianus Forbes of Stoliczka 1865, pl. 5 (figs 1, 1a); fide Matsumoto et al. 1986: 5.)

Much younger are records from the Valudavur Formation of Pondicherry.

Eubaculites vagina comes from the Anisoceras beds of various authors and is prob-

ably from a horizon equivalent to the lower part of the Upper Maastrichtian Abathom-

phalus mayaroensis (planktonic foraminiferan) Zone (Rajagoplan 1965; Govindan

1972). Eubaculites lyelli probably occurs at an even higher horizon, the lower part of

the Trigonarca Beds of Kossmat, horizon D of Warth (1895) (e.g. Baculites vagina

var. ootacodensis Stoliczka of Kossmat 1895, pl. 19(5) (fig. 16)).

Madagascar. The records from Madagascar (Collignon 1971) indicate the presence of

E. vagina, E. labyrinthicus (as E. ootacodensis), E. simplex and E. latecarinatus (as Baculites

occidentalis and E. ambindensis) at the following localities (horizons): E. vagina—localities

504, 671; E. simplex—localities 503, 666; E. latecarinatus—localities 504, 665-7; and

E. labyrinthicus—localities 503, 504.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 193

We are unable to place these localities in sequence, but note that E. vagina, E. late-

carinatus and E. labyrinthicus co-occur at locality (horizon) 504.

Japan. No typical Eubaculites has been recorded, but for doubtful Baculites occiden-

talis and B. regina Obata & Matsumoto from uncertain Upper Campanian to lower Maas-

trichtian of south-west Japan (in Matsumoto & Obata 1963).

United States. The oldest recorded Eubaculites is E. occidentalis, which occurs in the

Upper Campanian—Lower Maastrichtian of California and British Columbia (Matsumoto

1959; Ward 1978a, 1978b). Eubaculites carinatus from the presumed Maastrichtian of

California was recorded as misidentified Baculites chicoensis Gabb (1864 pars) and

Eubaculites ootacodensis by Matsumoto (1959). In the Gulf Coast region, the Corsicana

Formation of north-east Texas yields Eubaculites carinatus and is equivalent to the mid-

Maastrichtian Globotruncana gansseri (planktonic foraminiferan) Zone (Kennedy & Cob-

ban unpublished). The Owl Creek Formation of Missouri and Mississippi yields common

E. carinatus (Conrad 1858; Stephenson 1955) and is referred to the upper part of the

G. gansseri zone by C. C. Smith (pers. comm. 1990). The Maastrichtian Prairie Bluff

Chalk in Alabama is the source of the type material of E. carinatus and E. labyrinthicus,

but these occurrences are difficult to date precisely as they are from phosphatic layers that

include elements from more than one horizon in the Maastrichtian, at least in some places.

In New Jersey, E. labyrinthicus occurs in the Maastrichtian middle part of Navesink

Formation, and EF. carinatus has been found reworked into the base of the succeeding

Palaeocene.

Europe. There is a well-substantiated belemnite zonation from the White Chalk of

north-west Europe, into which the scattered occurrences of Eubaculites carinatus can be

placed. This zonation is:

ZONE

ars Belemnella casimirovensis

Upper Maastrichtian

Belemnitella junior

fe peule Belemnella occidentalis

Lower Maastrichtian

Belemnella lanceolata

Eubaculites carinatus first appears at Neuberg, Steiermark, Austria, at a horizon

equivalent to the lower part of the Belemnella occidentalis Zone (Kennedy & Summes-

berger 1986: 200), that is to say, somewhere in the Globotruncana gansseri (planktonic

foram) Zone. The last appearance of EF. carinatus is in the Petites-Pyrénées of south-

eastern France (Kennedy et al. 1986), where the species occurs with forms of Hoplo-

scaphites constrictus that indicate the Belemnella casimirovensis Zone, and foraminifera

indicating the upper part of the Abathomphalus mayaroensis (planktonic foraminiferan)

Zone. There is also a mass occurrence of this species in the A. mayaroensis Zone,

approximately 40 m below the base of the Palaeocene at Hendaye (Pyrénées Atlantiques),

France. A specimen from the Maastricht area (Kennedy 1986c: 195, pl. 27 (figs 5—8)) is

from either the upper part of the B. junior Zone or the lower part of the B. casimirovensis

Zone.

It is difficult to determine absolute ranges for Eubaculites species from these data,

because of difficulties in long-distance correlation. Using Upper and Lower Maastrichtian

194 ANNALS OF THE SOUTH AFRICAN MUSEUM

in a north-west European sense, FE. carinatus is seen to be long-ranging, from middle

Lower to upper Upper Maastrichtian. Eubaculites vagina is probably lower Upper Maas-

trichtian, overlapping with E. carinatus, but not occurring with it in the Valudavur For-

mation in South India.

Evidence from Western Australia suggests E. simplex to be Upper Maastrichtian;

that it is older than E. vagina in South India might suggest lower Upper Maastrichtian.

Eubaculites latecarinatus is older than E. simplex in Western Australia, whereas

E. labyrinthicus occurs with E. carinatus and E. latecarinatus in Maastrichtian II in

Zululand.

The succession of Eubaculites may thus be:

E. carinatus (youngest)

E. carinatus, E. vagina

E. carinatus, E. simplex (2plus E. labyrinthicus)

E. carinatus, E. labyrinthicus, E. latecarinatus

E. carinatus

E. occidentalis (oldest).

Eubaculites labyrinthicus (Morton, 1834)

Figs 1-6, 7F-H, 36A-D, 52A-C

Baculites labyrinthicus Morton, 1834: 44, pl. 13 (fig. 10).

Baculites vagina Forbes var. Ootacodensis Stoliczka, 1866: 199, pl. 90 (fig. 14) (non fig. 15).

Baculites vagina var. Otacodensis Stoliczka: Kossmat, 1895: 157, pl. 19(5) (?fig. 15) (non fig. 16 =

E. carinatus). Cottreau, 1922: 180(72), pl. 9117) (fig. 11—11b). Diener, 1925: 63 (pars).

Baculites vagina Forbes: Spengler, 1923: 54, pl. 4 (fig. 9). Besairié, 1930: 222, pl. 26 (figs 12-13).

non Eubaculites otacodensis (Stoliczka): Spath, 1940: 49, pl. 1 (fig. 3), text-fig. 1b (= E. carinatus).

non Eubaculites ootacodensis (Stoliczka): Wright, 1957: L218, text-figs 245, 246a—c (= E. carinatus).

non Eubaculites ootacodensis (Stoliczka): Matsumoto, 1959: 166, pl. 43 (fig. 6), pl. 44 (figs 1-3), text-

figs 84-85 (= E. carinatus).

non Eubaculites ootacodensis (Stoliczka): Brunnschweiler, 1966: 27, pl. 1 (figs 9-14), text-figs 9-11

(= E. carinatus).

?Eubaculites sp. Leanza, 1967: 55, pl. 1 (figs 6-7).

Baculites simplex Kossmat: Collignon, 1971: 15, pl. 645 (figs 2388-2389).

non Eubaculites otacodensis Stol.: Collignon, 1971: 18, pl. 646 (fig. 2395) (? = E. carinatus).

non Eubaculites ootacodensis (Stoliczka): Riccardi, 1974: 388, pl. 1 (figs 1-7), pl. 2 (figs 1-4, 6), pl. 3

(figs 1-6), pl. 4 (figs 1-7), text-fig. 2 (= E. carinatus).

Eubaculites ootacodensis (Stoliczka): Klinger, 1976: 90 (pars) pl. 39 (fig. 3 only) (non fig. 1); non pl. 41

(figs 1-2), pl. 42 (figs 3, 8), text-fig. llc (= E. carinatus). Kennedy, 1986c: 194 (pars).

Eubaculites labyrinthicus (Morton): Kennedy & Henderson, 1992: 716, text-fig. 5D—-K.

Types

There are two syntypes (Fig. 1). The original of Morton (1834, pl. 13 (fig. 10)) is

ANSP 72868; the second specimen is ANSP 72869. Both are from the Prairie Bluff Chalk

of Alabama. The lectotype of Baculites ootacodensis is the original of Stoliczka (1866,

pl. 90 (fig. 14a-c)) (Fig. 5), GSI 406 and GSI 407 in the Collections of the Geological

Survey of India, from ‘the white gritty sandstone north of Ootacod, near Arrialoor’.

Material

SAM-PCZ8652 from Bed 3(C) and OUM-KX1828-1833, from Bed 7 (G) at local-

ity 20, Zululand, St Lucia Formation, Maastrichtian [-II.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 195

Fig. 1. Eubaculites labyrinthicus (Morton, 1834). A-C. Syntype,

ANSP 72868. D-F. Syntype, ANSP 72869. Both from

the Prairie Bluff Chalk of Alabama. Collection of the

Academy of Natural Sciences of Philadelphia. x 1.

Diagnosis

Whorl section cuneiform, with flattened dorsum and fastigiate venter, lacking siphonal,

tabulate keel; lateral ornament consists of two rows of tubercles, linked by ribs of variable

Strength.

Description

This is a rare species. In addition to the types, we have seen four other fragments

from the Prairie Bluff Chalk. All are phosphatic internal moulds; they range from 8,0 mm

to 17,5 mm in whorl height. The largest specimen (syntype ANSP 72868), figured by

Morton (1834, pl. 13 (fig. 10)), is wholly septate throughout (Fig. 1A—C). The expansion

rate is moderate, the whorl section compressed, with intercostal whorl breadth to height

ratio 0,58. The dorsum is flattened and barely convex. The dorsolateral margin is. nar-

rowly rounded, the dorsal flanks flattened, mid-flank region broadly rounded, ventral

flanks flattened and convergent, and the venter fastigiate. Strong nodes, elongated parallel

to the length of the shell, perch on the dorsalmost flank, two occurring in a distance equal

to the whorl height. A low, broad rib connects the node to a similarly elongated but

slightly obliquely aligned mid-lateral tubercle. The mid-lateral tubercles give rise to deli-

cate riblets and striae, also present on the interspaces, and projected forwards to intersect

196 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 2. Eubaculites labyrinthicus (Morton, 1834). A-D. OUM KX1831. E-H. OUM KX1832.

Both from Bed 7 (G), locality 20, Zululand, St Lucia Formation, Maastrichtian Il. Both x 2.

the siphonal line of the mid-venter at an acute angle. The dorsum is decorated by feebly

convex ribs and striae. The suture has broad, bifid, rectangular saddles, L/U with variable

degree of incision, E/L narrow and bifid, and U broad and bifid.

A cast of the lectotype of Baculites ootacodensis, here considered a junior synonym

of E. labyrinthicus, shows it to be a wholly septate internal mould, 56 mm long (GSI 406,

Fig. 5), with a maximum preserved whorl height of 15,5 mm. The intercostal whorl

breadth to height ratio is 0,62, the dorsum broad and very feebly convex to flat. The

dorsolateral margin is narrowly rounded, the dorsal flanks flattened in intercostal section,

the mid-flank region broadly rounded, the ventral flanks flattened, converging to a bluntly

CRETACEOUS FAUNAS FROM SOUTH AFRICA 197

Fig. 3. Eubaculites labyrinthicus (Morton, 1834). A-C. OUM KX1830. D-G. OUM KX1829.

Both from Bed 7 (G), locality 20, Zululand, St Lucia Formation, Maastrichtian II. Both x 2.

fastigiate venter. There are two ribs in a distance equal to the whorl height, parts of four

ribs being preserved on the specimen. They begin at small dorsolateral nodes, perched on

the dorsolateral margin, are elongated parallel to the length of the shell, and are coarse,

transverse and feebly prosiradiate on the inner flank, strengthening into a mid-lateral

tubercle that is elongated in a slightly prorsiradiate direction. These tubercles give rise to

a progressively weakening prorsiradiate rib that sweeps forwards and declines before

reaching the venter. The flanks are otherwise smooth, but the mid-venter is distinctly

crenulated.

198 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 4. Eubaculites labyrinthicus (Morton, 1834). Smooth variants with fastigiate venter, probably of

E. labyrinthicus. A-D. OUM KX1828. E-G. OUM KX1833. Both from Bed 7 (G) at locality 20,

Zululand, St Lucia Formation, Maastrichtian II. Both x 2.

The Zululand specimens referred to the species (Figs 2-4, 52A—C) are all rather

small body chambers, with a maximum preserved whorl height of 14,5 mm, and a whorl

breadth to height ratio of 0,65. They have weaker ornament than the types of both

Eubaculites labyrinthicus and E. ootacodensis, rather matching specimens from the

Prairie Bluff Chalk, such as those shown in Figure 1; in fact, some specimens are nearly

smooth (Figs 3-4). Material from Madagascar, here referred to E. labyrinthicus, includes

specimens larger than any known from the Prairie Bluff Chalk or the Valudavur Forma-

tion (e.g. Collignon 1971: 15, pl. 645, (fig. 2388), as Baculites simplex—herein Fig. 6;

Cottreau, 1922: 180, pl. 9 (fig. 11), as Baculites vagina var. otacodensis). In these, lateral

CRETACEOUS FAUNAS FROM SOUTH AFRICA 199

A B C D E

Fig. 5. Eubaculites labyrinthicus (Morton, 1834). Cast of lectotype of E. ootacodensis

(Stoliczka, 1866), the original of Stoliczka, 1866, pl. 90 (fig. 14), from Ootacod. Collection

of the Geological Survey of India, no. 406 x 1.

ornament consists predominantly of prominent crescentic ribs, with little or no sign of

development of tubercles. We assume these to be adults of forms with bituberculate

ornament in the juvenile stage. Given more material, it may be possible to separate those

forms with lateral ribbing only and fastigiate venter from those with lateral ribbing, two

rows of tubercles and fastigiate venter. The relationship of these predominantly ribbed to

ribbed and bituberculate forms may be analogous to that of E. carinatus to E. vagina.

Discussion

Eubaculites labyrinthicus has been a neglected species, but comparison of the type

and other material from the Prairie Bluff Chalk with the lectotype of E. ootacodensis and

the Zululand and Madagascan material referred to the species, show this to be a distinctive

form, the combination of predominantly binodose flanks and a fastigiate venter distin-

guishing it from all other species referred to the genus. Eubaculites vagina (Forbes, 1846)

(Figs 8—20) has binodose flanks, but the type population shows it to have a broad, flat

tabulate venter at a size where E. labyrinthicus is fastigiate. Some variants of E. carinatus

develop an incipient binodose condition (e.g. Brunnschweiler 1966, text-fig. 11), but the

broad, tabulate ventral keel is immediately distinctive.

Eubaculites ootacodensis has been widely misinterpreted. In his original account,

Stoliczka (1866: 199) referred to this species as differing from E. vagina in that ‘lateral

tubercles in this variety begin to grow at a much earlier stage, and the dorsal edge is

somewhat narrower than in many Pondicherry specimens; further, the siphuncle often lies

nearer to one edge than to the other; this, however, is not constant in all specimens’. The

last statement is difficult to reconcile with a fastigiate venter. Kossmat (1895: 157 (61),

pl. 19(5) (figs 15, 16)) figured, as Baculites vagina var. ootacodensis, a specimen from

Otacod, and one from Rautankupam (Pondicherry). The latter is a large Eubaculites

carinatus. The former is distinctly binodose but is damaged and lacks the venter

(Fig. 7F—H); we presume it to be a near-adult E. labyrinthicus but it could be a variant of

E. carinatus. Most subsequent authors took Kossmat’s figures to represent the species,

200 ANNALS OF THE SOUTH AFRICAN MUSEUM

assuming it to have a tabulate venter. As indicated in the synonymy, most of these

citations refer to Eubaculites carinatus.

Spengler (1923) described three baculitid species from Assam. The first, Baculites

vagina (Spengler 1923: 54, pl. 4 (fig. 9)) from Tharia Ghat, is a typical representative of

E. labyrinthicus. The second species from Assam (below Maomluh), described and fig-

ured by Spengler (1923: 54—55, pl. 4 (fig. 2a—b)) as Baculites sp. (cf. vagina Forbes var.

simplex Kossmat), has perfectly smooth flanks, a compressed whorl section, and a nar-

rowly rounded venter. This is similar to the lectotype of Eubaculites simplex, but for the

shape of the venter. We are not sure if this is a Eubaculites at all; it may be a Baculites

sensu stricto. The third species of Spengler (1923: 55, pl. 4 (fig. 8)) from Tharia Ghat,

described and figured as Baculites cf. B. anceps Lamarck, has an inflated whorl section,

but Spengler noted that ‘The siphonal side is keel-shaped’. Again, we are not sure if it

indeed is a Eubaculites at all—if it is, it might be a smooth variant of E. vagina or some

other species.

Baculites rioturbioensis Hiinicken, 1965 (p. 58, pl. 2 (figs 1-2), pl. 3 (figs 7-8),

pl. 7 (fig. 1), pl. 8 (figs 6—-8)) is from Santa Cruz Province, where Hiinicken tentatively

dated it as uppermost Campanian or basal Maastrichtian. Baculites rioturbioensis has a

trigonal whorl section with a flattened dorsum and fastigiate venter as in E. labyrinthicus.

In some specimens of B. rioturbioensis, Htinicken (1965: 59) noted signs of depressions

on either side of the siphuncle—possibly indications of an incipient ventral keel. Major

differences between B. rioturbioensis and E. labyrinthicus are the lateral ornament. In

B. rioturbioensis the flanks are ornamented by low, broad folds, in contrast to the gener-

ally well-defined, crescentic binodose flank ribs of E. labyrinthicus.

Baculites argentinicus Weaver (1927: 429, fig. 2) is probably a senior synonym of

B. rioturbioensis, but is based on poorly preserved material and best regarded as a nomen

dubium. Hiinicken & Covacevich (1975: 146, pl. 2 (figs 1-3), text-fig. 5a—b) described,

as Baculites? sp. B, a form with a fastigiate venter and low ribs from Bahia Las Tablas,

Quiriquina Island. This looks very much like E. /abyrinthicus. It occurs at the same

stratigraphic level (Nivel 12) as E. carinatus (= B. lyelli).

The lectotype of Eubaculites vagina var. simplex Kossmat (1895, pl. 19(5) (fig. 13))

(Fig. 7I-K), by subsequent designation of Kennedy (1986c: 195), has a fastigiate

venter like that of the lectotype of E. labyrinthicus, but lateral ornament is very faint,

and the whorl section is not as inflated as that of typical E. ootacodensis. Kennedy

(1986c: 194) regarded it as a synonym of E. ootacodensis (= E. labyrinthicus herein),

but subsequent work on Australian faunas (Henderson et al. 1992) showed E. simplex

to be a distinct, very compressed species (whorl breadth to height ratio 0,46—0,70) with

a tear-shaped to pyriform whorl section, with or without a distinct keel, the venter

being fastigiate or narrowly tabulate. Smooth variants of E. labyrinthicus (e.g. Fig. 4),

differ from E. simplex mainly in having a more inflated whorl section, but juvenile

specimens are difficult to separate satisfactorily. The other specimen referred to

E. simplex by Kossmat (1895, pl. 19(5) (fig. 14)) has an inflated whorl section, prominent

lateral ribbing and indications of a tabulate keel; it is best referred to E. carinatus, as

suggested by Kennedy (1986c: 196). Eubaculites simplex has a more compressed whorl

section and smooth flanks.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 201

A B C

Fig. 6. Eubaculites labyrinthicus (Morton, 1834). The original of Collignon’s (1971, pl. 645

(fig. 2388)) E. simplex from locality 666, Antsoha (Belo sur Tsiribihina), Madagascar. This specimen

shows prominent development of ‘E. ootacodensis’ type of ribbed ornament on flanks in lieu of

bituberculate as in typical E. labyrinthicus. x 1.

202

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 7. A-E. Eubaculites carinatus (Morton, 1834). Paralectotype of Eubaculites

simplex (Kossmat, 1895) the original of Kossmat, 1895, pl. 19 (5) (fig. 14) from

Ootacod. GSI 14820. F-H. Eubaculites labyrinthicus (Morton, 1834). Cast of

original of Kossmat, 1895, pl. 19 (5) (fig. 15) from Ootacod. GSI 14821.

I—K. Eubaculites simplex (Kossmat, 1895). Cast of lectotype, the original of

Kossmat, 1895, pl. 19(5) (fig. 13), from Arrialoor; GSI 14819. All specimens in

the collections of the Geological Survey of India. All x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 203

Occurrence

Maastrichtian, Prairie Bluff Chalk in Alabama, Navesink Formation in New Jersey,

U. S. A. Maastrichtian near Ootacod, South India. Locality 20 of Kennedy & Klinger

(1975: 283), Beds 3 (C) and 7 (G) of Klinger (1976, fig. 13), St Lucia Formation, Maas-

trichtian II, Zululand. Lower Maastrichtian, zone of Pachydiscus gollevillensis and

P. neubergicus of Madagascar (as Baculites simplex).

Eubaculites vagina (Forbes, 1846)

Figs 8—20

Baculites vagina Forbes, 1846a: 144, pl. 10 (fig. 4a—c). D’Orbigny, 1850: 215. Stoliczka, 1866: 198

(pars), pl. 91 (figs 1-6), non pl. 90 (fig. 14) (= E. labyrinthicus); non pl. 90 (fig. 15) (= E. cari-

natus). Crick, 1898: 78, pl. 17 (fig. 5). Diener, 1925: 63 (pars).

Baculites ornatus d’Orbigny, 1847: pl. 3 (figs 3-6).

Baculites vagina Forbes. a) Typische Form Kossmat, 1895: 155 (50), pl. 19 (5) (fig. 17).

non Baculites vagina Forbes. Forbes in Darwin, 1846b: 126, pl. 5 (fig. 3) (= E. carinatus).

non Baculites vagina Forbes. Kossmat, 1895: pl. 19(5) (fig. 14) (= E. carinatus).

non Baculites vagina var. Cazadorana Paulcke, 1907: 11, pl. 16 (fig. 5—5b). (= Baculites sp.).

non Baculites vagina Forbes. Boule, Lemoine & Thévenin, 1907: 45(65), pl. 8(15) (fig. 3) (= Bacu-

lites sp.).

non Baculites sp. cf. vagina Forbes. Woods, 1917: 36, pl. 20 (fig. 5a—d) (= ?B. rectus).

non Baculites vagina Forbes. Spengler, 1923: 54, pl. 4 (fig. 9) (= E. labyrinthicus).

non Baculites cf. vagina var. otacodensis Stoliczka. Crick, 1924: 140, pl. 9 (figs 4-5) (= ?E. carinatus).

non Baculites cf. vagina var. simplex Kossmat. Crick, 1924: 140, pl. 9 (figs 6-7) (= ?E. carinatus).

non Baculites vagina Forbes. Crick, 1924: 139, pl. 9 (figs 1-3) (= E. carinatus).

non Baculites vagina Forbes. Spath, 1926: 80.

non Baculites vagina Forbes. Wetzel, 1930: 90, pl. 10 (figs 3-4) (= E. carinatus).

non Baculites vagina Forbes. Basse, 1931: 20, pl. 2 (figs 6-10) (= ?Baculites sp.).

non Baculites vagina Forbes var. Van Hoepeni Venzo, 1936: 116(58), pl. 10(6) (figs 11-12) (= Baculites

vanhoepeni).

non Eubaculites vagina (Forbes). Spath, 1940: 49, text-fig. la (= E. carinatus).

non Eubaculites vagina (Forbes): Brunnschweiler, 1966: 29, pl. 1 (fig. 7), pl. 2 (figs 1-14), text-figs

12-14 (= E. carinatus).

Eubaculites vagina (Forbes): Collignon, 1971: 15, pl. 645 (fig. 2391), p. 18, pl. 646 (fig. 2394). Klinger,

1976: 87, pl. 35 (figs 1-4), pl. 36 (figs 1-4), pl. 37 (figs 1-5), pl. 38 (figs 1-3, ?fig. 4), pl. 39

(fig. 2), pl. 42 (fig. 1, ?fig. 4), pl. 43 (figs 5-12). Kennedy, 1977, text-fig. 31 (1-2). Klinger &

Kennedy, 1980: 299, fig. 5a—c. Kennedy 1986c: 194. Kennedy & Henderson, 1992: 712, pl. 5

(figs 10-13), pl. 7 (figs 1-18), pl. 8 (figs 1-12), pl. 9 (figs 1-13), pl. 10 (figs 1-3), text-fig. 1C, F.

Eubaculites Kennedy, 1989, fig. 17i-}.

Type

The lectotype of E. vagina is the original of Forbes (1846a, pl. 10 (fig. 4)) from the

Valudavur Beds of Southern India, BMNH C49762 (herein Fig. 8), designated by

Kennedy & Henderson (1992: 713). Klinger (1976: 88) referred to it as “holotype’—this

is not considered a valid lectotype designation.

Diagnosis

Eubaculites with tabulate siphonal keel and bituberculate lateral ornament; some rare

variants may be smooth.

204 ANNALS OF THE SOUTH AFRICAN MUSEUM

A B Cc

Fig. 8. Eubaculites vagina (Forbes, 1846). A-C. BMNH C49762, the lectotype, the original

of Forbes (1846, pl. 10 (fig. 4)); Geological Society Collection R10488. From the Valudavur

Formation of southern India. x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 205

Fig. 9. Eubaculites vagina (Forbes, 1846). A-C. BMNH C77593. Adult specimen with typical

bituberculate lateral ornament. From the Valudavur Formation of southern India. x 1.

ANNALS OF THE SOUTH AFRICAN MUSEUM

206

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CRETACEOUS FAUNAS FROM SOUTH AFRICA

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208 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

Full descriptions of the species are given by Klinger (1976) and Kennedy &

Henderson (1992) and need not be repeated. As yet, no adult specimens of E. vagina have

been found in Zululand, and, in retrospect, we suspect that the juvenile specimens referred

to this species by Klinger (1976, pl. 42 (fig. 4)) (herein Fig. 52A—C) are better referred

to E. labyrinthicus.

Discussion

One of us (Klinger 1976) previously described and figured syn- and topotype

material from South India, and described the variation shown by this species. Due to

unforeseen circumstances, the original figures were reduced so as to be practically

unrecognizable. These specimens are here refigured (Figs 8-15).

Eubaculites vagina 1s unique in the genus Eubaculites, in having two rows of lateral

tubercles combined with a tabulate ventral keel. Specimens with distinct bituberculate

ornament, here regarded as being typical of E. vagina, include BMNH C77593 (Fig. 9),

C51141 (Fig. 1O0D-F) and C51143 (Fig. 10A-—C), C77594 (11A-C), and C2583

(Fig. 11D-—-F). Some specimens are nearly smooth, with no lateral ornament, e.g. C51142

(Fig. 1SA—C), C51145 (Fig. 15I-J)) and the specimen figured by Stoliczka (1866, pl. 91

(figs 1—2)) (herein Fig. 16). We are not quite sure if these are, in fact, smooth E. vagina, or

Eubaculites latecarinatus; we think E. vagina, as they occur with typical representatives of

that species. Also, typical populations of E. latecarinatus are, as yet, unknown from South

India. BMNH C51140 (Fig. 13) and the specimen figured by Stoliczka (1866, pl. 91

(fig. 4)) (herein Fig. 19) differ from other specimens of E. vagina in their denser

and more oblique, though weaker lateral tuberculation. This may possibly be a

feature of the adult body chamber. The differences in size between these specimens and

the lectotype, which also appears to be adult, may be an indication of dimorphism. How-

ever, the largest known specimens of E. vagina are all significantly smaller than the

largest E. carinatus and E. latecarinatus. We are not sure if this is a consistent feature or

only due to the relatively small number of adult E. vagina known.

Until now, unequivocal adult bituberculate shells of E. vagina have only been found

in South India and in Madagascar—the doubtful Zululand specimens reported by Klinger

(e.g. 1976, pl. 42 (fig. 4)) are juveniles and better referred to E. labyrinthicus. Collignon

(1971: 15, pl. 645 (fig. 2391), p. 18, pl. 646 (fig. 2394)) (herein Fig. 20) recorded E. vagina

from the “Lower? Maastrichtian of Madagascar, Zone a Pachydiscus gollevillensis et

P. neubergicus. This is true E. vagina, unlike the totally incorrect interpretation of the

species by Boule er al. (1907: 65, pl. 8(15) (fig. 3)) and Basse (1931: 20) (fide Spath

1953: 46). These Madagascan specimens appear to be true Baculites. The Australian speci-

mens referred to E. vagina by Brunnschweiler (1966: 29, pl. 1 (fig. 7), pl. 2 (figs 1-14),

text-figs 12-14) are all typical representatives of E. carinatus.

The specimen figured by D’Orbigny (1847, pl. 3 (fig. 3)) as Baculites ornatus is a

composite synthetograph. Nevertheless, it clearly shows typical E. vagina bituberculate

lateral ornament. Stoliczka (1866: 199) commented that “B. ornatus . . . probably is from

Pondicherry itself’. The fact that the vast collections from the Quiriquina Formation of

Stinnesbeck (1986: 207) yielded only E. lyelli but no E. vagina (ornatus) confirms

Stoliczka’s suspicions. Hiinicken & Covacevich (1975: 159-160) discussed the

CRETACEOUS FAUNAS FROM SOUTH AFRICA 209

Fig. 12. Eubaculites vagina (Forbes, 1846). A-—C. BMNH C51146. Body chamber fragment

with part of the aperture preserved. D-F. BMNH C51147. Specimen with prominent

bituberculate ornament. Both specimens from the Valudavur Formation of southern India.

Both x 1.

210

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 13. Eubaculites vagina (Forbes, 1846). A-C. BMNH C51140, part of body chamber, from

the Valudavur Formation of Pondicherry, southern India. x 0,75.

211

CRETACEOUS FAUNAS FROM SOUTH AFRICA

InNAephyeA 94} WO, ae suouttoads IV

‘T X [IV ‘eIpuy usoyjnos Jo uoneuroy

UOHeLeA syIosdsenuI puke ojeuasojuO Jo asuLI SuNeIsNT[I suowttoeds Jo soiias V

69S¢L) HNING “O-W

‘SPITSO HNING “Tf 69S€L0 HNING IHD “prlISO HNN ‘4-d ‘6rIISO HN O-V “(9Or8I ‘seqiog) vuI3na sanynovqny “py ‘3tq

DD: ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 15. Eubaculites vagina (Forbes, 1846). A-C. BMNH C51142, a smooth form of E. vagina showing

fine striae on shell and smooth surface on internal mould. D-E. BMNH C51150. F-H. BMNH C77599,

a body chamber fragment with crowded ornamentation. I-J. BMNH C51145, another smooth form. All

specimens are from the Valudavur Formation of southern India. All x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 213

A B Cc D

Fig. 16. Eubaculites vagina (Forbes, 1846). A smooth, non-ornate variety of

the species. Cast of the original of Stoliczka, 1866, pl. 91 (fig. 2), from Pondicherry.

Collection of the Geological Survey of India, GSI 408. x 1.

systematic position of B. ornatus, stating that the type of B. ornatus could not be found in

the D’Orbigny collections, but concluded that (1975: 160) it was a synonym of E. vagina.

Eubaculites vagina has been reported from the Ukrainian SSR (see Kennedy 1986a:

1015; 1986c: 194) on the basis of a crushed baculitid from the Maastrichtian flysch

described and figured by Wisniowski (1907: 199, pl. 17 (fig. 9)) as Baculites anceps.

Admittedly, the specimen does show bituberculate lateral ornament, but this is due to a

longitudinal fracture and subsequent displacement of ribbing (°. . . dargestellten Gehduse,

welches leider in der Langsrichtung zerbrochen ist und infolgedessen eine kleine Ver-

schiebung beider Teile erlitt?-—Wisniowski 1907: 199).

Baculites labyrinthicus Morton (1834: 44, pl. 13 (fig. 10); see above) also has bitu-

berculate flank ornament but a fastigiate rather than tabulate venter. Baculites binodosus

Noetling, 1897 (p. 77, pl. 23 (fig. 3)) (herein Fig. 56) is based on a single fragment; it has

a single feeble row of dorsolateral tubercles and a feebly tabulate venter. It may represent

a further distinct species of Eubaculites, probably close to E. simplex.

Occurrence

Maastrichtian of South India and Madagascar; only dubious juveniles, best referred

to E. labyrinthicus, are known in Zululand.

ANNALS OF THE SOUTH AFRICAN MUSEUM

214

OOM EE

(OW a rep eee BE 2

repo erattn

capt

pari

Fig. 17. Eubaculites vagina (Forbes, 1846). A—D. Cast of original of Stoliczka, 1866, pl. 91

E-I. Cast of original of Stoliczka, 1866, pl. 91 (fig. 3). GSI 410.

(fig. 1), part, GSI 408.

J—M. Cast of original of Stoliczka, 1866, pl. 91 (fig. 1), part, GSI 408. All from Pondicherry,

Collection of the Geological Survey of India. x 1.

Dales

CRETACEOUS FAUNAS FROM SOUTH AFRICA

ines mcenpeptettittimenemneageette

LORE

fig. 5)

herry;

5 fol Dil ©

4) from Pondic

1866

GSI 411. Both in the collections of the Geological Survey of India. x 1.

inal of Stoliczka

rig

ka, 1866, pl. 91 (fig.

A-C. Cast of o

inal of Stolicz

1846).

Fig. 18. Eubaculites vagina (Forbes,

from Pondicherry; GSI 412. D. Cast of or

ANNALS OF THE SOUTH AFRICAN MUSEUM

216

peter

sheiercectcecactnonnnnannennmntn tere

porter

PTE

—_—

oo UO

=o!

os a4

ae,

~ 0

St tS

Gay

o= Of

Re)

—

en, oe

2 0

O) we

10ns O

, 1846).

(Forbes

ites vagina

pl. 91 (fig. 4) from Pondicherry. Collect

Fig. 19. Eubacul

GSI 411.

CRETACEOUS FAUNAS FROM SOUTH AFRICA De),

Fig. 20. Eubaculites vagina (Forbes, 1846). A-—C. The original of Collignon (1971, pl. 645

(fig. 2391)) from locality 504, Mont Ambinda-Mikoboka (Manera), Madagascar. D-E. The

original of Collignon (1971, pl. 646 (fig. 2394)) from locality 671, Antsoha (Bela Sur

Tsiribihina), Madagascar. All x 1.

218 ANNALS OF THE SOUTH AFRICAN MUSEUM

Eubaculites carinatus (Morton, 1834)

Figs 7A-E, 21-30, 31A—G, 32-35, 36E-F, 37-38, 42A, 52G—H

Baculites carinatus Morton, 1834: 44, pl. 13 (fig. 1). Stephenson, 1955: 134, pl. 24 (figs 5-9).

Baculites vagina Forbes: Darwin, 1846: 126. Forbes in Darwin, 1846b; pl. 5 (fig. 3). Steinmann, 1895:

89, pl. 6 (fig. 4), text-figs 8-10. Kossmat, 1897), pl. 6 (fig. 4). Wilckens, 1904: 188. Diener, 1925:

63 (pars). Wetzel, 1930: 90, pl. 10 (figs 3-4). Hoffstetter et al., 1957: 300, 302.

Baculites Lyelli d’Orbigny, 1847, pl. 1 (figs 3-7); 1850: 215. Olsson, 1944: 104, pl. 16 (figs 3-5), text-

cra

Baculites tippaensis Conrad, 1858: 334, pl. 35 (fig. 27).

Baculites spillmani Conrad, 1858: 335, pl. 35 (fig. 24).

Baculites anceps Lamarck. Binckhorst, 1861: 42, pl. 5d (fig. 3a—d).

Baculites chicoensis Gabb, 1864: 80 (pars), pl. 14 (fig. 29, 29a); non pl. 17 (fig. 27, 27a), non pl. 14

(fig. 27b).

Baculites vagina var. Otacodensis Stol.: Kossmat, 1895: 157(61) (pars), pl. 19(5) (fig. 16), (2non fig. 15

= ?E. labyrinthicus).

Baculites vagina var. simplex Kossmat, 1895: 156(60) (pars), pl. 19(5) (fig. 14 only).

non Baculites vagina var. nov. cazadorianus Paulcke, 1907: 11, pl. 16 (fig. 5—5b) (= Baculites sp.).

non Baculites vagina var. cazadoriana Paulcke: Diener, 1925: 63.

Baculites sheringomensis Crick, 1924: 139, pl. 9 (figs 1-3).

Baculites cf. vagina var. otacodensis Stoliczka: Crick, 1924: 140, pl. 9 (figs 4—5).

Baculites cf. vagina var. simplex Kossmat: Crick, 1924: 140, pl. 9 (figs 6-7).

Eubaculites otacodensis Stoliczka. Spath, 1940: 49, pl. 1 (fig. 3), text-fig. 1b. Wright, 1957: L218,

figs 245, 246a—c. Collignon, 1971: 18, pl. 646 (fig. 2395).

?Baculites anceps (Lamarck) d’Orbigny: Haas, 1943: 13 (figs 16-17 only).

Eubaculites lyelli (d’ Orbigny): Matsumoto & Obata, 1963: 97. Hiinicken & Covacevich, 1975: 149, pl. 1

(figs 5-12), pl. 2 (figs 4-9), pl. 3 (figs 1-8), pl. 4 (figs 1-8), pl. 5 (figs 1-4), text-figs 6-28. Stin-

nesbeck, 1986: 207, pl. 9 (figs 6-8), text-fig. 24D, d. Kennedy & Summesberger, 1986: 197, pl. 14

(figs 1-5, 9-14). Kennedy, 1986a: 1016, pl. 1 (figs 1-3), pl. 2 (figs 3-8), pl. 3 (figs 2-8, 13-21).

Kennedy, 1986), fig. 10c, d. Kennedy, 1986c: 195, pl. 27 (figs 5-8), pl. 32 (figs 13-14). Immel,

ISTE WA,

Eubaculites argentinicus (Weaver): Leanza, 1964: 95, pl. 1 (figs 1-5), text-fig. 1.

Eubaculites sp. Garcia & Camacho, 1965: 72, pl. 1 (fig. 4).

Eubaculites vagina (Forbes): Brunnschweiler, 1966: 29, pl. 1 (fig. 7), pl. 2 (figs 1-14), text-figs 12-14.

Eubaculites ootacodensis (Stoliczka): Brunnschweiler, 1966: 27, pl. 1 (figs 9-14), text-figs 9-11. Matsu-

moto, 1959: 166, pl. 43 (fig. 6), pl. 44 (figs 1-3), text-figs 84a—b, 85a—b. Riccardi, 1974: 388, pl. 1

(figs 1-7), pl. 2 (figs 1-4, 6), pl. 3 (figs 1-6), pl. 4 (figs 1-7), text-fig. 2; 1980: 11-14, figs 1-2.

Klinger, 1976: 90 (pars), pl. 39 (fig. 1, non fig. 3), pl. 41 (figs 1-2), pl. 42 (figs 3, 8), non pl. 43

(fig. 1), text-fig. llc.

Eubaculites kossmati Brunnschweiler, 1966: 31, pl. 2 (figs 15-17), pl. 3 (figs 1-7), text-fig. 15.

Eubaculites multicostatus Brunnschweiler, 1966: 32, pl. 3 (figs 8-12), text-fig. 16.

Eubaculites carinatus (Morton, 1834): Henderson et al., 1992: 150, figs 6F—G, 14-16, 17A—C, G-—J,

18-20. Kennedy & Henderson, 1992: 716, text-fig. SA—C.

Type

Holotype, by monotypy, is ANSP 72866 in the collections of the Academy of Natu-

ral Sciences, Philadelphia, the original of Morton (1834, pl. 13 (fig. 1)), from the Prairie

Bluff Chalk of Alabama (Fig. 21). The lectotype of Baculites lyelli d’Orbigny, 1847, by

subsequent designation by Kennedy (1986a: 1016) is MNHP R1020a (refigured by Ken-

nedy 1986a, pl. 1 (figs 1-3)) (herein Fig. 22F—G); paralectotypes are MNHP R1020b—c

(formerly D’Orbigny collection no. 7206) (herein Fig. 22D—E). These specimens are pre-

sumably the basis of D’Orbigny’s highly idealized figures (1847, pl. 1 (figs 3—7)). The

locality is given as ‘Conception’, but it is more likely Quiriquina Island, as also later

stated by D’Orbigny’s (1850: 715) Prodréme entry.

— Lee

CRETACEOUS FAUNAS FROM SOUTH AFRICA 219

Material

We have numerous specimens, including SAS H54/11 from locality 132, degraded

cliff and foreshore platform 300 m north-north-east of the northern jetty at Charter’s

Creek Rest Camp, St Lucia Formation, Maastrichtian I; SAM-—PCZ7070, PCZ7073,

PCZ7427, PCZ7442, PCZ7429, PCZ7788, PCZ8317—8330, SAS H54/11, SAS Z2204

from locality 134, cliffs and foreshore section 1,2 km south of Charter’s Creek Rest

Camp, St Lucia Formation, Maastrichtian I; SAM—PCZ7047, PCZ8651, SAS H63/12,

H63/1, H61/20, OUM KX1835-1838, SAS Z2204 from an unspecified horizon at local-

ity 133; H60F/2, SAS H60F/14, H60H/17, Bed F; SAS H60H/17, Bed H, locality 133,

cliff section and beach platform below Charter’s Creek Rest Camp, St Lucia Formation,

Maastrichtian I; PCZ8831 from locality 125, foreshore platform north of Fanies Island,

eastern shores of the southern peninsula, Maastrichtian a (ex Campanian IV);

SAM-PCZ8647, PCZ8650, SAS H48/1, SAS L97a—b from locality 131, low cliffs and

foreshore sections 3, 1 km north of Charter’s Creek Rest Camp, St Lucia Formation,

Maastrichtian Il; SAM—PCZ8311, PCZ8648, loose at locality 20; SAS H163C/4, Bed 3

(C); H163D/1, OUM KX1834, Bed 4 (D); SAS H20/5, H163E/5, Bed 5 (E); SAS H163G,

OUM KX1825, KX1827, KX1839-1840, Bed 7(G) at locality 20, section at junction of

old course of the Mfolozi River, the present course and the unnamed stream draining

south from Lake Mfuthululu, St Lucia Formation, Maastrichtian [-—II.

Dimensions

Specimen Wb Wh Wb/Wh Tapering

SAS Z2204a 8 14 0,57 —

SAS H163D/1 10 16 0,62 a

1] 20 0,55 11,4 (D = 35)

SAM-PCZ7442 10 12) 0,53 —

SAM-PCZ8320 9 7 0.53 —

12 20 0,60 1,3 © = 55)

SAM-PCZ8322 12 19 0,63

10 14 0,71 12,5 (D = 40)

SAM-—PCZ8323 1] 7 0.65 —

10 14 0,71 8,8 (D = 34)

SAM-—PCZ8328 10 16 0,62

9 13 0,69

7 10 0,70 (20 (UD = 25)

SAM-—PCZ8330 9 13 0,69 —

SAM-PCZ8319 11 16 0,69 —

9 13 0,69 10 (D = 30)

SAM-PCZ8326 9 13 0,69

7 10 0,70 340) (UD) = 23)

SAM-—PCZ8329 9 13) 0,69

8 11 0,72 Lbs (ID) = 117)

SAM-PCZ8327 9 13 0,69

7 25 0,74 13,5 (D = 26)

220 ANNALS OF THE SOUTH AFRICAN MUSEUM

SAM-PCZ8321 9 14 0,64 —

SAS H63/1 21 42 0,50 =

24 47 ES) AS) (UD) = 102)

SAM-—PCZ7047 24 43 0,59

19 34 0,56 10,6 (D = 85)

SAM-—PCZ7427 20 34 0,56 ——

14 25 0,56 8,1 (D = 94)

SAS 97 = 44

— 34 — 20) (DY = i)

SAS H60H/17 8 SII 0,58

Diagnosis

Eubaculites with tabulate siphonal keel and lateral ornament consisting of ribbing

only.

Description

The holotype of Eubaculites carinatus (Fig. 21) is a phosphatic internal mould of a

body chamber and parts of two camerae, 47,5 mm long, with a whorl breadth to height

ratio of 0,74. The whorl section is pyriform, with a flattened dorsum, narrowly rounded

dorsolateral margin and broadly rounded flanks, with greatest breadth below mid-flank.

The outer flanks are convergent, and separated by a narrow longitudinal groove from

narrowly rounded ventrolateral shoulders; the venter is narrow and bears a tabulate keel.

The rib index is 2,5. The ribs are broad, strong and concave, as wide as the interspaces,

extending from the dorsolateral margin to the middle third of the flank. On the ventral

third of the flank the ribs weaken, break down into riblets and striae and project strongly

forwards to the ventrolateral margin, where they flex back and are feebly convex. They

strengthen markedly at the ventrolateral shoulder and are coarse and distant on the venter,

which is markedly serrate in profile; ventral ribs are approximately twice as numerous as

those on the flank.

Typical Zululand representatives of this species are easily identified by the pynform

whorl section, tabulate venter and the crescentic flank ribs. There is, however, consider-

able variation in the strength of the flank ribs, rib index, and whorl breadth to height ratio.

The shell may grow to a great size (e.g. Figs 25, 26C, 36F)—the maximum whorl

height measured is 80 mm. At the smallest diameter available (PCZ8318) (Fig. 28B), the

whorl section is tear-shaped, with a narrow, rounded venter. Flattening of the keel only

takes place at whorl heights varying between 15 mm and 20 mm; development of the

typical tabulate keel takes place even later. Tapering is rapid in small specimens; large

specimens have essentially parallel ventral and dorsal sides and a very slow rate of taper

as can be seen in the list of dimensions.

Flank ornament typically consists of crescentic ribs, arising, and protruding above

the dorsal edge, most prominent at mid-flank, and, in adult specimens, disappearing near

the ventrolateral furrows demarcating the keel. The stage of appearance, strength and

density of ribbing is extremely variable as can be seen from the figured specimens.

Prominent flank ornament is best shown in PCZ7047 (Fig. 24A—C), whereas PCZ8651

(ex H63/1) (Fig. 27) merely shows lateral undulations, and is morphologically transitional

to E. simplex.

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Fig. 21. Eubaculites carinatus (Morton,

1834). The holotype, ANSP 72866, from

the Prairie Bluff Chalk of Alabama. x 1.

Fig. 22. Eubaculites carinatus (Morton, 1834). A—C. Unnumbered

specimen, presumably from India. D-—-E. Paralectotype of

Baculites lyelli d’ Orbigny, 1847, from Quiriquina Island, Chile;

MNHP R1020b-c. F-—G. Lectotype of Baculites lyelli

d’Orbigny, 1847, from Quiriquina Island, Chile; MNHP R1020a.

All specimens housed in the collections of the Muséum

National d’ Histoire Naturelle, Paris. x 1.

221

222 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 23. Eubaculites carinatus (Morton, 1834). SAM-PCZ8647 from locality 131,

Zululand, St Lucia Fm, Maastrichtian II. x OMS:

CRETACEOUS FAUNAS FROM SOUTH AFRICA 223

Fig. 24. Eubaculites carinatus (Morton, 1834). A—-C. SAM—PCZ7047 from locality 133,

Zululand, St Lucia Fm, Maastrichtian I. D. SAM—PCZ8322. E. SAM—PCZ8648. Both loose

from the top of the section at locality 20, Zululand, St Lucia Fm, Maastrichtian II.

‘TX log ‘ev uURTYOINSeRY ‘Wy BION] 1S “puR[N[NZ CZ] Aieoo] WoIy TEESZOd-WVS “A

] ueYoIsee] “Wy LION Ig “puRININZ “PET AITVo]T WOIJ 6POSZId-WWS “WV ‘(HEsT “UOMOW) smPULiD? SaiimoPand ‘SC ‘SIy

ANNALS OF THE SOUTH AFRICAN MUSEUM

224

CRETACEOUS FAUNAS FROM SOUTH AFRICA BOS

Fig. 26. Eubaculites carinatus (Morton, 1834). A. SAM-—PCZ7427, specimen with very weak

lateral ornament, transitional to E. latecarinatus, from locality 134, Zululand, St Lucia Fm,

Maastrichtian I. B. SAM—PCZ7073 from locality 134. C. SAM—PCZ8650 from locality 131,

Zululand, St Lucia Fm, Maastrichtian Il. All x 1.

ANNALS OF THE SOUTH AFRICAN MUSEUM

226

‘| X ‘T uenyommseryy ‘Wy elon] 1g “pueyn[nZ ‘ce AyI[eoo] WOIF

‘snpulap2aIv] “A OY euoNTsueNn AT[estsoToydsour “WOJ payuwoureuto APyeom (T/E9H X28) 1S98ZOd-WVS “(pEQ] ‘UOMOJ)) SNIDULIDI SalyNoDgny “LZ ‘Sty

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Fig. 28. Eubaculites carinatus (Morton, 1834). A. SAS H48/1 from locality 131, Zululand,

St Lucia Fm, Maastrichtian I. B. SAM—PCZ8318. C—D. SAM-—PCZ8319. E-F. SAM—PCZ8326.

G. SAM-—PCZ8324. H. SAM-—PCZ8329, all from locality 134, Zululand, St Lucia Fm, Maas-

trichtian II. I. SAS H163G from Bed 7 (G) at locality 20. J—-L. SAS Z2204a, specimen tran-

sitional to E. simplex from locality 133, Zululand, St Lucia Fm, Maastrichtian I.

M. SAM-—PCZ8320 from locality 134, Zululand, St Lucia Fm, Maastrichtian I. N. SAM-—

PCZ8323, from locality 134. All x 1.

ae,

228 ANNALS OF THE SOUTH AFRICAN MUSEUM

Parts of the suture line (from PCZ7047) are shown in Figure 38 and illustrated

photographically in Figure 24A—C.

Discussion

The ribbed Zululand Eubaculites were originally identified by Kennedy & Klinger

(1975: 281) and described as E. ootacodensis by Klinger (1976). However, as discussed

above, this is a synonym of E. labyrinthicus, which has binodose flanks and a fastigiate

venter at small diameter. In large specimens the lateral ornament of E. labyrinthicus and

some of E. carinatus 1s virtually identical, and they can only be distinguished by their distinct

fastigiate venter and tabulate ventral keels, respectively (compare e.g. Fig. 36A—D and

Fig. 35, specimens from Madagascar). In typical specimens, E. carinatus 1s easily identi-

fied in the adult stage by the conspicuous crescentic flank ribbing. Eubaculites vagina has

a similar, sometimes more angular whorl section, but ornament typically consists of two

distinct rows of tubercles—one at mid-flank and the other near the dorsolateral edge.

Eubaculites latecarinatus differs from E. carinatus, typically having smooth flanks. How-

ever, smooth specimens of E. carinatus do occur (see Kennedy & Summesberger 1986,

pl. 14 (figs 6-8)). Individually these are indistinguishable from E. latecarinatus, and

smooth variants of E. vagina, but populations are distinct enough.

We have a number of small specimens that are weakly ribbed to smooth but with

compressed whorl section, with Wb : Wh ratio of 0,4—0,5. Klinger (1976) referred these

compressed specimens to E. compressum Brunnschweiler, 1966. Eubaculites compres-

sum, as interpreted by Klinger (1976), included Brunnschweiler’s (1966) Eubaculiceras

and Cardabites species. This 1s a rather mixed bag containing costate (Eubaculiceras) and

non-costate (Cardabites) forms, some with tabulate venter, e.g. Eubaculiceras compres-

sum (sensu Brunnschweiler 1966), Cardabites tabulatus, and others with fastigiate venter,

e.g. Eubaculiceras fastigiatum and Cardabites scimitar. SAS H60F/2 (Klinger 1976,

pl. 43 (fig. 13)) shows the transition from a fastigiate venter at one end to tabulate at the

other. They all have in common a compressed whorl section with Wb : Wh ratio of less

than 0,5. Henderson et al. (1992) restudied the Australian material, and also regarded all

of these species as synonyms (plus Baculites sp. of Hiinicken 1965: 64, pl. 2 (figs 3-4),

pl. 3 (figs 5-6), pl. 5 (fig. 5), pl. 6 (figs 6-7), pl. 8 (figs 2-5) and Eubaculites simplex

Kossmat of Collignon (1971: 15, pl. 645 (figs 2388—2389)), for which they used the name

Eubaculites simplex Kossmat, 1895 (pars, p. 156, pl. 19 (fig. 13 non 14)). The lectotype

is shown in Figure 7I-K. The Australian material is very compressed, smooth or with

faint dorsal undulations and a narrowly fastigiate to narrowly tabulate venter.

Of all the species of Eubaculites recognized, E. carinatus has the longest strati-

graphic range, through much of the Lower and Upper Maastrichtian. Assemblages of

E. carinatus from different geographic regions and different stratigraphic levels all show

a very wide range of variation. The excellent illustrations by Htinicken & Covacevich

(1975) of the Quiriquina material (Fig. 37) and West Australian material, described and

figured by Brunnschweiler (1966) and Henderson et al. (1992), show the wide variation

in density and strength of nbbing of E. carinatus. Brunnschweiler (1966: 32, pl. 3

(figs 8-12), text-fig. 16) named the more densely ribbed forms E. multicostatus. Kennedy

& Summesberger (1986: 197, pl. 14 (figs 6-8)) figured an almost smooth individual of

E. carinatus (as E. lyelli) from the Maastrichtian of Neuberg, Steiermark, Austria.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 229

Fig. 29. Eubaculites carinatus (Morton, 1834). A-B. OUM KX1836. C-D. OUM KX1837.

E-G. OUM KX1835. All from locality 133, Zululand, St Lucia Fm, Maastrichtian I.

ANIL 3 2D,

Kossmat (1897b: 73) noticed that ‘The Chilean Baculites vagina is recognizable by

the fact that the swellings of the ribs are a little nearer to the siphonal part of the shell

than in the Indian type, for this reason the section appears to be more oval.’ We do not

know if Kossmat had Indian Eubaculites carinatus or E. ootacodensis in mind when

making this statement. Both Indian E. ootacodensis figured by Stoliczka (1866, pl. 90

(fig. 14)) and E. carinatus figured by Kossmat (1895, pl. 19(5) (fig. 16)) do show the ribs

to be most strongly developed near the dorsal edge, but we do not know if this is a

constant feature of Indian E. carinatus, for there are not enough illustrations of Indian

material to substantiate this view.

230 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 30. Eubaculites carinatus (Morton, 1834). A-D. OUM KX1839. E-H. OUM KX1840.

Both from Bed 7 (G), locality 20, Zululand, St Lucia Fm, Maastrichtian II. Both x 2.

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Fig. 31. A-G. Eubaculites carinatus (Morton, 1834). A. SAS H20/5 from Bed 5 (E) at locality 20,

Zululand, St Lucia Fm, Maastrichtian II. B. SAS H63/12 from locality 133,

Zululand, St Lucia Fm,

Maastrichtian I. C-D. SAS H60H/17 from Bed H at locality 133. E-G. SAS H54/11 from locality 132,

Zululand, St Lucia Fm, Maastrichtian I. HJ. Eubaculites latecarinatus (Brunnschweiler, 1966), SAS H61/13

from locality 133. All x 1.

ANNALS OF THE SOUTH AFRICAN MUSEUM

Diey)

‘7 X [I ueNYonseryy

‘uly BION] 1g ‘pueln{nz ‘Oz Aryeooy “(D) L Ped Wosy L7ZIXA WNO ‘A-V “(PEST ‘UOMOW) smUIADI saIynovgny “ZE “BIq

CRETACEOUS FAUNAS FROM SOUTH AFRICA 233

Fig. 33. Eubaculites carinatus (Morton, 1834). A-D. OUM KX1826. E-H. OUM KX1825. Both from

Bed 7 (G), locality 20, Zululand, St Lucia Fm, Maastrichtian II. Both x 2.

234 ANNALS OF THE SOUTH AFRICAN MUSEUM

si OIE

Oe “Hy

“

Fig. 34. Eubaculites carinatus (Morton, 1834). A-C. OUM KX1834 from Bed 4, locality 20,

Zululand, St Lucia Fm, Maastrichtian II. D-E. OUM KX1838, from locality 133, Zululand,

St Lucia Fm, Maastrichtian I. All x 2.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 235

Fig. 35. Eubaculites carinatus (Morton, 1834). The original

of Collignon’s (1971, pl. 646 (fig. 2395)) Eubaculites

otacodensis from locality 503-504, Soatana-Mikoboka

(Manera), Madagascar. Note similarity of lateral ornament to

that of large E. labyrinthicus, also from Madagascar in

Figure 36A—D.

As in E. latecarinatus, there is great variation in the absolute size of E. carinatus.

Some of the Argentinian specimens before us (Fig. 36F) are as large as the largest E. late-

carinatus we have seen.

The specimens from Mozambique identified by Crick (in Teale 1924: 139) as belonging

to the group of Baculites vagina include ribbed forms that probably belong to the present

species.

Eubaculites carinatus has not been previously recorded from Madagascar, but the

original of Collignon’s (1971) E. otacodensis in part belongs to the present species

aige3 5):

Occurrence

Of the known species of Eubaculites, E. carinatus has the widest geographic distri-

bution and is a good indicator for the Maastrichtian Stage. It first occurs in Zululand in

Maastrichtian a, associated with typical Lower Maastrichtian Pachydiscus neubergicus—

as it also does at Neuberg, Steiermark in Austria—and ranges to Maastrichtian II in

Zululand, and the upper Upper Maastrichtian in the Petites-Pyrénées in southern France,

as well as in coastal sections in the Biscay region. Eubaculites carinatus is abundant in the

Southern Hemisphere in the South Gondwanid Province, where it has been recorded from

South India, Western Australia, Zululand, Mozambique, Madagascar, possibly Angola

(Fig. 55SF—H), Chile and Argentina. In the Northern Hemisphere it is common in the Ow]

Creek, Prairie Bluff and Corsicana Formations in Missouri, Mississippi, Alabama, and

Texas, and is rare as remanié fossils at the base of the Palaeocene in New Jersey. It also

occurs, albeit not in such great numbers, in California, the Netherlands, south-east France,

northern Spain and Austria.

236 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 36. A-D. Eubaculites labyrinthicus (Morton, 1834). Specimen from locality 503, Soatana-

Mikoboka (Manera), Madagascar, donated by the late Gen. M. Collignon. E-F. Eubaculites

carinatus (Morton, 1834). Two specimens from Neuquén Basin, Argentina. Collections of the

Division Paleozoologia de Invertebrados, Museo de Ciencias Naturales, Universidad Nacional de

la Plata, Argentina; courtesy Dr A. C. Riccardi. All x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA Doe

Fig. 37. Eubaculites carinatus (Morton, 1834). Block of specimens from Quiriquina Island, Chile. Collection

of Museum fiir Naturkunde, Berlin. x 1.

238 ANNALS OF THE SOUTH AFRICAN MUSEUM

O 10

O 5

ee eee ce ee

Fig. 38. Eubaculites carinatus (Morton, 1834). Part of suture line of SAM—PCZ7047.

Eubaculites latecarinatus (Brunnschweiler, 1966)

Figs 26A, 39-41, 42B-C, 43-49, 50A, 53A

Eubaculites otacodensis (Stoliczka): Spath, 1940: 49 (pars), text-fig. Ic.

Giralites latecarinatus Brunnschweiler, 1966: 33, pl. 3 (figs 13-14), pl. 4 (figs 1-5), text-figs 17-18.

Giratites quadrisulcatus Brunnschweiler, 1966: 35, pl. 4 (figs 11-14), text-fig. 20.

Eubaculites ambindensis Collignon, 1971: 18, pl. 646 (fig. 2393).

Baculites occidentalis Meek: Collignon, 1971: 15, pl. 645 (fig. 2390).

Eubaculites latecarinatus (Brunnschweiler): Klinger, 1976: 91, pl. 40 (figs 1-2), pl. 41 (fig. 3), pl. 42

(figs 2, 6), pl. 43 (figs 3-4), text-fig. 1ld—e. Klinger & Kennedy, 1980: 296, figs 2-4, 5D. Hender-

son et al., 1992: 159, figs 22L—N, 23N-P.

Type

Holotype is the original of Brunnschweiler (1966, pl. 4 (figs 2-4)), CPC 2718 from

the nodule bed at the top of the Korojong Calcarenite; Western Australia.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 239

Material

Numerous specimens, including SAS H163D, Bed 4 (D), SAM—PCZ7938, Bed 7 (G)

at locality 20, section at junction of old course of the Mfolozi River, the present course

and the unnamed stream draining south from Lake Mfuthululu, St Lucia Formation,

Maastrichtian I-II]; SAM—PCZ7029, PCZ7041—7045, PCZ7048—7056, PCZ7064, PCZ7072,

PCZ7074, PCZ7172-7176, PCZ7291—7298, PCZ7975, PCZ7979, SAS H60, H60/1.2, H61/6,

H61/13, H62/5, all from locality 133; SAS H60F/11 from Bed F at locality 133; SAS H60H,

H60H/1, H60H/4, Bed H at locality 133, cliff section and beach platform below Charter’s

Creek Rest Camp, Lake St Lucia, Zululand, St Lucia Formation, Maastrichtian I; SAM-—

PCZ7038, PCZ7058—-7059, PCZ7061, SAS H54B/15, Bed B, from locality 132, degraded

cliff and foreshore platform 300 m north-east of northern jetty at Charter’s Creek Rest Camp,

Lake St Lucia, Zululand, St Lucia Formation, Maastrichtian I; SAM—PCZ7065—7066,

PCZ7068-—7069, PCZ7075, PCZ7429, PCZ7432, PCZ7709, PCZ7980, PCZ8320—8330,

PCZ8457-8458, SAS Z2204 from locality 134, cliffs and foreshore section, 1,2 km south

of Charters Creek Rest Camp, St Lucia Formation, Maastrichtian I; SAS L97 from

locality 131, low cliff and foreshore section 4,6 km north of Charter’s Creek Rest Camp,

St Lucia Formation, Maastrichtian Il; SAM—PCO5908-5914, all from locality C in the

Alphard Group, offshore, Cape Province, Maastrichtian ?a.

Dimensions

Specimen Wb Wh Wb/Wh Tapering

SAS 22204 19 28 0,68 —

SAS H60F/11 L235) IQ) 0,66 =

10 16 0,62 Yo7 (WD = 13)

SAS H61/6 18 DES) 0,7 —

SAS H61/13 17 27) 0,63 —

14 23 0,61 16 (D = 25)

SAS H62/5 18 26 0,69 —

13 19 0,68 14,2 (D = 49)

SAS H60/1.2 — SZ

4] 5,2 CD = Zi)

SAM-—PCZ7068 Zi) 44 0,61

SAM-—PCZ7069 29 49 0,59

24 44 0,54 oS) (D777)

SAS H54B/15 46 Vz 0,64

40 65 0,61 6,0 (D = 117)

SAM-PCZ7269 — 65

64 0,41 (D = 240)

SAM -PCZ7066 — 68 =

66 2,3 (D = 88)

SAM-—PCZ7980 = 90 ae

82 4.6 (D = 172)

SAS H163D 28 44 0,63

19 28 0,69 1226. (D125)

240 ANNALS OF THE SOUTH AFRICAN MUSEUM

SAS H60 <= 68 —

oF Srl CD= 2.90)

SAM-PCZ7298 == 88 —

Sy// 38) (UD) = ead)

Diagnosis

Eubaculites with tabulate ventral keel; no lateral ornament. May grow to a large size.

Description

Tapering in the early stages is rapid, but decreases with growth, such that large

specimens have venter and dorsum essentially parallel in side view. The whorl section in

the adult stage is distinctly pyriform with a broad, flat dorsum and a serrated, tabulate

keel. The flanks are slightly inflated and nearly parallel. Maximum width is at, or just

ventral of, mid-flank. A distinct longitudinal ridge at the middle of the dorsum is present

in some specimens. The flanks are completely smooth. The suture is minutely frilled with

phylloid foliole elements in the saddles.

Discussion

This species is characterized by its tabulate venter and smooth flanks. It is the

commonest Eubaculites in Zululand, and is conspicuous by virtue of its large size—the

estimated length of some specimens was in excess of 2 metres.

Specimens reach maturity at large and small whorl heights, indicating size-related

dimorphism—compare, e.g. Figures 39, 40 and 41. Unfortunately, complete apertures are

very rare in the available material; part of one is preserved in H60H/1 (Fig. 46).

Smooth individuals of E. vagina (e.g. Stoliczka 1866, pl. 90 (fig. 1); herein

Figs 15A—C, I-J, 16) and of E. carinatus (Kennedy & Summesberger 1986, pl. 14

(figs 6—-8)) occur and are inseparable from E. latecarinatus, apart perhaps from details in

whorl section. Populations of the species are easily distinguished, as discussed above.

The specimen figured by Collignon (1971: 15, pl. 645, (fig. 2390)) as Baculites

occidentalis Meek seems to be a juvenile specimen of Eubaculites simplex or E. latecari-

natus (Fig. 5SA—B).

We have been inclined to regard smooth and compressed Eubaculites, described by

Klinger (1976: 92, pl. 43 (fig. 2)), as E. compressum as variants of the present species,

but revision of the Australian material (Henderson et al. 1992) has shown E. latecarinatus

and these compressed forms, for which the earliest available name is E. simplex (Kossmat,

1895), to be morphologically and stratigraphically separated. Eubaculites simplex is much

more compressed than E. latecarinatus, may have feeble flank undulations or be smooth,

and has a fastigiate to narrowly tabulate ventral keel.

Occurrence

Eubaculites latecarinatus is most abundant in Maastrichtian I of Zululand, but

onshore extends into Maastrichtian II. It is imprecisely located in the Maastrichtian of the

offshore Alphard Group, but may appear as early as Maastrichtian a (ex Campanian IV).

In Madagascar it occurs in the Lower Maastrichtian Zone of Pachydiscus neubergicus and

P. gollevillensis. The occurrences in Western Australia can be dated no more precisely

than Maastrichtian.

241

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Fig. 39. Eubaculites latecarinatus (Brunnschweiler, 1966). A-C. SAS H163D from Bed 4 (D) at

locality 20, Zululand, St Lucia Fm, Maastrichtian [-II. x 1.

242

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 40. Eubaculites latecarinatus (Brunnschweiler, 1966). A—B. SAS H60H/4 from Bed H,

locality 133, Zululand, St Lucia Fm, Maastrichtian I. x 0,75.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 243

Fig. 41. Eubaculites latecarinatus (Brunnschweiler, 1966). A. SAS H60H from locality 133,

Zululand, St Lucia Fm, Maastrichtian I. B. SAS H54B/15 from Bed B at locality 132, Zululand,

St Lucia Fm, Maastrichtian I. A x 0,40; B x 0,85.

244 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 42. A. Eubaculites carinatus (Morton, 1834). SAS L97, specimen with weak lateral

ornament, transitional to E. latecarinatus, from locality 131, Zululand, St Lucia Fm,

Maastrichtian II. B—C. Eubaculites latecarinatus (Brunnschweiler, 1966). SAM—PCZ7432 from

locality 134, Zululand, St Lucia Fm, Maastrichtian I. Both x 1.

245

CRETACEOUS FAUNAS FROM SOUTH AFRICA

‘pueln[nZ

PET Aypeoo] Wos Pog

‘| X Mog ‘| uenyomserpy ‘Wy eion] 1S

LSvV8ZOd-WVS —

690LZOd-WVS “A ‘O-V (9961

JoflamyosuUNIg) SNjpUulADpIaIv] Saj1jnovqny

er “314

246 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 44. Eubaculites latecarinatus (Brunnschweiler, 1966). A. SAM—PCZ7068. B. SAM—PCZ8458, the

largest known specimen, slightly deformed, viewed from the venter to show the size of the crenulate,

tabulate venter. A—B. From locality 134, Zululand, St Lucia Fm, Maastrichtian I. C-D. SAM—PCZ7938;

from Bed 7 (G) at locality 20, Zululand, St Lucia Fm, Maastrichtian J-II. All x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA DAG,

Fig. 45. Eubaculites latecarinatus (Brunnschweiler, 1966). A—B. SAM—PCZ7975 from locality 133.

C. SAS H54B/15 from Bed B at locality 132. Both from Zululand, St Lucia Fm, Maastrichtian I.

A-B x 1; C x 0,85.

248 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 46. Eubaculites latecarinatus (Brunnschweiler, 1966). SAS H60H/1. Adult |

specimen with part of aperture preserved. Bed H, locality 133, Zululand, St Lucia |

Fm, Maastrichtian I. x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA

Peeetecetsernrceseresay,

AAR LEL EEA PURI LAAETLED DEPLETED OTE TRE

ea He

)

dge on dorsal side. Both from locality C (TBD 1336) on the offshore

1an fr

Fig. 47. Eubaculites latecarinatus (Brunnschweiler, 1966). A-C. SAM—PCO5909. D. SAM—PCO5908

note the prominent med

Alphard Group, southern Cape, South Africa. x 1.

ANNALS OF THE SOUTH AFRICAN MUSEUM

250

TL 2S NY

“SISUapUIquiD SajynIvgny S UOUsIT[OD Jo sdAyuds poimsiyun uy ‘q-D “seoseSepeyy ‘(esiouRy\]) PYOGOYIP-epurquiy joy ‘pOS Aipeoo, wor

sisuapulquip sayynovqngy (€6¢7 3) 9/9 Id ‘[L6[) S,UoUsTJOD Jo jeuIsuO oY, “q—-V “(996] ‘Jo[lomyosuUNIg) smpusD2aID] salyNovgny ‘gp ‘Btq

CRETACEOUS FAUNAS FROM SOUTH AFRICA DES |

O 10

_—

—_a =e —

Fig. 49. Eubaculites latecarinatus (Brunnschweiler, 1966). Parts of the suture line.

DSO ANNALS OF THE SOUTH AFRICAN MUSEUM

Eubaculites simplex (Kossmat, 1895)

Figs 7I-K, 50B, 51, 52D-—F, I-N, 53B, 54-55

Baculites vagina Forbes n. var. simplex Kossmat, 1895: 156, pl. 19(5) (figs 13a—b) (non figs 14a-c)

(= E. carinatus).

Baculites sp. Hiinicken, 1965: 64, pl. 2 (figs 3-4), pl. 3 (figs 5-6), pl. 5 (fig. 5), pl. 6 (figs 6-7), pl. 8

(figs 2—5).

Giralites simplex (Kossmat): Brunnschweiler, 1966: 34, pl. 4 (figs 6-10), text-fig. 19.

Eubaculiceras compressum Brunnschweiler, 1966: 36, pl. 4 (figs 15-17), pl. 5 (figs 1-3), text-fig. 21.

Eubaculiceras fastigiatum Brunnschweiler, 1966: 37, pl. 5 (figs 7-9), text-fig. 22.

Cardabites tabulatus Brunnschweiler, 1966: 38, pl. 5 (figs 12-15), text-fig. 23.

Cardabites scimitar Brunnschweiler, 1966: 38, pl. 5 (figs 16-21), text-fig. 24.

Eubaculites compressum (Brunnschweiler): Klinger, 1976: 92, pl. 42 (fig. 10a—b), pl. 43 (figs 2a-—c,

13a—c, 14a-c).

Eubaculites simplex (Kossmat): Henderson et al., 1992: 153, 21, 22A—K, O-P, 23A—F, K—M, 24-25.

Kennedy & Henderson, 1992: 717, text-fig. 6D-F.

Types

Lectotype, designated by Kennedy (1986c: 195), is the original of Kossmat (1895,

pl. 19 (figs 13a—b)), Geological Survey of India Type Collection, GSI 14819 (Fig. 7I-K).

The original of Kossmat (1895, pl. 19 (figs 14a—c)) 1s GSI 14820 in the same collections,

and is a Eubaculites carinatus. Both are from the Arrialoor Group of Otacod, South India.

Material

SAS H163C/3 (Bed C), H163D/3 (Bed D), H163F/2 (Bed F), and SAM—PCZ7939

and PCZ7912 (Bed G) at locality 20, section at junction of the old course of the Mfolozi

River, the present course, and the unnamed stream draining south from Lake Mfuthululu,

St Lucia Formation, Maastrichtian I-II]; SAS H66/2, H66/3, from locality 135, foreshore

A B

Fig. 50. Comparison of whorl sections of large examples of Eubaculites

latecarinatus and E. simplex. A. Eubaculites latecarinatus (Brunnschweiler,

1966), SAM-—PCZ7979. B. Eubaculites simplex (Kossmat, 1895), SAM—PCZ7939.

Both x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA US)

Fig. 51. Eubaculites simplex (Kossmat, 1895). A. SAS H163D/3 from Bed 4 (D). B. SAS Z2204m.

C-D. SAS H163C/4 from Bed 3 (C). E-F. SAM—PCZ7939 from Bed 7 (G). All from

locality 20, Zululand, St Lucia Fm, Maastrichtian I-II. All x 1.

D354 ANNALS OF THE SOUTH AFRICAN MUSEUM

K L M N

Fig. 52. A-—C. Eubaculites labyrinthicus (Morton, 1834). SAS H163G/7 from Bed 3 (C) at locality 20,

Zululand, St Lucia Fm, Maastrichtian I-II. D-—-F, I-N. Eubaculites simplex (Kossmat, 1895).

D. SAS H66/2 from locality 135, Zululand, St Lucia Fm, Maastrichtian I. E-F. SAS H60F/12

from Bed F at locality 133, Zululand, St Lucia Fm, Maastrichtian I. I-J. SAS 163F/2 from

Bed 6 (F) at locality 20. K-N. SAS Z2204c from locality 133. G—-H. Eubaculites carinatus

(Morton, 1834). G. SAM-—PCZ8311, loose from locality 20. H. SAS H163D/1 from Bed 4 (D)

at locality 20. All x 1.

CRETACEOUS FAUNAS FROM SOUTH AFRICA DS)

Fig. 53. A. Eubaculites latecarinatus (Brunnschweiler, 1966). SAM—PCZ7980 from locality 134,

Zululand, St Lucia Fm, Maastrichtian I. B. Eubaculites simplex (Kossmat, 1895), SAM-—

PCZ7980, a body chamber fragment with part of the aperture preserved; from Bed 7 (G) at

locality 20, Zululand, St Lucia Fm, Maastrichtian I-II. A. x 0,45; B. x 0,70.

256 ANNALS OF THE SOUTH AFRICAN MUSEUM

ita

Fig. 54. Eubaculites simplex (Kossmat, 1895). A. Large specimen, SAM—

PCZ7980 (see also Fig. 53B) showing details of aperture, from Bed 7 (G) at

locality 20, St Lucia Fm, Maastrichtian II.

CRETACEOUS FAUNAS FROM SOUTH AFRICA

75)

F G H

Fig. 55. A-E. Eubaculites simplex (Kossmat, 1895). The original of Collignon’s (1971, pl. 645

(fig. 2390)) Baculites occidentalis from locality 665-7, Route Antsoha-Andranomana

(Ambavaloza) (Belo sur Tsiribihina), Madagascar. x 1. C—E. SAS H60A/21, specimen showing

transition from fastigiate to tabulate venter from locality 133, St Lucia Formation,

Maastrichtian Il. F—H. Eubaculites? sp., specimen from Angola figured by Haas (1943, fig. 16)

as Baculites anceps. Collections of the American Museum of Natural History, courtesy of

Dr N. Eldredge.

258

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 56. Eubaculites? binodosus (Noetling, 1897). Cast of holotype,

GSI 3097, of Baculites binodosus, the original of Noetling (1897,

pl. 23 (fig. 3)) from Dés Valley, Baluchistan.

outcrops in Makakatana Bay, east of the settlement, St Lucia Formation, Maastrichtian I;

SAS H60F/5, H60/2, H66F/15, H60F/21, all from Bed F at locality 133; SAS H60A/21

from Bed A; and SAS Z2204 from an unspecified horizon at locality 133; SAS Z2204c

and SAS H60HA/21, Bed A, locality 133, cliff section and beach platforms below Char-

ter’s Creek Rest Camp, Lake St Lucia, Zululand, St Lucia Formation, Maastrichtian I.

Dimensions

Specimen

SAS H60F/12

SAS H60/2

SAS H60F/21

SAS H60F/5

SAS H163F/2

SAM-—PCZ7939

SAM-—PCZ7912

Wb/Wh

0,53

0,56

0,44

0,60

0,46

0,56

0,50

0,53

Tapering

7,3 (D = 55)

CRETACEOUS FAUNAS FROM SOUTH AFRICA 259

A 8

Fig. 57. Baculites sp. (nov.?) aquilaensis

Reeside. The original of Collignon (1970,

pl. 639 (fig. 2358)) from the Upper

Campanian of locality 227-2, Mokotibe

(Antsalova) Madagascar. x 1.

Diagnosis

Compressed Eubaculites (Wb/Wh 0,46—0,70) with flanks that are smooth or with

broad, faint dorsal undulations. Whorl section tear-shaped to pyriform, with or without a

distinct keel, and venter fastigiate or narrowly tabulate.

Description

The lectotype (Fig. 7I-K) as represented by a cast, is 39 m long, with a maximum

preserved whorl height of 19,5 mm and a whorl breadth to height ratio of 0,64. It retains

recrystallized shell, and appears to be a fragment of body chamber; venter and dorsum are

parallel when viewed in profile. The dorsum is broad and very feebly convex. The dorso-

lateral margin is narrowly rounded, the dorsal two-thirds of the flank very broadly

rounded, the ventral third converging to a fastigiate venter, demarcated from the flanks by

a marked longitudinal depression. There is no ornament. Zululand material referred to this

Species consists predominantly of fragmentary juvenile specimens. These all show the

very compressed whorl section, and have generally no, or extremely weak flank ornament.

H60F/21 (Fig. 55C—E) shows the transition from a fastigiate venter at the smaller end to

tabulate at the larger. In all the other specimens, the venter is narrowly tabulate both with

and without crenulations.

Two large specimens, PCZ7939 (Fig. 51E-F)—part of the phragmocone—and

PCZ7980 (Figs 53B, 54A)—part of a body chamber with the aperture preserved, with

compressed whorl section—are possibly adults of the species. These indicate that E. sim-

plex may reach maximum sizes comparable to those of E. latecarinatus and E. carinatus.

Discussion

Henderson et al. (1992) referred an assemblage of 132 specimens from the Miria

Formation of Western Australia to this species. They described the shell as straight or

260 ANNALS OF THE SOUTH AFRICAN MUSEUM

with a slight endogastric curvature, and a very low expansion rate. The whorl section is

fairly to very much compressed, with a whorl breadth to height ratio of 0,46—0,7, the shell

surface smooth or with broad, weak undulations on the dorsal part of the flanks, 2-4 in a

distance equal to the whorl height. Dorsum and venter are completely smooth. The whorl

section is tear-shaped to weakly pyriform, the flanks broadly convex, dorsum feebly

convex, and venter fastigiate or with a tabulate keel which may be bordered on either side

by a longitudinal groove in some specimens. A collection of 27 specimens from a single

locality included eight with a clearly tabulate venter and eight with a rounded to fastigiate

venter that is narrowly arched in some but blade-like in others. Two specimens showed a

change from fastigiate to tabulate with growth. Shape of venter is not a function of growth

stage, as both large (whorl height in excess of 20 mm) and small specimens (whorl height

less than 20 mm) may have either tabulate or fastigiate venter.

Patagonian material described by Hitinicken (1965) as Baculites sp. and possessing a

keeled, fastigiate venter, is referred here to E. simplex. Baculites vagina var. cazadorianus

Paulcke (1907: 11, pl. 16 (figs 5, 5a, 5b)) may perhaps be conspecific, but its range of

variation is not known and the sole illustrated specimen is more like B. anceps in whorl

profile than E. simplex.

The lack of significant flank ornament, very compressed whorls, and fastigiate to

very narrowly tabulate venter separates populations of E. simplex from E. vagina, E. cari-

natus and E. labyrinthicus, described above. Smooth variants of E. carinatus and

E. labyrinthicus may be difficult to separate, as individuals, from smooth E. simplex with

tabulate venter, other than by their wider venters. Populations of E. simplex and E. late-

carinatus can be separated by the presence of individuals with fastigiate venters in the

former at a size where E. latecarinatus always has a tabulate venter, whereas this species

has a much wider venter than morphs of E. simplex with a tabulate venter. Differences in

whorl section in large specimens of E. simplex and E. latecarinatus are shown in

Figure 50.

Occurrence

Maastrichtian of South India, Western Australia, Patagonia, and Zululand (South

Africa). This species does not extend into the highest Maastrichtian.

ACKNOWLEDGEMENTS

Financial aid to Klinger from the Foundation for Research and Development, South

Africa, and to Kennedy from the Trustees of the Sir Henry Strakosh Bequest, Royal

Society, and Natural Environment Research Council (U. K.) is gratefully acknowledged.

We thank the staff of the Geological Collections, University Museum, Oxford, Depart-

ment of Earth Sciences, Oxford, and South African Museum, Cape Town, for technical

assistance. Dr A. Ayyasami of the Geological Survey of India supplied casts of Indian

type material, and the Academy of Natural Sciences of Philadelphia loaned Morton’s type

of Eubaculites for study. Dr W. A. Cobban (Denver) permitted us to study his North

American Eubaculites collections, Drs R. A. Henderson (Townsville), and K. J.

McNamara (Western Australian Museum) those from the Miria Formation, and Dr N.

Eldridge, American Museum of Natural History, some of the Angolan material described

CRETACEOUS FAUNAS FROM SOUTH AFRICA Doi

by Haas. D. Phillips, M. K. Howarth and H. G. Owen (London) assisted us with work on

Forbes’ specimens from South India in the collections of the Natural History Museum.

We also thank Dr W. A. Cobban and Prof. Emeritus T. Matsumoto for their con-

structive comments on the text.

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HERBERT CHRISTIAN KLINGER

WILLIAM JAMES KENNEDY

CRETACEOUS FAUNAS FROM ZULULAND

AND NATAL, SOUTH AFRICA.

THE HETEROMORPH AMMONITE GENUS

EUBACULITES SPATH, 1926

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FiscHER, P. H. 1948. Données sur la résistance et de la vitalité des mollusques. Journal de conchyliologie 88 (3): 100-140.

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

expérimentale et générale 74 (33): 627-634.

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

Magazine of Natural History (13) 2 (17): 309-320.

Koun, A. J. 1960b. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bulletin of

the Bingham Oceanographic Collection, Yale University 17 (4): 1-S1.

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

pologische Ergebnisse einer Forschungsreise im westlichen und zentralen Suid-Afrika ausgefiihrt in den Jahren

1903-1905 4 (15). Denkschriften der medizinisch-naturwissenschaftlichen Gesellschaft zu Jena 16: 269-270.

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 102 + Band

January 1993 Januarie

Part 7 Deel

/0yip NOVI We

ON THE AFFINITIES OF COBBANOSCAPHITES

COLLIGNON, 1969 (CRETACEOUS AMMONOIDEA)

WILLIAM JAMES KENNEDY

HERBERT CHRISTIAN KLINGER

Cape Town Kaapstad

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D1824

ON THE AFFINITIES OF COBBANOSCAPHITES COLLIGNON, 1969

(CRETACEOUS AMMONOIDEA)

WILLIAM JAMES KENNEDY

Geological Collections, University Museum, Oxford

HERBERT CHRISTIAN KLINGER

Department of Invertebrate Palaeontology, South African Museum, Cape Town

(With 2 figures)

[MS accepted 25 May 1992]

ABSTRACT

The genus Cobbanoscaphites Collignon, 1969, from the Lower Campanian of Madagascar, is not

a heteromorph ammonite of the superfamily Scaphitaceae, as originally described, but a pachydiscid

microconch and a synonym of Menuites Spath, 1922.

CONTENTS

PAGE

ITCCLICHIOR § cmb oy eects sd AOS See | GT ee ace 265

CODD TUTOR ce 8 os See A Se ae, cs 267

Syste malerpalacomlolOsy 6 con. fo ee se aes es ee Sea wed and 267

PG Kn Wile GOCIMCM Ser erie” = eat S UN ese Pek tans vic wk aye Ns 270

INGLE CMCC Sameer Wty te ia Fe Shia ye ee cys era OS See whe od 270

INTRODUCTION

In the course of a revision of the South African representatives of the heteromorph

ammonite superfamily Scaphitaceae Gill, 1871, the authors recently examined the Mada-

gascan representatives of the group described by Collignon (1963-1971), and currently

housed in the Département des Sciences de la Terre, Dijon. The most distinctive form

referred to the scaphites by Collignon was the genus Cobbanoscaphites Collignon, 1969,

originally described and diagnosed as follows:

COBBANOSCAPHITES MENABENSIS nov. gen. nov. sp. G. T. (1) D: 0,080.—H:

0,043. (0,52).—E: 0,055. (0,69).—O: 0,023. (0,59) (?).

Ce Scaphites (unique exemplaire) parait représenter un nouveau genre, défini par un

ensemble de caractéres empruntés a différents autres genres.

Ammonites massive, épaisse, subsphérique, a flancs plats fortement convexes et

région externe un peu plus large que ceux-ci. Section largement semi-lunaire.

Ombilic difficile 4 apprécier (probablement peu large) parce que, l’exemplaire étant

creux, |’enlevement de ce qui subsiste de la gangue ferait tomber les tours internes

265

Ann. S. Afr. Mus. 102 (7), 1993: 265-271, 2 figs.

266 ANNALS OF THE SOUTH AFRICAN MUSEUM

C D

Fig. 1. Menuites menu (Forbes, 1846). A—B. Paralectotype, BMNH C47599. C—D. Lectotype,

BMNH C51048, from Pondicherry, South India. All figures are x 1.

ON THE AFFINITIES OF COBBANOSCHAPHITES 267

retenus par celle-ci; mais il est éminemment infundibuliforme (voir la Fig. de face).

Ornementation de cotes basses, espacées, prenant librement naissance au sommet de

la fosse ombilicale, et aboutissant, sur la chambre d’habitation seulement, a six trés

gros tubercules externes (qui n’existent pas sur la partie cloisonnée). Cloisons a

premier lobe latéral trifide; deuxieme lobe étroit; selles hautes et étalées.

Rapports et Differences —L’ornementation de cdtes est celle de Desmoscaphites (cf.

Reeside, Eagle Sandstone. Pl. 22, Fig. 8), tandis que les tres gros tubercules exter-

nes sont exactement ceux de Acanthoscaphites (cf. Nowak, Reeside, Cobban); mais

ils n’existent que sur la chambre d’habitation. La cloison a le premier lobe de

Desmo-scaphites, tandis que l'ensemble rappelle celle de Clioscaphites (cf. Cobban.

Scaphitoid Cephalopods of the Colorado Group. Pl. 20-21).

Zone a Karapadites karapadensis. Sous-Zone a Maorites Aemilii. Gisement 191 de

la Coupe de Berere II (Belo sur Tsiribihina). Coll. M. Collignon, 1954.

(1) Le Genre est dédié a Mr. W. A. Cobban, Paléontologiste Américain, auteur de

nombreux Travaux sur le Genre Scaphites. (Collignon 1969: 51, pl. 533 (fig. 2097)).

Examination of the holotype, and only known specimen, showed that, rather than

being a distinctive scaphite, it is an adult microconch pachydiscid, referable to the genus

Menuites Spath, 1922, as is discussed below.

CONVENTIONS

All dimensions given below are in millimetres; D = diameter, Wb = whorl breadth,

Wh = whorl height, U = umbilical diameter.

Figures in parentheses are dimensions as a percentage of the total diameter.

The suture terminology is that of Wedekind (1916), as reviewed by Kullmann &

Wiedmann (1970), E = external lobe, L = lateral lobe, U = umbilical lobe, I = internal

SYSTEMATIC PALAEONTOLOGY

Class CEPHALOPODA

Order AMMONOIDEA Zittel, 1884

Suborder AMMONITINA Hyatt, 1889

Superfamily DESMOCERATACEAE Zittel, 1884

Family Pachydiscidae Spath, 1922

Genus Menuites Spath, 1922

[= Neopachydiscus Yabe & Shimizu, 1926; Anapachydiscus Yabe & Shimizu, 1926;

Besairieites Collignon, 1931; Cobbanoscaphites Collignon, 1969]

Type species. Ammonites menu Forbes, 1846 (p. 111, pl. 10 (fig. 1)), by original

designation of Spath (1922: 123).

Discussion

The type species of Menuites, M. menu, is based on a lectotype and six paralecto-

types from the Upper Maastrichtian Valudavur Formation of Pondicherry, South India,

268 ANNALS OF THE SOUTH AFRICAN MUSEUM

and has been revised by Kennedy & Henderson (1992: 430, pl. 14 (figs 1-15), text-

fig. 12c; see Fig. 1A—D herein). It is a small form reaching a maximum known diam-

eter of 70 mm (Kennedy & Henderson 1992, pl. 14 (figs 10—15)). The phragmocone

has a depressed reniform whorl section, and an ornament of umbilical bullae from a

diameter of 15mm onwards, that give rise to groups of fine, crowded irregular ribs

and growth lines, while additional ribs intercalate between. Strong ventrolateral tuber-

cles that alternate on either side of the venter appear on the last part of the phragmo-

cone at a diameter of 40 mm, and persist on to the adapical part of the body chamber.

Ribbing weakens on the body chamber, which is densely lirate, whereas ventro-

lateral tubercles are absent on the last 90°—120° sector. There are two constrictions

on the lectotype, one 45° before, and one immediately preceding the aperture

(Fig. 1C—D). A single constriction precedes the aperture in a paratype (Fig. 1A—B).

Similar tuberculate pachydiscids referred to Menuites are known from the Lower

Campanian to Upper Maastrichtian and, in a number of cases, occur with much larger

pachydiscids referred to the genus Anapachydiscus Yabe & Shimizu, 1926. These co-

occurring pairs have identical early developmental stages, and constitute dimorphic pairs,

as demonstrated by Cobban & Kennedy (in press) for Campanian species from the U.S.

Western Interior. In northern Hokkaido in Japan, a distinctive Menuites, described by

Matsumoto (1984: 17, pl. 5 (fig. 1), text-fig. 5) as M. sanadai, occurs in Upper Cam-

panian rocks associated with typical large specimens of the type species of Anapachydis-

cus, A. fascicostatus (Yabe 1921) (in Yabe & Shimizu 1921: 57, pl. 8 (fig. 5), pl. 9

(figs 2-5); see Matsumoto 1984: 14, pl. 4 (figs 1-2), pl. 5 (fig. 2), pl. 8 (fig. 7), text-fig. 4).

These co-occurring Menuites and Anapachydiscus have similar early developmental

stages and are clearly dimorphs, so that Anapachydiscus is a junior synonym of Menuites.

That small, tuberculate pachydiscids are microconchs of larger non-tuberculate forms was

demonstrated by Kennedy & Summesberger (1984) and discussed by Kennedy (1986);

current work on South African pachydiscids shows it to be present in Pachydiscus

(Neodesmoceras) Matsumoto, 1938.

Menuites menabensis (Collignon, 1969)

Fic. 2A—D

Cobbanoscaphites menabensis Collignon, 1969: 51, pl. 533 (fig. 2097).

Type

Holotype, by monotypy, is the original of Collignon, 1969: 51, pl. 533 (fig. 2097),

from the Upper Campanian Maorites aemilii [sic] Subzone of the Karapadites

karapadensis Zone, Gisement 191 de la Coupe de Berere II (Belo sur Tsiribihina),

Madagascar.

Dimensions

D Wh Wh Wb: W U

Holotype 80,0(100) 51,5 (64,3) 35,0 (43,8) 1,47 24,5 (30,6)

ON THE AFFINITIES OF COBBANOSCHAPHITES 269

SSR AE

Meee

Shee

Fig. 2. Menuites menabensis (Collignon, 1969). The holotype, the original of Collignon (1969, pl. 533

(fig. 2097)), from the Lower Campanian of Belo sur Tsiribihina, Madagascar. All figures are x 1.

270 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

The holotype is a slightly distorted internal mould, two-thirds of the last whorl is

body chamber, and a short section of the adapertural phragmocone is also preserved.

Coiling is moderate involute, the umbilicus comprising an estimated 30 per cent of the

diameter, with a broadly rounded wall and shoulder. On the phragmocone, the whorl

section 1s very depressed, reniform, with a whorl breadth to height ratio greater than two

(the specimen is defective); the only detectable ornament is distant, blunt umbilical bul-

lae. The body chamber is much better preserved, with a depressed, reniform whorl

section, and whorl breadth to height ratio of 1,47 at the adapertural end. Six low, broad,

straight prorsiradiate ribs arise on the umbilical wall, and strengthen into long umbilico-

lateral bullae. The bullae give rise to single narrow, straight ribs that link to very strong,

blunt conical ventrolateral tubercles. A pair of delicate ribs loops between these tuber-

cles, which are opposite rather than alternate. The interspaces between the ribs are

decorated by delicate, distant, prorsiradiate riblets, straight and prorsiradiate on the flank,

and feebly convex across the venter. They become increasingly prominent on the adaper-

tural end of the specimen, where there is an incipient constriction flanked by two of these

riblets.

The suture is imperfectly preserved but there is a large ventral lobe, a deeply incised

E/L, and L/U2 with narrow stems, and deeply incised L.

Discussion

A comparison of Figures 1A—D and 2A—D demonstrates the striking resemblance

between Cobbanoscaphites menabensis and Menuites menu. Any doubts that Cobbano-

scaphites might not be a pachydiscid, rather than a scaphite, are removed by a consid-

eration of the suture. Although imperfectly preserved (Fig. 2), the degree and nature of

the incisions are typically pachydiscid, as is the very narrow stem of the saddles.

Because the inner whorls of the holotype are so poorly preserved, it is not possible

to link Menuites menabensis with its corresponding macroconch.

Occurrence

As for the type.

ACKNOWLEDGEMENTS

Kennedy acknowledges the financial support of the Natural Environment Research

Council (U.K.), and the technical assistance of the staff of the Geological Collections,

Oxford University Museum, and Department of Earth Sciences, Oxford. Klinger acknow-

ledges financial support from the Foundation for Research Development (S.A.) and tech-

nical assistance from the South African Museum.

REFERENCES

Coppan, W. A. & Kennepy, W. J. (in press.) Dimorphic pachydiscid ammonites (Menuites) from the

Upper Campanian (Upper Cretaceous) of the Western Interior of the United States. Professional

Papers. United States Geological Survey.

CoLLicNon, M. 1931. Faunes Sénoniennes du Nord et de l’Ouest de Madagascar. Annales géologiques

du Service des mines, Madagascar 1: 7-64.

ON THE AFFINITIES OF COBBANOSCHAPHITES WALA |

CoLLicNon, M. 1963. Atlas des fossiles caractéristiques de Madagascar (Ammonites). X. Albien.

Tananarive: Service Géologique.

CoLiicNnon, M. 1964. Atlas des fossiles caractéristiques de Madagascar (Ammonites). XI. Cenomanien.

Tananarive: Service Géologique.

Co.iicnon, M. 1965a. Atlas des fossiles caractéristiques de Madagascar (Ammonites). XII. (Turonien).

Tananarive: Service Géologique.

CoLLicnon, M. 1965b. Atlas des fossiles caractéristiques de Madagascar (Ammonites). XIII. (Coniacien).

Tananarive: Service Géologique.

CoLLicNon, M. 1966. Atlas des fossiles caractéristiques de Madagascar (Ammonites). XVI. Santonien.

Tananarive: Service Géologique.

CoLiiGNon, M. 1969. Atlas des fossiles caractéristiques de Madagascar (Ammonites). XV. (Campanien

inférieur). Tananarive: Service Géologique.

CoLiicnon, M. 1970. Atlas des fossiles caractéristiques de Madagascar (Ammonites). XVI. (Campanien

moyen; Campanien supérieur). Tananarive: Service Géologique.

CoLiicNon, M. 1971. Atlas des fossiles caractéristiques de Madagascar (Ammonites). XVII. (Maés-

trichtien). Tananarive: Service Géologique.

Forses, E. 1846. Report on the fossil Invertebrata from southern India, collected by Mr. Kaye and

Mr. Cunliffe. Transactions of the Geological Society of London (2) 7: 97-174.

Git, T. 1871. Arrangement of the families of Mollusks. Smithsonian Miscellaneous Collections 227:

xvi, 1-49.

KENNEDY, W. J. 1986. Campanian and Maastrichtian ammonites from northern Aquitaine, France. Special

Papers in Palaeontology. Palaeontological Association of London 36: 1-145.

KENNEDY, W. J. & HENDERSON, R. A. 1992. Non-heteromorph ammonites from the Upper Maastrichtian

of Pondicherry, south India. Palaeontology 35 (2): 381-442.

KENNEDY, W. J. & SUMMESBERGER, H. 1984. Upper Campanian ammonites from the Gschliefgraben

(Ultrahelvetic, Upper Austria). Beitrdge zur Paldontologie von Osterreich 11: 149-206.

KULLMAN, J. & WIEDMANN, J. 1970. Significance of sutures in phylogeny of Ammonoidea. Paleontologi-

cal Contributions, University of Kansas 44: |—32.

Matsumoto, T. 1938. A biostratigraphic study on the Cretaceous deposits of the Naibuchi Valley, South

Karahuto. Proceedings of the Imperial Academy of Japan 14: 190-194.

Matsumoto, T. 1984. Some ammonites from the Campanian (Upper Cretaceous) of northern Hokkaido.

Special Papers. Palaeontological Society of Japan 27: 1-93.

SpATH, L. F. 1922. On the Senonian ammonite fauna of Pondoland. Transactions of the Royal Society of

South Africa 10: 113-147.

WEDEKIND, R. 1916. Uber Lobus, Suturallobus und Inzision. Zentralblatt fiir Mineralogie, Geologie und

Paldontologie 1916: 185-195.

YABE, H. & Suimizu, S. 1921. Notes on some Cretaceous ammonites from Japan and California. Scientific

Reports of Tohoku Imperial University, Sendai (2) 5: 53-59.

Yase, H. & Suimizu, S. 1926. A study of the genus ‘Parapachydiscus’ Hyatt. Proceedings of the Imperial

Academy of Japan 2: 171-173.

ZITTEL, K. A. voN 1884. Handbuch der Palaeontologie. 1, Abt. 2; Lief 3, Cephalopoda: 329-522. Munich

& Leipzig: R. Oldenbourg.

; ch

ae pink ap

: ei 2 aan ,

a a oly

Rey

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WILLIAM JAMES KENNEDY

HERBERT CHRISTIAN KLINGER

ON THE AFFINITIES OF COBBANOSCAPHITES

COLLIGNON, 1969 (CRETACEOUS AMMONOIDEA)

VOLUME 102 PART 8 FEBRUARY 1993 . a ISSN 0303-2515

OF THE SOUTH AFRICAN

MUSEUM

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

expérimentale et générale 74 (33): 627-634.

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. 1960b. 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-

pologische Ergebnisse einer Forschungsreise im westlichen und zentralen Stid-Afrika ausgefiihrt in den Jahren

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ANNALS OF THE SOUTH AFRICAN MUSEUM

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Part 8 Deel

THE BASKETWORK OF SOUTHERN AFRICA.

PART 2.

BASKETWORK OF THE KHOISAN AND THE DAMA

E. M. SHAW

Cape Town Kaapstad

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THE BASKETWORK OF SOUTHERN AFRICA.

PART:

BASKETWORK OF THE KHOISAN AND THE DAMA

E. M. SHAW

African Studies and Anthropology, South African Museum, Cape Town

(With 18 figures)

[MS accepted 9 December 1991]

ABSTRACT

In Part 1 of this study, the techniques and materials used in southern African basketwork were

described. Part 2 describes the objects made, their uses, and the techniques and materials used by the

Khoisan and the Dama. The study is based on the few extant museum specimens and on the literature.

Virtually no fieldwork was undertaken; this part is intended merely to gather what evidence remains.

It might be expected that basketwork, because of its comparative lightness, would have featured

prominently in the material culture of people constantly on the move. With three notable exceptions,

however—mats, traps and sieves—it did not.

CONTENTS

PAGE

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INTRODUCTION

The people whom early anthropologists designated Khoisan embraced two major

cultural divisions—the San (Bushmen), who were semi-nomadic hunters and food-

gatherers, and the Khoikhoi (Hottentots), who were herders and moved about follow-

ing the grazing for their cattle, sheep and goats.

Formerly, San hunter-gatherers seem to have been spread, doubtless in small groups,

Pie

Ann S. Afr. Mus. 102 (8), 1993: 273-301, 18 figs.

274 ANNALS OF THE SOUTH AFRICAN MUSEUM

over the whole of southern Africa, but it is only in the Kalahari and adjacent areas that

some remain as distinct groups today.

By early historic times, the Khoikhoi were living mainly in the coastal area below

the escarpment, from the Swakop River in the north to the Great Fish River in the east.

While some of them still recognize their original affiliations, none live in a tribal situation

today.

People who are constantly on the move tend not to have an elaborate material

culture, particularly if, like the San, they have no means of transport other than them-

selves. Because of its comparative lightness, one would expect basketwork to have fea-

tured more in the culture, but with three notable exceptions—mats, traps and sieves—it

evidently did not.

The Dama of Namibia, formerly called Bergdama or Bergdamara, are people about

whose origins very little is known. All that is known for certain is that for many gener-

ations they were associated often, but not always, as serfs or slaves, with the Nama, whose

language they speak. Whether this association took place before or after their entry into

Namibia is not known. In Namibia their form of subsistence was hunting and gathering

veld foods. Later, when the Herero entered the country, they too enslaved groups of

Dama. One would have expected, therefore, that the material culture of the Dama would

have been strongly influenced by those with whom they came in contact, neither of whom

had specialized in basketwork. Very little basketwork has been recorded from the Dama.

The text is arranged under headings of class of object, and the sources are

arranged in date order of sojourn of the author, or of the museum acquisition, where

known. Date of publication follows in square brackets, and it stands alone if the date

of sojourn is not known. Sources are grouped as: “Early’—up to 1899, and ‘Recent’ —

from 1900. Because so many of even the quite early sources are quotations from

even earlier sources, eye-witness accounts only are quoted as far as can be judged.

As far as possible, the place where the author was when he saw the object described

is noted. The localities follow, therefore, the travel routes of the respective authors.

This arrangement does not, of course, imply that the objects described by them

occurred in that particular place only. In figure captions and text, numbers in italics

refer to technique numbers allocated in Shaw (1992).

SAN

MATS

Roofing mats were made of sedge stems, in the straight-sewn technique. Generally,

one or two mats were used for each dwelling, which, as Daniell (1820, pl. 26) depicts

(Fig. 2), was little more than a windscreen. One mat would be placed on the side from

which the wind was coming, in a semicircle supported by two or three sticks; the other

might form a roof. When the group moved on, the mats could be rolled up with the sticks

and carried. This type of shelter is also described by Le Vaillant (1803) from just north

of the Orange River.

The mats seem to have been identical with those of the Khoikhoi. Judging by the

frequency with which they are mentioned in the folk tales and in the records of early

travellers, the mats were in common use by the southern San who lived away from

mountains, and it should not therefore be considered that those who used them were

Fig. 1. A. Straight sewing, sins

single lattice wrap, 5c. D. Wick

4a. F. Wefts taken back diagot

G next row, loop at edge, 85e. H.

taut at edge, 85d. J. Wrapping

oversewing over one, multiple f

foundation, 17a. M. Beginning

work: closed ring, 38a. O. Dias

in colour; coiled work, /5/c. Q

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Fig. 1. A. Straight sewing, single foundation, 8. B. Wicker: plain open twine, 5j. C. Wicker:

single lattice wrap, 5c. D. Wicker: chequer weave, 5a, E. Plain close twine weave, single warp,

4a. F. Wefts taken back diagonally, extra twist at edge, 850. G. Sewing strand taken down to

next row, loop at edge, 85e. H. Plain open twine, 4c. I. Sewing strand taken down to next row,

taut at edge, 85d. J. Wrapping—over two/four and under one/two, 76a. K. Coiled work: simple

oversewing over one, multiple foundation, 7b. L. Coiled work: plain beeskep sewing, multiple

foundation, 17a. M. Beginning of coiled work: bent foundation, 37a. N. Beginning of coiled

work: closed ring, 38a. O. Diagonal oversewing of edge, round coil, 97d. P. Geometric pattern

in colour; coiled work, 75/c. Q. Square plait, 7g. R. Finish: sewing strand threaded back, 84d.

SSS

Cpe

S. Straight-sewn work: decorative band by crossing foundation elements, 128a. T. Coiled

work: simple oversewing over two coils of multiple foundation, /Jc. U. Simple plait, 7a.

BASKETWORK OF SOUTHERN AFRICA 2A

Fig. 2. San windscreen, Daniell ( 1820, pl. 26), 1804, near Prieska.

impoverished Khoikhoi. In fact, Burchell illustrates from south of the Orange River, an

apparently far from impoverished settlement, with full, if small, huts with withy domes,

covered with mats (Burchell 1822). According to Burchell, some San immediately south

of the Orange River painted their mats lengthwise with stripes of red ochre.

Mats, presumably the same as those above, were used in the Lake Ngami and

Wankie areas for roofing, and in the Okavango area for roofing or as windscreens.

According to Seiner (1910), those in the Okavango area were obtained from the

Mbukushu, but no other author reports this and the hut depicted is of Khoikhoi style, and

the mats are straight-sewn.

Techniques

Fabric—straight-sewn, single foundation (Fig. 1A, 8).

Ornamentation—painting of mats by some southern San.

No further information.

Tools

A bone awl.

Materials

Foundation—sedge (Cyperus sp.).

Sewing—cord of inner bark or other plant fibre; antelope sinew.

276 ANNALS OF THE SOUTH AFRICAN MUSEUM

Makers

Women.

Records

Early: Gordon 1777 [1988: 132], near Steynsburg. Le Vaillant 1783-1784 [1803:

167], just north of Orange River near Fish River in Namibia. Barrow 1797 [1804: 272,

275], Colesberg. Daniell 1802 [1820, pl. 26], near Prieska. Burchell 1811-1813 [1822b:

frontispiece, p. 141], between De Aar and Orange River. Campbell 1813 [1822a: 105],

near Kuruman. Backhouse 1839 [1844: 203], Shiloh, eastern Cape. Arbousset & Daumas

[1846: 249, 358], near Morija. Bleek & Lloyd c. 1873 [1911: 183, 185, 203, 205], Katkop,

north Cape. Stow before 1880 [1905: 43-44], ‘plains’.

Recent: Seiner [1910: 342, 345, 359], Okavango. Dornan [1925: 90], Ngami,

Wankie. Hirschberg [1933: 128], general.

San huts or shelters, as described in the literature, did not have formal frames, but

Lebzelter (1934, pl. 4:4) shows a !Kung hut frame on which side lattices are attached by

single wrapping. A similar use of lattice wrapping was seen among the !Kung of Nama

Pan in 1975.

No further information.

Records

Early: nil.

Recent: Lebzelter 1926-1928 [1934, pl. 4:4], northern Namibia.

Field survey: 1975, Tsumkwe.

SIEVES

In the south, sieves were used for sifting small bulbs, seeds, ants’ larvae and roots.

According to Burchell, they were about 900 mm by 600 mm and were made in the same

way as roofing mats ‘except that the rushes are not so close together’, and that they were

‘extended by a stick tied along each side and by others fastened to them transversally’. In

areas where there were a lot of small bulbs and roots, the earth was sifted through the

sieves So as to obtain the roots quickly. No other author reported this, but among the //n/ke

of Gordonia, D. F. Bleek collected sieves that, except in size and the fact that the warps

are thin reeds, fit this description (Fig. 3).

In the south-western Kalahari, sieves were used for sorting white ants’ larvae or

melon seeds, and for sifting them from the ashes after roasting or, according to Kling

(1925 correspondence), for laying out and cooling cooked meat. In that area they were

made of thin reeds, in a flexible style without the supporting sticks, so that they could be

rolled like a mat. According to Steyn (1971: 284) they are made, in the Ghanzi area,

exclusively for sifting roasted melon seeds.

The flexible sieves (Fig. 4) are said to have been used in the central Kalahari also,

for sifting roasted larvae of termites or roasted grass seeds. No sieves were seen among

the !Kung of Tsumkwe in 1975, nor did Marshall record their use among the !Kung of

Nyae Nyae, or Lee record having seen them among the !Kung of Dobe.

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278 ANNALS OF THE SOUTH AFRICAN MUSEUM

Techniques

(a) Fabric: open twine wicker (Fig. 1B, 57); one strand open lattice wrap (Fig. 1C, 5c)

with lattices in three places, over four and under two; two rows chequer (Fig. 1D, 5a).

Method of work: not described.

Beginning: knotted wefts.

Shaping: natural increase or decrease of warps.

Shape: roughly rectangular.

Edges: warp—none; several rows of close twine (Fig. 1E, 4a); weft—one weft strand

wound round last warp which may be reinforced, then back diagonally (Fig. 1F, 850);

weft twined round last warp for short distance; one edge knotted and cut, other taken

down with loop (Fig. 1G, 85e).

(b) Fabric: open twine (Fig. 1H, 4c).

Method of work: not described.

Beginning: knotted weft.

Shaping: none.

Shape: rectangular.

Edges: warp—several rows of close twine (Fig. 1E, 4a); weft—at one edge weft

taken down to next row, with (Fig. 1G, 85e) or without (Fig. 11, 85d) a loop, and

returned, once only; at other edge knotted.

Finishings: ends knotted together.

Decoration: none.

Ornamentation: none.

Method of work: not described.

Beginning: knotted sewing strand.

Shaping: none.

Shape: rectangular.

Edges: sewing: sewing strand carried down straight (Fig. 11, 85d) or with a loop (Fig.

1G, 85e) and then back, continuously; foundation: sewing strand wrapped once over

two and under one foundation strand (Fig. 1J, 76a).

Finishings: a knot.

Decoration: none.

Ornamentation: none.

Tools

Not known.

Materials

Warp—thin reed stems; thin withies; weft—thin thongs, thin bark-fibre cord; sinew.

Makers

Not known.

BASKETWORK OF SOUTHERN AFRICA 279

Records

Early: Burchell 1811-1813 [1822b: 20], near Prieska. Bleek & Lloyd 1873 [1911:

85], Katkop, N. Cape. Stow before 1880 [1905: 59], Koesberg, N. Cape.

Recent: SAM-1552, 1911, Gordonia. Wit. Univ. 2441, no date, Gemsbok Park.

Drury 1921 [1935: 94], Sandfontein. SAM—3679, 1921, Nharo, Sandfontein. SAM—4284,

1925, Kalahari. Dornan [1925, facing p. 48], Kalahari. Kling 1925, correspondence, east

of Gochas. Fourie [1928: 102], Namibia. Hirschberg [1933: 128], Oas, Rietfontein,

Sudonibsaup. SAM—7551, 1936, N. E. Kalahari. Maccrone 1937: 252, pls 97, 98; near

Auob and Nossop Rivers. Afr. Mus. 43/165, 1943, Gobabis. Marshall 1951-1961 [1976],

Nyae Nyae, Namibia. Lee 1963-1965 [1965], Dobe, northern Namibia. SAM—9141, 1965,

Nharo, Ghanzi. SAM—9333, 1966, Nharo, Ghanzi. Steyn 1968 [1971: 284-285], Ghanzi.

SAM-9621, 1969, Nharo, Kalkfontein.

Field survey: 1975, Tsumkwe.

FISH-TRAPS

Basketwork fish-traps were used by people who lived along the Orange River and

the Fish River in Namibia and probably near other rivers and marshes. They are described

by early travellers as neatly made funnel-shaped or pointed baskets, about 183 cm long

and 45-61 cm at the greatest diameter. The warps of those seen by Barrow (1801) near

the Orange River were alternately of ‘reeds’ and twigs of taaibos (Rhus sp.), light and

dark respectively, and ‘gave a pretty effect’. Other descriptions were that they were made

of ‘stick grass’ (this often meant sedge) and that they resembled the eel-baskets of Europe

Fig. 5. Eel-pot, England, c. 1860. Photo: A. Heseltine, Shire Publications Album 92; copyright

Institute of Agricultural History, University of Reading.

(Fig. 5). No authors, however, describe the inner valve, which is visible in the eel-baskets,

and is common in other African fish-traps (Fig. 6). According to Barrow (1801), they

were used either individually by men standing in the river, or were placed mouth for-

wards, in gaps in stone walls or reed fences, across or at the edge of the water. Alexander

(1838) saw men sitting in deep water holding the traps, while other men chased the fish

into them. These two original sources have been quoted by several other authors.

280 ANNALS OF THE SOUTH AFRICAN MUSEUM

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Fig. 6. Ambo fish-trap, SAM-6124, Oshikango, 1940, typical for southern Africa. Diameter of

mouth 465 mm.

Fig. 7. Drawing of a San fish-trap. Van Rippen 1918, fig. 24.

BASKETWORK OF SOUTHERN AFRICA 281

According to Dornan (1925), he saw the same type of funnel-shaped basketwork

fish-traps used by San people in the north-east, along the Botletle River, placed in gaps

in stone weirs, and facing upstream. Sometimes a group of men would sit in a row holding

the traps, while others went upstream and chased the fish into the traps. This description

does resemble that of Alexander, but there is nearly a century between them.

In 1917, Van Rippen (1918) had a fish-trap made for him by a 42-year-old San man,

who claimed that his people used to make such traps (see Fig. 7). Van Rippen does not

state where he travelled, nor where he was at the time. As far as can be seen, the technique

is open lattice wrap wicker (Fig. 1C), and the trap is said to have been made of ‘taai

boschje’. The valve is clearly visible. Van Rippen questions whether this is of ‘pure

Bushman origin’.

Techniques

Fabric: ‘woven’; lattice wrap (or twine) wicker.

Shape: ‘pointed’ or ‘funnel-shaped’; a cone flattened at the narrow end. No further

information.

Tools

No information.

Materials

Warps—‘reeds’ and twigs of taaibos (Rhus sp.); taaibos alone; ‘stick grass’; Aristida

sp.: wefts—cord made of “bruised rushes’ or of the inner bark of mimosa (Acacia sp.).

Makers

Men (according to Van Rippen).

Records

Early: Barrow 1797-1798 [1804: 290], general, [p. 300], Orange River west of

Philippolis. Lichtenstein 1803-1806 [1930: 55], Orange River. Alexander 1835 [1838a:

237, 238], Fish River (Namibia). Moffat 1817-1818 [1842: 55], Namaqualand. Hahn

[1870: 103], Orange and Zuga Rivers, partly quoting Barrow. Stow before 1880 [1905: 93],

general, quoting Barrow.

Recent: Seiner [1910: 345], Okavango River. Van Rippen 1917 [1918: 92, 93], no

locality. Dornan [1925: 51, 106], Botletle River.

MISCELLANEOUS

BASKETS

No distinctive basketry containers have been recorded as used by the San. The only

actual baskets seen among them were Tswana-style garden baskets made by Sarwa near

Serowe in 1962, for sale to Tswana. At the temporary camp where these were seen, they

were also in use. The technique and the materials were those used by the local Tswana.

They were made by women and the tool used was an awl.

In addition to those made near Serowe, Ebert (1978) recorded a flourishing industry

in Tswana-style baskets among the Sarwa along the Nata River, north-west of Francis-

town—again for use at home as well as for sale. Four types are made, three by women

282 ANNALS OF THE SOUTH AFRICAN MUSEUM

—the garden basket, a beer storage vessel and a basket with lid and handle, and one by

men—a winnowing basket or tray.

Stow (1905) stated that ants’ eggs, after they had been sieved and sorted, were put

into a small grass basket or a skin bag. He does not mention where or among whom he

saw this and no confirmation of it has been found.

Records

Early: Stow before 1880 [1905: 59].

Recent: Ebert 1977 [1978: 69-83], Nata River.

Field survey: 1962, Serowe.

STORAGE BINS

According to Lee (1965), each !Kung family in the Dobe area made a storage bin of

‘branches and palm fronds woven together in the crotch of a young tree’, in which to keep

foods. The bin is not further described and may not be basketwork. Bins were not seen

among the !Kung of Tsumkwe in 1975.

No further information.

Records

Early: nil.

Recent: Lee 1963-1965 [1965: 180].

Field survey: 1975, Tsumkwe.

HATS

Some people in the north-east made and wore hats (Fig. 8). These were rather

roughly made in an eighteenth century ‘sailor’ shape. Hats are said to have been worn by

!'Kung women and other San in Namibia.

Techniques

Fabric: simple or beeskep oversewing (Fig. 1K, //b, L, 17a) over multiple or simple

coil foundation.

Method of work: not seen.

Beginning: bent foundation (Fig. 1M, 37a); closed ring (Fig. IN, 38a).

Shaping: placing of coil.

Shape: ‘sailor’.

Edge: diagonal oversewing around coil in opposite direction (Fig. 10); leather

binding.

Finishing: ends worked in.

Decoration: sewing: geometric patterns in different coloured material (Fig. 1P, 149f) .

Ornamentation: nil.

Tools

Not seen.

Materials

Foundation: grass; reed.

Sewing: strands of bark; palm leaf.

Makers

Generally not known, but among the !Kung, men.

BASKETWORK OF SOUTHERN AFRICA 283

Fig. 8. San hat, SAM—7089, northern Kalahari, c. 1936. Diameter of crown 175 mm.

Records

Early: Alexander 1835 [1838: 236], Namibia.

Recent: Lebzelter 1926-1928 [1934: 36, pl. 7: 5, 6], northern Namibia. SAM-—7089,

1936, north-east Kalahari.

ORNAMENTS

Plaited grass ornaments are mentioned by Smith (1836 [1975]). A bangle from Lake

Chrissie is an eight-strand square plait of Digitaria grass, but may have been a borrowing

from the Swazi.

The Nharo made necklets and bangles of Scirpus nodosus and Kyllinga alba (Bleek

1928). According to Passarge (1907), those in the Ghanzi area were “woven spirally’ by

children. According to Hirschberg (1933), at Rietfontein, ornaments were made from the

lengths between the nodes of a grass stem, split in two and ‘interlaced’.

Grass ornaments are common throughout southern Africa.

Techniques

Fabric: eight-strand square plait (Fig. 1Q, 7q); spiral plait; ‘interlaced’.

Finishing: oversewn.

Materials

Flowering stems of Digitaria sp., Scirpus nodosus Rottb., Kyllinga alba Nees, etc.

Records

Early: Smith 1834-1836 [1975: 147], Riet River near Philippolis. Passarge 1896

[1907: 90-91], Ghanzi.

284 ANNALS OF THE SOUTH AFRICAN MUSEUM

Recent: Schultze 1903-1905 [1907: 657-658], southern Botswana. Bleek 1921

[1928: 67], Sandfontein. SAM-—1795, 1922, Lake Chrissie. Hirschberg [1933: 3], Oas,

Rietfontein, Sudonibsaup.

PLAITED ROPE

There is an isolated report of a plaited grass rope used as a hand-hold on a ladder by

San honey-gatherers in southern Lesotho.

Technique

‘Plaited’.

Material

‘Grass’.

Records

Early: nil.

Recent: Walton [1956: 15-16].

KHOIKHOI

The material culture of the Khoikhoi was not uniform throughout the three large

areas where the major groups of Khoikhoi lived—that is, east of the Gamtoos River,

between the Gamtoos and the Olifants Rivers, and north of the Olifants River. It seems

best in this discussion to group the people of the first two areas together, and to separate

those of the north, the Nama. One item in particular, however, was common to all—their

type of dwelling, which was a framework hut of beehive shape, roofed with the long mats

that were their main item of basketwork. The great advantage of the mats was that in wet

weather the sedge stems, of which they were made, swelled and made the cover water-

tight, while in dry weather the slight gaps between the stems allowed the air to circulate.

MATS

Gonaqua, Cape Khoikhoi, Korana, Griqua

The only articles of basketwork that all Khoikhoi appear to have made and used were

mats made of the long inflorescence stems of sedges (Fig. 9). Their main and universal

purpose was for roofing the huts (Figs 10, 11, 12). The huts, some large, some small,

consisted of a framework of previously bent poles, planted in a circle and tied where they

crossed. Mats of various sizes, according to their position, were laid over the frame and

tied on to it. The order in which the mats were disposed was not the same everywhere. A

single small mat was tied above the lintel of each of the doorways, back and front, and

rolled up or let down to open or close the doorway. When the owner moved, the hut was

dismantled and poles and mats were transported, tied to each side of a pack-ox (Fig. 13).

The sedge stems were prepared by drying in the sun and damping for use if neces-

sary. To make the mats, the stems were laid side by side and joined by straight-sewing.

Holes were pierced in the stems a short distance apart and a thin cord of sedge, of the

inner bark of an acacia species, sinew, or—as early as 1772—of European pack-thread,

was threaded through the holes.

BASKETWORK OF SOUTHERN AFRICA

Fig. 9. Sedge, Cyperus textilis, Montagu area. Photo: W. van Rijssen.

Fig. 10. Le Vaillant’s drawing of the arrangement of mats on a Khoi hut near

Great Fish River, 1780-1803 [1973, pl. 58].

286 ANNALS OF THE SOUTH AFRICAN MUSEUM

\ <

Fig. 11. Korana hut, Daniell 1820, pl. 1B, Read’s Drift, Orange River near Prieska, 1804.

Fig. 12. Gonaqua village, Le Vaillant 1790, pl. 12, west of Great Fish River, 1780-1785.

BASKETWORK OF SOUTHERN AFRICA 287

Fig. 13. Korana preparing to move house, Daniell 1820, pl. 20, banks of Orange River, 1804.

According to Burchell (1822), the women of Genadendal made mats for sale as well

as for their own use.

Remains in a cave on the Bredasdorp coast (Grobbelaar & Goodwin 1952) suggest

that the mats were also used as sleeping mats. For adults skins were more generally used,

but children slept on mats (Engelbrecht 1936).

Techniques

Fabric: straight-sewn (Fig. 1A, 8).

Method of work: foundation elements laid side by side and joined by sewing through

holes pierced at short intervals. The sewing ran lengthwise and the full length of the mat

was sewn before the next row was commenced.

Beginning: not recorded.

Shape: rectangular.

Shaping: none.

Edge: top and bottom: a single or double row of twine (Fig. 1E, 4a); sides: sewing

carried down to next row taut (Fig. 11, 85d) or leaving a loop for tying (Fig. 1G, 85e).

Finishing: loose ends knotted at edge.

Decoration: none.

Ornamentation: none.

Tools

An awl of bone or wood; a needle of ostrich bone or wood.

288 ANNALS OF THE SOUTH AFRICAN MUSEUM

Materials

Foundation: stems of Cyperus textilis at Genadendal, or Scirpus inanis near Fraser-

burg, and other sedges elsewhere. The Korana used a less durable material for mats for

children to sleep on.

Sewing: two-ply twisted cord of crushed sedge stems, the inner bark of Acacia

capensis Or Acacia karroo, a soft outer bark, sinew, or pack-thread.

Makers

Women made both mats and sewing strand.

Records

Early: Dapper 1668 [1933: 57, 59], Cape. Schreyer 1679 [1965: 93], Cape. Graeven-

broeck 1695 [1933: 251], Cape. Kolben 1705-1713 [1738: 220-221, 236], Cape. Thun-

berg 1773 [1986: 206], Cogmans Kloof; [p. 238], Van Stadens River. Sparrman

1772-1776 [1785a: 207-209], near Swellendam. Le Vaillant 1780-1785 [1790a: 159-160],

near Mossel Bay; [1790b: 39], near Great Fish River. Somerville 1801 [1979]: 89, south

of Griquatown. Daniell 1804-1805 [1820, pl. 20], Orange River, south of Griquatown.

Burchell 1811-1813 1822a: 83, Genadendal; 186-187, Riet River; 228-229, pl. 5, north

bank of Orange River, east of Prieska; [pp. 279, 281-282], Vaal River, east of junction

with Orange; [1822b: 88], Seekoe River, north of Sneeuwberg. Pitt Rivers Museum:

A: M. 645-651, 1822, Genadendal, Fraserburg. Campbell 1820 [1822b: 346], east of

Kuruman. Thompson 1823-1824 [1827a: 120, 122], east of junction Vaal and Orange;

[1827b: 46-47], near Aughrabies Falls. Smith 1834a [1839: 80], Riet River, east of

Philippolis. Backhouse 1838-1840 [1844: 433], Griquatown. Mackenzie [1871: 499],

general. Stow before 1880 [1905: 240, general: 276], near junction Vaal and Orange.

Recent: Engelbrecht 1927 [1936: 93-96], Bloemhof. Afr. Mus. M48/53, 1948,

Bloemhof. Grobbelaar & Goodwin 1952: 102-103, Skipskop Cave, Bredasdorp. Wells

1965: 81, Gamtoos Valley.

Nama

Mats (Fig. 14) of the same type as described above were made for the roofing of

huts. At least 15 to 20 were required for a single covering, and the making took a long

time. Some Nama dyed the sedge stems to part of their length by wrapping them in

bundles with old damp kraal manure. The stems could then be arranged in the mats to

make patterns with the dyed portions (Fig. 15). Mats were also made for babies to sleep

on, or were sometimes used as a floor covering. The roofing mats, with the hut frame-

work, could be dismantled and moved. They were said to last about 10 to 15 years but

the original and the dyed colour faded before that. Ollp (1884) noted that women made

enough to replace worn examples and also to sell. Their making and use have been

described fully by several authors (see particularly Haake 1982). By the end of the nine-

teenth century they were becoming rare because the sedges were difficult to obtain.

Substitute coverings had to be used, such as rough mats of Aristida grass, skins or

sacking.

According to Thompson (1827), rough mats were also used for fishing. (See p. 292.)

BASKETWORK OF SOUTHERN AFRICA 289

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290 ANNALS OF THE SOUTH AFRICAN MUSEUM

Techniques

Fabric: straight-sewn (Fig. 1A, 8).

Method of work: sedge gathered green, allowed to dry, trimmed, and soaked over-

night when about to be used. Stems are pierced at 50-70 mm intervals with a flat awl

(Fig. 16A) which is turned on its side from time to time to let the needle (Fig. 16C) with

the sewing thread (Fig. 16B) go through easily. The Nama south of the Orange River are

said to have taken the stems one or even up to four at a time, while those north of the

river took about 20. The awl moves away from the worker and the needle towards.

Finally, the ridges are tidied with the wooden handle of an awl and emphasized with a

special grooved tool (Fig. 17).

Beginning: the foundation strands are laid on the ground in a row and holes for the

thread are made at intervals. The work may commence at the centre of the width of the

mat, with a knotted loop.

Shaping: none.

Shape: rectangular.

Edges: foundation: single or double row of plain twine (Fig. 1E, 4a), in twos or

threes.

Sewing: carried down with loops (Fig. 1G, 85e).

Fig. 16. A. Nama awl for making mats, SAMUCT-—23/137, Fransfontein, 1923. Length 210 mm.

B. Nama string for sewing mats, SAMUCT—23/137, Fransfontein, 1923. Thickness 4 mm.

C. Nama needle for sewmg mats, SAMUCT-—23/138, Fransfontein, 1923. Length 210 mm.

BASKETWORK OF SOUTHERN AFRICA 291

Finishing: foundation strands cut off about 15 mm from the twining; sewing strands

knotted at beginning and threaded back (Fig. 1R, 84d) at end.

Decoration: crossed foundation strands (Fig. 1S, /26a); arrangement of same

material dyed.

Ornamentation: none.

Tools

A long thin awl (Fig. 16A) made from the shin bone of giraffe, or latterly of iron,

together with a needle made of the leg bone of an ostrich; an iron needle (Fig. 16C); a

thorn or piece of sedge to which the thread has been attached; a knife for cutting; a tool

for smoothing ridges (Fig. 17).

Fig. 17. Nama tool for emphasizing ridges of mat, SAMUCT-—23/152, Fransfontein, 1923. Width

across flat head 70 mm.

Materials

Foundation: sedge—Cyperus sp., Juncus sp., Scirpus sp.

Sewing thread: inner bark of Acacia polyacantha, A. karroo or other spp.; flexible

stems of Salix capensis (willow); Asclepius fruticosus; grain-bag string, to obtain which

the bags may be bought especially. The fibre for the thread is soaked in water or milk,

then chewed, to soften it, then twisted on the leg into two- or more ply thread (Fig. 15B).

Dye: old kraal manure.

Makers

Women.

Records

Early: Barrow 1797-1798 [1804: 389-390], Kamiesberg. Thompson 1822-1824

[1827b: 56, 59, 63, 64-65], Pella, Gams, general south of the Orange. Alexander 1835-—

1837 [1838a: 98], between Kleinsee and Orange River mouth; [pp. 171, 194], Warmbad.

292 ANNALS OF THE SOUTH AFRICAN MUSEUM

Shaw 1815-1837 [1840: 23], Kamiesberg (Lily Fountain) and general south of Orange

River. Ridsdale 1844-1847 [1883: 87], Nisbetts Bath (Warmbad). Andersson 1854 [1856:

326], Great Namaqualand. Hahn 1867: 306, general Namibia. Ollp 1865-1876 [1884: 28],

Gross-Namaland. Stow before 1880 [1905: 253], near Pella. Schinz 1884-1887 [1891:

86], Keetmanshoop. Von Francois 1889-1893 [1896: 209], general.

Recent: Schultze 1903-1905 [1907: 227-231, 241-242], ‘Great Namaqualand’.

Hoernlé 1913 [1987: 71-74], Walvis Bay. Hoernlé 1922 [1987: 127, 128], Fransfontein.

Hoernlé 1913-1922 [1923: 24], Namaqualand. SAMUCT-—23/131, 1923, Fransfontein.

Vedder [1928: 125], general Namibia; [1934: 51-52], general Namibia. Ferreira 1974:

1-6, general. SAM—9389, 1977, Leliefontein (Kamiesberg). Du Pisani 1981 [1983: 9, 11,

15], Kuisib River. Haake 1982: 80-84, general Nama. Van der Merwe 1984: 1-4, Lelie-

fontein.

STRAINERS

Nama

The only record of the use of a strainer was by the Topnaars of Walvis Bay for

straining boiled nara fruit (Acanthosicyos horrida) in the process of making nara cakes.

It appears from Schultze’s figure to be a deep bowl-shaped basket of open twine with a

thick coil once round the top and continuing to make a handle. It is very roughly made.

The coil is oversewn and the handle bound round with a two-ply twisted cord of the same

material. This strainer is also mentioned in Hoernlé’s 1913 diary.

Techniques

Fabric: open twine (Fig. 1H, 4c).

Edge: thick coil and handle, oversewn two-ply twisted cord.

Materials

Sedge.

No further information.

Records

Early: mil.

Recent: Schultze 1903-1905 [1907: 198-199], figure, Walvis Bay. Hoernlé 1913

[1987: 71], Walvis Bay.

FISH-TRAPS

Nama

There is no record of the use of traps by any Khoi other than Nama.

Basket traps, according to Wikar and Hoernlé, or ‘rush mats’, according to

Thompson, were used by Nama for fishing in shallow streams and in the Orange River.

They were placed across the stream with men standing behind them while other men

drove the fish towards the traps. Nama who lived away from the rivers, claimed, when

talking to Alexander, that they did not eat fish.

BASKETWORK OF SOUTHERN AFRICA 293

Materials

Acacia karroo; Rhus sp.

No further information.

Records

Early: Gordon 1779 [1988: 326], an island in the Orange River. Wikar 1779 [1935:

115], near Aughrabies Falls. Thompson 1821-1824 [1827b: 64-65], near Orange River

mouth. Alexander 1835-1837 [1838b: 64], near Orange River mouth. Von Francois

1889-1893 [1896: 211], general Namibia.

Recent: Schultze 1903-1905 [1907: 198-199], probably Orange River. Hoernlé 1923

[1923: 25], Namibia. Du Pisani 1981 [1983: 4], Kuisib River and general.

MISCELLANEOUS

BASKETS

Gonaqua

The Gonaqua did not themselves make baskets but obtained them from the Xhosa.

They were used as containers for milk and as pails for milking. Two such baskets, one

globular and lidded (Stockholm, 626), one flanging to a wide mouth (Stockholm

1799.2.101) are preserved in the State Ethnographic Museum in Stockholm, presented

respectively by Thunberg and Sparrman. There is no record of the use of baskets by Cape

Khoikhoi or Griqua.

Records

Early: Thunberg 1772 [1793: 102-103, 238], Gamtoos River. Sparrman 1772-1776

[1785a, pl. 1.1; 17855: 34, 35], Little Sundays River. Stockholm: 626, 1799, Gamtoos

River; Stockholm: 1799.2.101, 1799, ? Little Sundays River. Le Vaillant 1782-1783

[1790a: 375-376; 1790b: 4, 22], Great Fish River.

Korana

The Korana, similarly, obtained baskets from the Tswana. They did, however, fit a

basketwork neck onto containers of skin (Fig. 18).

Techniques

Fabric: multiple foundation, simple oversewing over two (Fig. IT, //c).

Method of work: not seen.

Beginning: start sewing into the skin, then add foundation.

Shaping: placing of coil.

Shape: circular band.

Edge: none.

Finishing: just end.

Decoration: none.

Ornamentation: none.

294 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 18. Korana flask for butter fat, SAM—4450, Barkly West, 1926. Diameter of bowl 240 mm,

mouth 113 mm.

Tools

Not known.

Materials

Foundation: split root of katbos (?Lycium hirsutum) or ‘rushes’ or thin withies.

Sewing: split root of katbos, or ‘rushes’ or latterly sedge and string.

Makers

Not recorded but surely men.

Records

Early: nil.

Recent: SAM-4450, 1926, Barkly West. Engelbrecht 1927 [1936: 101, pl. 8],

Bloemhof.

Nama

According to Vedder, baskets were made of ‘suitably flexible roots and twigs’, but

there is no information regarding their making or use.

Records

Early: nil.

Recent: Vedder 1928: 126—127.

BASKETWORK OF SOUTHERN AFRICA 295

HATS

Korana

Hats were worn by both men and women. They had circular or conical crowns and

flat brims. Narrow leather straps were sewn at the sides to tie under the chin.

Techniques

No information, but judging from the description and materials, coiled sewn on a

multiple foundation (Fig. 1K, 7/b).

Materials

Foundation: grass.

Sewing: rosyntjiebos (Grewia flava).

Makers

Men.

Records

Early: nil.

Recent: Engelbrecht 1927 [1936: 106], Bloemhof. Smith 1966: 403.

ROPE

Korana

There is no record of other branches of the Khoikhoi using plaited ropes, but it is

unlikely that they did not do so. Twined string, which is much used, is not interlaced and

therefore does not come under the definition of basketwork.

Plaited rope of wildebeest tail hair was made by the Korana for tying over the mats

of a hut. Later, horsehair was used.

Technique

Simple plait (Fig. 1U, 7a).

Materials

Wildebeest hair; horsehair.

No further information.

Records

Early: nil.

Recent: Engeibrecht 1927 [1936: 94—95], Bloemhof.

DAMA

BASKETS

1. A special narrow-mouthed basket in which locust meal is stamped and stored. It is hung

in a tree and lasts for one or two years. This may or may not be the ‘little bag’ mentioned

by Hahn (1984).

296 ANNALS OF THE SOUTH AFRICAN MUSEUM

Techniques

No information.

Tools

No information.

Materials

Grass; Cyperus marginatus; Stipagrostis namaquensis.

Makers

No information.

Records

Early: Hahn 1846-1851 [1984: 386], Central Namibia.

Recent: Lebzelter 1926-1928 [1934: 131], Okambahe, Namibia. Steyn & Du Pisani

1985: 44-45, Ugab River Valley, Namibia.

2. A flat basket for which no purpose is given.

Techniques

‘Woven’, ‘made tight with bast’.

Tools

No information.

Materials

Acacia bast and thin sticks split in half; Stipagrostis namaquensis.

Makers

Men.

Record

Early: nil.

Recent: Lebzelter 1926—1928 [1934: 178], general.

SIEVES

A shallow ‘woven’ basket through which ants’ eggs are sifted out of the sand for

food.

No further information.

Records

Early: nil.

Recent: Schwarz [1928: 201].

The lack of further information about Dama techniques makes it hard to judge

whether there is any item of basketwork that could be considered truly Dama.

BASKETWORK OF SOUTHERN AFRICA 297

SUMMARY

There is no doubt that basketwork was not a major feature of Khoisan or Dama

culture, with the exception of the mat which was universal among the Khoikhoi and used

by some San but not by the Dama. The technique of the mats was straight, flat sewing

with single foundation strands. The twined technique was known and was used for traps

along the rivers, for sieves, especially for flexible sieves, and occasionally for the edging

of mats. Apart from the latter, edges and finishings were merely the most practical way

of dealing with a problem. The only decoration recorded is the Nama dyeing of sedge

stems, to make a chequered pattern on the finished hut. For ornamentation, there is

Burchell’s (1822) statement that the San south of the Orange used red ochre to paint

longitudinal stripes on their roofing mats, but there is no confirmation of this.

A bone awl was used throughout for making holes for the sewing strands to go

through the sedge stems of the mats. The bone used was said to have been that of the

ostrich or the giraffe. No other tool is recorded among the San, but the Khoikhoi used a

needle for the actual sewing, and the Nama used a stone to smooth and sharpen the

needles and a wooden tool for smoothing and emphasizing the ridges made by the sewing.

By the eighteenth century, when most of the records began, iron awls with wooden

handles, and later, iron needles, were in use.

The material of importance beyond all others was the sedge—Cyperus and Scirpus

spp. The long smooth stems formed the foundation for the mats and for some sieves north

of the Orange River. They were sometimes twisted into a cord for binding or sewing. So

important were the sedges that it is possible that their availability may have been a factor,

additional to grazing, in directing the semi-nomadic movements of the Khoikhoi. Burchell

(1822: 279) notes an occasion when he was taken further than he wished to go to camp,

so that his people could gather the ‘mat rush’. It is also possible that the people in drier

areas may have obtained the material through trade.

Today, sedge continues to flourish in the south because circumstances have brought

about the disappearance of the mat-house. In Namaland, however, where the domed

hut has remained in use until recent times, the sedges have become scarce with over-

exploitation, and sacking and other materials have replaced sedge mats.

Throughout, Khoisan women seem to have been the manufacturers, except that the

men did the rare coiled sewn work and the twilled winnowing baskets, which appear to

have been a foreign borrowing.

Information for the Dama in earlier times is scant, and as the situation is now, it

would be hard to find any more information about basketwork that could be considered

truly Dama. Although the Dama settlement plan is similar to that of the Nama

(G. Lindhardt, pers. comm. 1989), they did not share the Khoikhoi type of mat-covered

dwelling, nor apparently did they use mats for any other purpose.

CONCLUSION

The country in which most of the Khoisan and the Dama lived was certainly not

lacking in material suitable for basketwork. But hunter-gatherers and pastoralists have

access to animal skins for carrying and for storing food and do not, therefore, need

baskets as containers. The use throughout of sewn mats for roofing by the Khoikhoi, and

298 ANNALS OF THE SOUTH AFRICAN MUSEUM

their more limited use by the San, must be considered a particular cultural trait, as indeed

were the huts themselves. The non-existence of the mat hut, or the mats, among the Dama

is equally significant. The sieves and traps, while important, are more localized, as are the

occasional borrowings from Bantu-spéakers.

ACKNOWLEDGEMENTS

Thanks are due to Margret Carey (London) and Mike Wilson (South African

Museum), for reading the manuscript, to Patricia Davison and Gerald Klinghardt for

suggestions made, to the Institute of Agricultural History, University of Reading, for

Figure 5, and to Bill van Rijssen for providing the photograph of Figure 9. June Hosford

was responsible for the other photographs. The late Leone Phillip drew most of the

techniques and Cedric Hunter drew some techniques and arranged the figures.

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paintings 1781-1784. Cape Town: Houses of Parliament.

LICHTENSTEIN, H. 1928. Travels in South Africa in the years 1803, 1804, 1805, 1806. Cape Town: Van

Riebeeck Society (Van Riebeeck Society Publications 10).

LICHTENSTEIN, H. 1930. Travels in South Africa in the years 1803, 1804, 1805, 1806. Cape Town: Van

Riebeeck Society (Van Riebeeck Society Publications 11).

Maccrong, I. D. 1937. A note on the tsamma and its uses among the Bushmen. Bantu Studies 11 (3):

251-252.

MackKENZIE, J. 1871. Ten years north of the Orange River. Edinburgh: Edmonton & Douglas.

MarsHaLL, L. 1976. The !Kung of Nyae Nyae. Cambridge, Mass.: Harvard University Press.

MorraT, R. 1842. Missionary labours and scenes in southern Africa. London: John Snow.

Oxtp, J. 1884. Angra Pequena und Gross Namaland. Elberfeld: Frederichs.

PassarGE, S. 1907. Die Buschmdnner der Kalahari. Berlin: Reimer.

RWSDALE, B. 1883. Scenes and adventures in Great Namaqualand. London: Woolmer.

ScHInZ, H. 1891. Deutsch Siidwest Afrika. Leipzig: Schwarz.

SCHREYER, J. 1681. Neue Ost-Indiaansche Reiz Beschreibung. Leipzig: C. J. Wohlfart. (Part reprinted

(1965) in translation as Johan Schreyer’s description of the Hottentots, 1679. In: Quarterly Bulletin

of the South African Library 19 (3): 88-101).

SCHULTZE, L. 1907. Aus Namaland und Kalahari. Jena: Fischer.

ScHwarz, E. H. L. 1928. Kalahari and its native races. London: Witherby.

SEINER, F. 1910. Die Buschmanner der Okavango- und Sambesigebietes der Nord-Kalahari. Globus 97

(22): 341-345, (23): 357-360.

SHAW, B. 1840. Memorials of South Africa. London: Mason.

SHaw, E. M. 1922. The basketwork of southern Africa. Part 1. Technology. Annals of the South African

Museum 100 (2): 53-248.

SmitH, A. 1834a. The diary of Dr. Andrew Smith. Edited by P. R. Kirby. Cape Town: Van Riebeeck

Society (Van Riebeeck Society Publications 20) (1939).

SmitH, A. 1834b. The diary of Dr. Andrew Smith. Edited by P. R. Kirby. Cape Town: Van Riebeeck

Society (Van Riebeeck Society Publications 21) (1939).

Smit, A. 1975. Andrew Smith’s journal. Edited by W. Lye. Cape Town: Balkema & South African

Museum.

SmitH, C. A. 1966. Common names of South African plants. Memoirs of the botanical survey of South

Africa. 35: 1-642.

SOMERVILLE, W. 1979. William Somerville’s narrative of his journey to the eastern Cape Frontier and

to Lattakoe, 1799-1802. Edited by Edna & Frank Bradlow. Cape Town: Van Riebeeck Society (Van

Riebeeck Society Publications 2nd series 10).

SPARRMAN, A. 1785a. A voyage to the Cape of Good Hope, towards the Antarctic polar circle, and round

the world: but chiefly into the country of the Hottentots and Caffres, from the year 1772, to 1776 1.

Dublin: White, Cash & Byrne.

SPARRMAN, A. 1785b. A voyage to the Cape of Good Hope, towards the Antarctic polar circle, and round

the world: but chiefly into the country of the Hottentots and Caffres, from the year 1772, to 1776 2.

Dublin: White, Cash & Byrne.

300 ANNALS OF THE SOUTH AFRICAN MUSEUM

Steyn, H. P. 1971. Aspects of the economic life of some nomadic Nharo Bushmen groups. Annals of

the South African Museum 56 (6): 275-323.

STeYN, H. P. & Du Pisani, E. 1984-1985. Grass seeds, game and goats: an overview of Dama subsistence.

Journal of the South West Africa Scientific Society 39: 37-52.

Stow, G. W. 1905. The native races of South Africa. London: Swan Sonnenschein.

TuHompson, G. 1827a. Travels and adventures in southern Africa. Edited by V. S. Forbes. Cape Town: Van

Riebeeck Society (Van Riebeeck Society Publications 48) (1967).

THompson, G. 1827b. Travels and adventures in southern Africa. Edited by V. S. Forbes. Cape Town: Van

Riebeeck Society (Van Riebeeck Society Publications 49) (1967).

THUNBERG, C. P. 1793. Travels at the Cape of Good Hope 1772-1775. Edited by V. S. Forbes. Cape

Town: Van Riebeeck Society (Van Riebeeck Society Publications, 2nd series 17) (1986).

VAN DER Merwe, H. M. 1984. Die vervaardiging van matjieshuise en inwoningspatrone in Leliefontein,

*n Namakwalandse landelike Kleurlinggebied. Koisis 5: 1-44.

VAN Ripren, B. 1918. Notes on some Bushman implements. Memoirs of the American Anthropological

Association 5: 75-97.

VEDDER, H. 1928. The Nama. In: The native tribes of South West Africa. Cape Town: Cape Times.

VepDpER, H. 1928. The Berg Damara. In: The native tribes of South West Africa. Cape Town: Cape Times.

VEDDER, H. 1934. Das alte Siidwest-Africa. Berlin: Warneck.

Von Francois, H. 1896. Nama und Damara. Magdeburg: Baensch.

WaLtTon, J. 1956. African village. Pretoria: Van Schaik.

WELLS, M. J. 1965. An analysis of plant remains from Scott’s Cave, in the Gamtoos Valley. South African

Archaeological Bulletin 20: 78-84.

Wikar, H. J. 1935. The journal of H. J. Wikar. Edited by E. E. Mossop. Cape Town: The Van Riebeeck

Society (Van Riebeeck Society Publications 15) (1935).

ABBREVIATIONS

Afr. Mus. Africana Museum, Johannesburg

SAM South African Museum, Cape Town

SAMUCT UCT collection now in South African Museum, Cape Town

Stockholm Folkens Museum, Stockholm

Wet University of Cape Town

Wit. Univ. Witwatersrand University, Johannesburg

BASKETWORK OF SOUTHERN AFRICA 301

Page numbers in bold refer to figures.

Acacia 281, 284, 291, 293, 296

Acanthosicyos 292

Aristida 281, 288

witli. 28. 287, 290, 291,

297

Bark 275, 281, 284

Barkly West 294

Baskets 281-282, 293-294,

295-296, 297

Bergdama 274

Bone 291, 297

Botletle River 281

Bredasdorp 287

Cord 297

Cyperus 275, 285, 288, 296, 297

Dama 274, 295-296, 297

Digitaria 283

Dobe 276, 282

Dwelling 274, 284

DyewZss, 291, 297

Eel-baskets 279, 279

Fish River, Great 274, 285, 286

Fish River, Namibia 279

Fish-traps 274, 279-281, 280,

292-293

Flask 294

Fransfontein 289, 290, 291

Gamtoos River 284, 293

Genadendal 287

Ghanzi 276, 283

Gonaqua 284, 286, 293

Gordonia 276, 277

Grewia 295

Griqua 284, 293

Hats 282-283, 283, 295

Herero 274

Huts 275, 276, 284, 285, 286,

288, 289, 295

Juncus 291

Kalahari 276, 283

Katbos 294

Khoikhoi 274, 275, 284-295,

297

Khoikhoi (Cape) 284, 293

Khoisan 273, 297

Korana 284, 286, 287, 293-294,

294, 295

‘Kung 276, 282

Kyllinga 283

Lake Chrissie 283

Lake Ngami 275

Lycium 294

Makers 276, 281, 282, 288, 291,

294, 295, 296

Materials 275, 278, 281, 282,

WS) PE Pero ZANE PEP,

293, 294, 295, 296

Mat-house 297

Mats 274-276, 284-292, 285,

289, 295, 297

Mbukushu 275

Men 281, 282, 294, 295, 296,

297

Nama 274, 284, 288, 289, 290,

291, 292, 294, 297

Namibia 274, 282

Nara 292

Needle 290, 297

Nharo 283

Nyae Nyae 276

Okavango 275

Olifants River 284

Orange River 274, 275, 279,

286, 287, 290, 292, 297

Ornamentation 275, 297

Ormaments 283

Oshikango 280

Painting of mats 275

Prieska 286

Neconks ZO, ZWD, Zsil, 222,

S25 vers 2x3, LD, LOD,

DB), YL, DS, PO, AI

Rhus 281, 293

Rope 284, 295

Rosyntjiebos 295

Salix 291

San 274-284, 283, 297

Sandfontein 277

Sarwa 281

NCUADES A383 Aer. 2911, 207)

Sedge 274, 275, 279, 284, 285,

288, 290, 291, 294, 297

Sieves 274, 276-279, 277, 296,

297

Sinew 275, 284

Stipagrostis 296

Storage bins 282

Strainers 292

String 290, 291

Swakop River 274

Taaibos 281

Mechniques; 2755, 278; 285 2382-

DRA DSi DIO 2922935.

295, 296

Tools 275, 287, 291, 291, 294

Topnaars 292

Traps 274, 297

Tsumkwe 276, 282

Tswana 281, 293

Walvis Bay 292

Wankie 275

Wildebeest tail hair 295

Windscreen 274, 275, 275

Women 275, 276, 281, 282, 287.

288, 291, 297

Xhosa 293

6. SYSTEMATIC papers must conform to the International 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.,

ete.

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°5S1'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.

E. M. SHAW

THE BASKETWORK OF SOUTHERN AFRICA.

| PART 2.

BASKETWORK OF THE KHOISAN:

AND THE DAMA

MK |

VOLUME 102 PART 9 FEBRUARY 1993 ISSN 0303-2515

way 1 3 1995

LIBRARIES

ANNALS

OF THE SOUTH AFRICAN

~~~ MUSEUM

CAPE ‘TOWN

INSTRUCTIONS TO AUTHORS

1. MATERIAL should be original and not published elsewhere, in whole or in part.

2. LAYOUT should be as follows:

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Title: informative but concise, without abbreviations and not including the names of new genera or species

Author’s(s’) name(s)

Address(es) of author(s) (institution where work was carried out)

Number of illustrations (figures, enumerated maps and tables, in this order)

(b) Abstract of not more than 200 words, intelligible to the reader without reference to the text

(d) Introduction

(e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents

(f) Summary, if paper is lengthy

(g) Acknowledgements

(h) References

(i) Abbreviations, where these are numerous.

3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced with

3 cm margins all round. First lines of paragraphs should be indented. Tables and a list of captions for

illustrations should be typed separately, their positions indicated in the text. All pages should be num-

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Major headings of the paper are centred capitals; first subheadings are shouldered small capitals;

second subheadings are shouldered italics; third subheadings are indented, shouldered italics. Further

subdivisions should be avoided, as also enumeration (never roman numerals) of headings and

abbreviations.

Footnotes should be avoided unless they are short and essential.

Only generic and specific names should be underlined to indicate italics; all other marking up

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4. ILLUSTRATIONS should be reducible to a size not exceeding 12 x 18cm (19 cm including

caption); the reduction or enlargement required should be indicated (and preferably uniform); orig-

inals larger than 35 x 47 cm should not be submitted; photographs should be rectangular in shape and

final size. A metric scale should appear with all illustrations, otherwise magnification or reduction

should be given in the caption; if the latter, then the final reduction or enlargement should be taken

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All illustrations, whether line drawings or photographs, should be termed figures (plates are not

printed; half-tones will appear in their proper place in the text) and numbered in a single series. Items

of composite figures should be designated by capital letters; lettering of figures is not set in type and

should be in lower-case letters. If Letraset is used authors are requested to use Helvetica-style letter-

ing, if possible.

The number of the figure should be lightly marked in pencil on the back of each illustration.

5. REFERENCES cited in text and synonymies should all be included in the list at the end of the

paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable):

(a) Author’s name and year of publication given in text, e.g.:

“Smith (1969) describes. . .’

‘Smith (1969: 36, fig. 16) describes... .”

‘As described (Smith 1969a, 1969b; Jones 1971)’

‘As described (Haughton & Broom 1927)...’

‘As described (Haughton et al. 1927)...”

Note: no comma separating name and year

pagination indicated by colon, not p.

names of joint authors connected by ampersand

et al. in text for more than two joint authors, but names of all authors given in list of references.

(b) Full references at the end of the paper, arranged alphabetically by names, chronologically within

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

that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a).

For books give title in italics, edition, volume number, place of publication, publisher.

For journal article give title of article, title of journal in italics (according to the World list of scientific periodicals. 4th ed.

London: Butterworths, 1963), series in parentheses, volume number, part number in parentheses, pagination (first and

last pages of article).

Examples (note capitalization and punctuation)

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

FiscHer, P. H. 1948. Données sur la résistance et de la vitalité des mollusques. Journal de conchyliologie 88 (3): 100-140.

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

expérimentale et générale 74 (33): 627-634.

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

Magazine of Natural History (13) 2 (17): 309-320.

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

the Bingham Oceanographic Collection, Yale University 17 (4): 1-51.

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

pologische Ergebnisse einer Forschungsreise im westlichen und zentralen Stid-Afrika ausgeftihrt in den Jahren

1903-1905 4 (15). Denkschriften der medizinisch-naturwissenschaftlichen Gesellschaft zu Jena 16: 269-270.

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 102 °&#Band

February 1993 Februarie

Part 9 Deel

HOW MANY SPECIES OF DITCTODON

WERE THERE?

GILLIAN M. KING

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na gelang van die

beskikbaarheid van stof

Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad 8000

OUT OF PRINT/UIT DRUK

1, 2(1-3, 5-8), 3(1—2, 4—5, 8, t—p.i.), 51-3, 5, 7-9),

6(1, t.—p.i.), 711-4), 8, 911-2, 7), 10(1-3), 1101-2, 5, 7, t—p.i.),

141-3), 15(4—-5), 24(2, 5), 27, 311-3), 32(5), 33,

36(2), 43(1), 45(1), 67(5), 84(2)

Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 142 3

Printed in South Africa by In Suid-Afrika gedruk deur Die

The Rustica Press, Pty., Ltd., Rustica-pers, Edms., Bpk.,

Old Mill Road, Ndabeni, Cape Old Millweg, Ndabeni, Kaap

D1873

HOW MANY SPECIES OF DIJCTODON WERE THERE?

GILLIAN M. KING

Department of Karoo Palaeontology, South African Museum, Cape Town

(With 3 figures and | table)

[MS accepted 26 October 1992]

ABSTRACT

Twenty species of the dicynodont genus Diictodon have been described. Although the genus is

distinguishable by reliable characters, the validity of the species-level taxonomy is questionable. The

present work re-evaluates the existing species and concludes that previous authors have not been able to

produce convincing species-level characters. A sample of well-preserved and well-prepared skulls in the

South African Museum is used to search for any other possible species-specific characters, but none is

found.

It is concluded that only one species of the genus, Diictodon galeops, is justifiable. The specific

longevity of this dicynodont is noted.

CONTENTS

PAGE

INTROGUC HOME ee eerste eerily ce rari Oe ey ae be 303

SPECIESHOMD ietodOnien rie ear pti se te eile wk Stas 307

INSSESSINEM@ Ol ChanaCtehS ys) ah 4 ies sass cae eee ees 313

SUIT eliny are ieep mee Uh ener 2ty RM iah tomtccrnia Clits sas 320

CON ClUIS(OME RE eee ek eos one ee ee eed Wala 's 320

ZACKMOWAE USENET (StMeua as eens ve ets Maat Me se oh sic yssaulp tiles 3

INGERCH COSI EN RR Ene ee aie crn a 8 a BAI

ADORE VAMONS merry ee rete ee ome a) ee ee tte A 5 AG 328

INDY XEINGIOS 9 “G 3-csto. 6 ei 8 as ace SCM Chances CREO RR aE 323)

INTRODUCTION

The dicynodont genus Diictodon is distinctive and well-characterized. Descriptions

of the genus have been given by Cluver & Hotton (1981), Cluver & King (1983) and King

(1988). The most distinctive feature of the skull is the prominent maxillary caniniform

process that is clearly demarcated from the anterior palatal rim by a deep notch (Figs 1,

2B). This feature is found in conjunction with small palatines that do not meet the pre-

maxilla on the palatal surface, a narrow intertemporal region in which the postorbitals

tend to cover the parietals (Fig. 2), prominent dentary tables of which the medial edge

stands proud as a ridge, and a weakly-developed lateral dentary shelf (Fig. 3). This suite

of characters permits easy identification of members of the genus.

Diictodon seems to have been a common element in the Karoo palaeoenvironment.

Many specimens of the genus are known (Smith 1980), together with trackways and

burrows that are probably attributable to this form (Smith 1987). Excellent postcranial

material is also available. Despite this, no detailed functional study of the genus has been

made, nor has a species-level revision of the taxonomy of the genus been carried out.

303

Ann S. Afr. Mus. 102 (9), 1993: 303-325, 3 figs, 1 table.

304 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 1. Three specimens of Diictodon in lateral view to show

the distinctive square-cut caniniform process. A. SAM-—10086.

B. SAM-K5105. C. SAM-K7730. Scale = 30 mm.

Owing to the wealth of material available, Diictodon offers an opportunity to examine the

effects of intraspecific variation in a fossil species and detailed studies of this aspect of

the genus are in progress (King in prep.). Stratigraphic information for specimens from

the South African Karoo Basin is also available and becoming increasingly refined (Kitch-

ing 1977; Keyser & Smith 1978; Rubidge in press), so the genus also offers the possibility

of studying changes in osteological morphology in time and space.

However, a prerequisite of such studies is an understanding of the species-level

taxonomy of the genus. Here the existing species are reviewed and their validity, based

HOW MANY SPECIES OF DIICTODON WERE THERE? 305

Fig. 2. Diictodon galeops SAM—K7673. A. Skull in dorsal view. B. Skull in palatal view.

Stipple indicates area of matrix. Scale = 30 mm.

306 ANNALS OF THE SOUTH AFRICAN MUSEUM

mdt Idt ds mf

Fig. 3. Diictodon galeops SAM-K7730. A. Lower jaw in dorsal view. B. Lower jaw in lateral view.

Stipple indicates area of matrix. Scale = 30 mm.

TABLE 1

Biostratigraphic schemes for the Karoo Sequence.

A. After Kitching (1977).

Triassic Cynognathus bio-zone

Lystrosaurus bio-zone

Permian Daptocephalus bio-zone

Cistecephalus bio-zone

Tapinocephalus bio-zone

B. After Rubidge (in press).

Triassic Cynognathus-—Diademodon assemblage zone

Lystrosaurus-Procolophon assemblage zone

Permian Dicynodon-Theriognathus assemblage zone

Cistecephalus-Aulacephalodon assemblage zone

Tropidostoma-Endothiodon assemblage zone

Pristerognathus—Diictodon assemblage zone

Tapinocephalus—Bradysaurus assemblage zone

Eodicynodon-Tapinocaninus assemblage zone

HOW MANY SPECIES OF DIJCTODON WERE THERE? 307

on the characters used by the naming authors, is assessed. In addition, a collection of

well-preserved and well-prepared skulls in the South African Museum is used to investi-

gate whether there are any species-specific characters that have been missed by previous

authors, and to facilitate a revision of the existing species.

Since it was hoped to draw some conclusions concerning the stratigraphic distribution

of Diictodon species, the most refined biostratigraphic scheme for the Beaufort Sequence

(Rubidge in press) has been used to assign the South African Museum sample skulls to

biostratigraphic assemblage zones (see Table 1). (I am extremely grateful for the help of Dr

R. M. H. Smith, South African Museum, in this regard.) However, the available locality

information for the existing type specimens does not always allow assignment of those

specimens to an assemblage zone in Rubidge’s scheme, and their assignments to bio-zones in

the previous stratigraphic scheme, as published in Kitching (1977), have been retained.

SPECIES OF DITCTODON

King (1988) gave a list of 20 species of the genus. Although it is clear that these

forms all belong within the genus (possessing the distinctive maxillary notch and other

features noted above), the validity of the species involved requires re-assessment.

The type species is Diictodon galeops, described by Broom (1913).

Diictodon galeops Broom, 1913

Diictodon galeops Broom, 1913: 453, fig. 15.

The type, AMNH 5308, is a skull from Slachtersnek, Somerset East District, Cape

Province, South Africa. This locality is recorded as being in the Cistecephalus bio-zone

of the Teekloof Formation, Karoo Sequence by Kitching (1977).

The feature which Broom (1913) felt distinguished this genus from the otherwise

similar Dicynodon was the large size and peculiar shape of the preparietal. This bone

surrounds the pineal opening in Diictodon galeops, whereas Broom considered that the

usual state in Dicynodon was for the preparietal to lie in front of the pineal. On this basis

he transferred the Dicynodon species D. kolbei and D. alticeps to Diictodon, and noted

that this might also be warranted for Dicynodon tigriceps. However, the former two

Species are now regarded as belonging in the genus Oudenodon, and the latter in

Aulacephalodon (King 1988).

Toerien (1953) discussed variation in the shape and position of the preparietal in

Diictodon (Dicynodon) grimbeeki and Diictodon sollasi. He concluded that sutural pattern

should be used as a generic or specific character only with caution, since in a sample of

otherwise similar skulls from one locality, continuous variation in pattern and size of the

preparietal was seen. However, Cluver & Hotton (1981) gave a revised diagnosis of the

genus based on more reliable characteristics also present in the type of Diictodon galeops.

Of the remaining 19 species, only one, Diictodon sesoma Watson, 1960, was origi-

nally described as Diictodon, the others having been previously included in the genera

Dicynodon, Oudenodon and Emydorhynchus.

Diictodon sesoma Watson, 1960

Diictodon sesoma Watson, 1960: 142, fig. 2.

The holotype of this species is a skull, lower jaw and postcranial skeleton from

Buffel’s River, Orange Free State, South Africa (UMZC R314). Watson (1960) included

308 ANNALS OF THE SOUTH AFRICAN MUSEUM

it in the genus Diictodon, because the pineal opening is surrounded by the preparietal, but

he felt that the material warranted allocation to a new species, owing to its shorter and

broader skull, wider occiput, relatively wider intertemporal region, and large canine.

The intertemporal region, as illustrated by Watson, is rather wide for Diictodon

(based on the sample of specimens in the South African Museum), although this appear-

ance can result when the skull roof is weathered and the postorbitals obliterated. The

specimen does appear to have a maxillary notch, however, although the maxillary rim in

front of the tusk appears to be broken. The pineal is slightly raised above the skull roof,

another feature also seen in some specimens attributed to Diictodon.

Diictodon feliceps (Owen, 1876)

Dicynodon feliceps Owen, 1876: 45, pl. 43.

Rhachiocephalodon feliceps (Owen, 1876) Seeley, 1898: 108.

Diictodon feliceps (Owen, 1876) Cluver & Hotton, 1981: 125.

The holotype, BMNH 47052, is from Fort Beaufort, Cape Province, South Africa

(Cistecephalus bio-zone, Kitching 1977).

Owen (1876) felt that there was no difficulty in accommodating this specimen in the

genus Dicynodon as then known but considered that a new species was warranted, based

on skull proportion, suture pattern and curvature of the canine tusk. Owen (1876, pl. 43

(fig. 1)) showed a clear notch in front of the caniniform tusk, a characteristic indicating

that the specimen can be accommodated within the genus Diictodon. The pinched-in

nature of the temporal region, with postorbitals approaching each other over the parietals,

is also typical of Diictodon.

Diictodon jouberti (Broom, 1905)

Dicynodon jouberti Broom, 1905: 331.

Sintocephalus jouberti (Broom, 1905) van Hoepen, 1934: 93.

Diictodon jouberti (Broom, 1905) Cluver & Hotton, 1981: 127.

The holotype skull (SAM-—695) is from Gouph (Koup) Tract, Beaufort West District,

South Africa. The locality is in a low to middle horizon of the Tapinocephalus bio-zone

(Kitching 1977).

Broom (1905) considered that the noteworthy features of the skull were that the

parietal, frontal and upper part of the nasal are practically in one plane; the interparietal

portion is about equal in breadth to the interorbital; and the jugal arch is unusually deep

in the region of the postorbital bar.

Broom (1905) described a series of skulls, some with large and others with smaller

canines. He considered this to be a sexual difference, since the two forms do not differ

consistently in size.

The type has a distinctly notched maxilla in front of the tusk typical of Diictodon.

Diictodon psittacops (Broom, 1912)

Dicynodon psittacops Broom, 1912: 869, pl. 92.

Diictodon psittacops (Broom, 1912) Cluver & Hotton, 1981: 129.

The holotype skull and skeleton (AMNH 5534) are from the Beaufort West com-

monage, Cape Province, South Africa. This locality is in the Cistecephalus bio-zone

according to Kitching (1977).

HOW MANY SPECIES OF DIICTODON WERE THERE? 309

Broom (1912) considered the distinguishing features of this new species to be the

narrow nasals, thickened upper part of the nasals, elevated prefrontal region of the orbital

margin, broad and flat frontal region, pineal foramen that is situated on an elevated

preparietal, postorbitals that approach each other behind the pineal opening and form a

ridge, small downwardly and forwardly directed tusk, broad and deep front portion of the

lower jaw, and the very small intramandibular foramen.

Diictodon ictidops (Broom, 1913)

Dicynodon ictidops Broom, 1913: 466, figs 5-6.

Pylaecephalus ictidops (Broom, 1913) van Hoepen, 1934: 93.

Diictodon ictidops (Broom, 1913) Cluver & Hotton, 1981: 130.

The holotype, AMNH 55110, is a skull from Beaufort West commonage, Cape

Province, South Africa (Cistecephalus bio-zone according to Kitching 1977).

Broom (1913) described a number of small skulls from the same locality as being

narrow with large, rounded nostrils. He noted that the septomaxilla does not show on the

side of the skull and that the tusks are variably expressed, being absent in at least one

specimen. The zygomatic arch beneath the postorbital bar was described as being very

deep. Broom noted that the preparietal does not surround the pineal foramen.

Diictodon palustris (Broom, 1913)

Emydorhynchus palustris Broom, 1913: 456, fig. 19.

Diictodon palustris (Broom, 1913) Cluver & Hotton, 1981: 130.

The holotype (AMNH 5512) is a skull from New Bethesda, Graaff-Reinet District,

Cape Province, South Africa. This is in the Daptocephalus bio-zone (Kitching 1977).

Broom (1913) noted the short preorbital part of the skull, the absence of tusks, the

apparent absence of the septomaxilla (or its reduction), the large preparietal, and the large

postorbital, which was considered to be unusual in that it was broad in front and narrow

_ posteriorly.

Diictodon testudirostris (Broom & Haughton, 1913)

Dicynodon testudirostris Broom & Haughton, 1913: 36.

Pylaecephalus testudirostris (Broom & Haughton, 1913) van Hoepen, 1934: 93.

Diictodon testudirostris (Broom & Haughton, 1936) Cluver & Hotton, 1981: 130, figs 22-26.

The holotype, SAM—2354, is a skull from Dunedin, Beaufort West District, Cape

Province, South Africa, a locality in the Cistecephalus bio-zone (Kitching 1977).

Distinguishing characters noted by Broom & Haughton (1913) are the very short

beak, the maxilla which almost reaches the orbit, the apparent absence of the septo-

maxilla, the reduced lachrymal and prefrontal, the large postorbitals that meet in the

midline, the large preparietal almost entirely in front of the pineal opening, the quadrato-

jugal that is not ankylosed to the quadrate.

Diictodon sollasi (Broom, 1921)

Dicynodon sollasi Broom, 1921: 648, figs 28-29.

Pylaecephalus sollasi (Broom, 1921) van Hoepen, 1934: 93.

Diictodon sollasi (Broom, 1921) Cluver & Hotton, 1981: 132.

The holotype (SAM-—7420) is from Biejiespoort, Victoria West District, Cape Prov-

ince, South Africa, in the Cistecephalus bio-zone according to Kitching (1977).

310 ANNALS OF THE SOUTH AFRICAN MUSEUM

Broom’s (1921) description was based on many skulls from the same locality. He

noted that immediately behind the plane of the nostrils there is a thickening of the bones,

which forms a low, round, button-like boss in the midline. Other features to which atten-

tion was drawn were the canine that had a thickened ridge above and behind it, and the

small preparietal. Broom (1921) distinguished Dicynodon sollasi from other species on

the basis of such characters as the delicate build of the skull, the feebler tusk, the degree

to which the postorbitals overlap the parietals, and the size of the preparietal.

Diictodon haughtonianus (von Huene, 1931)

Dicynodon haughtonianus von Huene, 1931: 30, fig. 25.

Diictodon haughtonianus (von Huene, 1931) King, 1988: 121.

The holotype (UT Von Huene 1931 Abb 25) is a skull and anterior part of lower jaw

from Bloukrans, Prince Albert, South Africa, in the Tapinocephalus bio-zone (Kitching

1977).

Von Huene (1931) distinguished this species from others by its smaller nasal opening

which lies further from the maxillary rim, the caniniform process which is more back-

wardly directed, the delicate postorbital bar, the shorter and wider postfrontal, and the

shape of the preparietal.

Diictodon rubidgei (Broom, 1932)

Dicynodon rubidgei Broom, 1932: 189, fig. 62.

Pylaecephalus rubidgei (Broom, 1932) van Hoepen, 1934: 93.

Diictodon rubidgei (Broom, 1932) Cluver & Hotton, 1981: 133.

The holotype (BMNH 47081) is a skull from the Graaff-Reinet commonage, Cape

Province, South Africa. This is in the Cistecephalus bio-zone (Kitching 1977).

Broom (1932) did not specifically allude to any distinguishing characters, but he

mentioned the large preparietal and small pineal opening, and the large postfrontal. He

noted that the postorbital is moderately large but its junction with the squamosal is not as

far back as in most species of Dicynodon.

Diictodon grimbeeki (Broom, 1935)

Dicynodon grimbeeki Broom, 1935: 7, figs 6-7.

Diictodon grimbeeki (Broom, 1935) Cluver & Hotton, 1981: 133.

The holotype (TM 253) is a skull from Leeuwpoort, Beaufort West District, Cape

Province, South Africa. The locality is in the Cistecephalus bio-zone (Kitching 1977).

Broom (1935) described a sample of 19 skulls from the locality and noted that the

morphology of the preparietal varies within the sample. A small boss behind the pineal

foramen was noted, as was the smooth rounded median boss on the snout formed by the

nasals and the premaxilla. Broom distinguished male and female skulls, noting that the

male skulls are bigger than those of the females, due to the much larger snouts of the

former. The males were considered to have tusks; the females no or rudimentary tusks.

Diictodon nanus (Broom, 1936)

Dicynodon nanus Broom, 1936: 379, fig. 25A.

Diictodon nanus (Broom, 1936) Cluver & Hotton, 1981: 133.

The holotype (TM 268) is a skull from Houd Constant, Graaff-Reinet District, Cape

Province, South Africa, in the Cistecephalus bio-zone (Kitching 1977).

HOW MANY SPECIES OF DIICTODON WERE THERE? a Et

Broom considered this to be a juvenile form. He noted the complete absence of

postfrontals, that the postorbitals do not meet over the parietals, and that a developing

canine (or possibly postcanine) is present behind the feebly developed caniniform

processes.

Diictodon huenei (Broili & Schroder, 1937)

Dicynodon huenei Broili & Schréder, 1937: 130, figs 14 (preoccupied).

Dicynodon broilii (Broili & Schréder, 1937) Boonstra, 1948: 57.

Oudenodon huenei (Broili & Schroder, 1937) Toerien, 1953: 97.

Anomodon huenei (Broili & Schréder, 1937) Keyser, 1975: 74, fig. 28.

The holotype (BSP 1934 VIII 46) is a skull from La-de-da, Beaufort West District,

Cape Province. The locality is in a high horizon of the Tapinocephalus bio-zone (Kitching

EST 7).

The main distinguishing features of this form mentioned by Broili & Schréder (1937)

are its small size (total skull length 75 mm), the broad intertemporal and interorbital

regions, the low bosses over the nostrils, the flat skull roof, the well-developed post-

frontals, the large preparietal, the postorbital incompletely covering the parietals, and the

beak-like dentary symphysis.

Diictodon broomi (Broili & Schroder, 1937)

Dicynodon broomi Broili & Schréder, 1937: 132, figs 5-13.

Diictodon broomi (Broili & Schroder, 1937) Cluver & Hotton, 1981: 134.

The holotype (BSP 1934 VIII 47a and b) comprises two skulls, one tusked, one

tuskless from La-de-da, Beaufort West District, Cape Province. The locality is in a high

horizon of the Tapinocephalus bio-zone (Kitching 1977).

The main distinguishing features of the species mentioned by Broili & Schréder

(1937) are the skull (length 91 mm), which is slightly larger than that of the other

specimens described in the same paper, the strongly developed preorbital region of the

skull with a median boss between the nasal openings, the interorbital distance which is

slightly broader than that of the intertemporal, the flat skull roof, the pineal foramen

surrounded by a ring of bone, the large tusk, and the postorbitals which converge behind

the pineal opening, covering the parietals.

Diictodon grossarthi (Broili & Schréder, 1937)

Dicynodon grossarthi Broili & Schréder, 1937: 161, figs 14-18.

Diictodon grossarthi (Broili & Schréder, 1937) Cluver & Hotton, 1981: 134.

The holotype (BSP 1934 VIII 48) is a skull from La-de-da, Beaufort West District,

Cape Province, in a high horizon of the Tapinocephalus bio-zone (Kitching 1977).

The main distinguishing features mentioned by Broili & Schroder (1937) are the

delicate snout, the convex skull roof, the absence of tusks, the low median boss over the

nostrils, the delicate postorbital bar, the palatine which reaches relatively far forward on

the palate, and the oval depression at the junction of the ectopterygoid, jugal and palatine.

The skull length is approximately 100 mm.

Diictodon anneae (Broom, 1940)

Dicynodon anneae Broom, 1940: 181, fig. 23.

Dicynodon whitsonae Toerien, 1954: 937.

Diictodon whitsonae (Toerien, 1954) Cluver & Hotton, 1981: 134.

Diictodon anneae (Broom, 1940) comb. nov.

Si ANNALS OF THE SOUTH AFRICAN MUSEUM

It should be noted that Toerien (1954: 937) renamed this species Dicynodon whit-

sonae, believing that Dicynodon anneae was preoccupied by the Russian form described

by Amalitsky (1922) as Dicynodon annae. Cluver & Hotton (1981: 134) followed Toe-

rien’s species reassignment and, in addition, referred Dicynodon whitsonae to Diictodon.

King (1988) also accepted Diictodon whitsonae as the valid species. However, these

authors failed to notice that the spelling of Broom’s original species, anneae, was distinct

from that of the Russian form, annae. The original species name given by Broom (1940)

is therefore valid. It is here referred to the genus Diictodon as Diictodon anneae.

The holotype (RC 42) is a skull from Wellwood, Graaff-Reinet District, Cape Prov-

ince, South Africa. The locality is in the Cistecephalus bio-zone (Kitching 1977).

The only noteworthy feature of this small tusked specimen from Broom’s (1940)

description is the fact that the postorbitals are well developed, but the posterior processes

are shorter than in most species.

Diictodon pseudojouberti (Boonstra, 1948)

Dicynodon pseudojouberti Boonstra, 1948: 60.

Diictodon pseudojouberti (Boonstra, 1948) Cluver & Hotton, 1981: 134.

The holotype (SAM-—774) is a skull from Prince Albert Road, Cape Province, South

Africa, in the Tapinocephalus bio-zone (Kitching 1977).

From Boonstra’s (1948) description, the points of distinction of this species appear

to be the relatively high and fairly broad skull, the fairly weak snout, the interorbital width

being approximately equal to the intertemporal, the convex intertemporal and interorbital

regions, the large and roughly oval preparietal, the well-developed postfrontal, and the

postorbitals that are large and overlap the parietals.

Diictodon vanderhorsti (Toerien, 1953)

Dicynodon vanderhorsti Toerien, 1953: 91, fig. 60.

Diictodon vanderhorsti (Toerien, 1953) Cluver & Hotton, 1981: 135.

The holotype (BPI 175) is a skull from Antjiesfontein, Prince Albert District, South

Africa, in a low horizon of the Tapinocephalus bio-zone (Kitching 1977).

Toerien (1953) noted that a low boss is present over the nostrils, and a low bony ring

surrounds the pineal foramen. He discussed variation within dicynodonts and suggested

that replacement canines are only found in young individuals, that nasal bosses increase

with the size of the individual, and that the pineal boss is characteristic of old males.

Diictodon tienshanensis (Sun, 1973)

Dicynodon tienshanensis Sun, 1973: 56, figs 1-6.

Diictodon tienshanensis (Sun, 1973) Cluver & Hotton, 1977: 179, pls 73-74.

The holotype (IVPP V.3260) is a skull from the north foot of the Tienshan moun-

tains, XinJiang Province, China. This is in the Upper Jijicao Group, perhaps equivalent to

either the Cistecephalus or Daptocephalus bio-zone (King 1992).

Sun (1973) did not note any features of the skull that might distinguish this species

from any other, apart from in the palate. Here the large interpterygoid fossa and long,

narrow interpterygoid foramen are specified. In addition, Sun noted that the palatine forms

a nodule at the anterior end, which projects internally and constricts the anterior end of

the internal nares, and a small boss also projects at the inner posterior side of the maxillary

process.

HOW MANY SPECIES OF DIJICTODON WERE THERE? Cu ie)

ASSESSMENT OF CHARACTERS

The features that previous authors have felt to be important in terms of recognizing

the above species can be categorized as follows:

Features of the pineal and preparietal

1.1 Pineal foramen surrounded by a ring of bone

1.2 Pineal foramen situated on an elevated preparietal

1.3. Size and shape of the preparietal

1.4 Position of the preparietal relative to the pineal opening

General skull features

2.1 Skull roof flat, concave or convex

2.2 Height of skull

2.3. Breadth of skull

2.4 Narrowness of skull

2.5 Length of skull

2.6 Whether skull is robust or delicate

2.7 Width of occiput

2.8 Width of intertemporal region relative to skull length

2.9 Width of intertemporal region relative to interorbital

2.10 Length of preorbital part of the skull

2.11 Whether snout is delicate or robust

Features relating to the nasals and nostrils

3.1 Width of nasals

3.2 Size and shape of nostrils

3.3. Position of nostril relative to maxillary rim

Features of the postorbital bones

4.1 Whether and how far the postorbitals approach each other behind the pineal opening

4.2 Shape of the postorbital

Features of the caniniform process and tusks

5.1 Direction of the caniniform process or tusk

5.2 Presence and size of canine tusks

Features relating to skull bosses

6.1 Whether a boss is present behind the pineal foramen

6.2 Whether separate nasal bosses are present

6.3 Whether median boss over the nostrils is present

Features of the lower jaw

7.1 Whether the dentary symphysis is beak-like

7.2 Whether front part of lower jaw is broad and deep

7.3 Size of intramandibular foramen

314 ANNALS OF THE SOUTH AFRICAN MUSEUM

Features relating to individual bones or specific areas of the skull

8.1 The septomaxilla does not show on the side of the skull

8.2 Presence, absence or reduction of the septomaxilla

8.3. Depth of the jugal arch

8.4 The prefrontal region of orbital margin is elevated

8.5 The frontal region is broad and flat

8.6 Maxilla almost reaches the orbit

8.7. The prefrontal is reduced

8.8 Quadrato-jugal not ankylosed to the quadrate

8.9 Whether the postorbital bar is delicate

8.10 Size and shape of the postfrontal

Features of the palate

9.1 The palatine reaches relatively far forward on the palate

9.2 Presence of an oval depression at the junction of the ectopterygoid, jugal and palatine

9.3 Size of interpterygoid fossa

9.4 Length and breadth of interpterygoid foramen

9.5. Whether a nodule is present on the anterior end of the palatine

9.6 Whether a small boss is present on the inner posterior side of the maxillary process

Several of these features may be dismissed at once, since it is now known that they

do not constitute valid specific characters, either because they vary within otherwise

similar forms, or because they are widespread among many different forms, or are likely

to have been produced by distortion or bad preservation.

The size and shape of the preparietal and its position with respect to the pineal

opening (1.3, 1.4) are examples of known variation within a group (Toerien 1953) and

may be dismissed.

Features 2.1—2.5, 2.7, 2.8, 3.2, 5.1, 9.3, and 9.4 may all be affected by distortion and

are therefore unreliable.

Features 8.1, 8.5, 8.6 and 8.7 occur in all specimens of Diictodon investigated.

Feature 8.8 (quadratojugal not ankylosed to the quadrate) is now known to be the case in

the majority of dicynodonts (see King 1981).

Feature 4.1 (whether and how far the postorbitals approach each other behind the

pineal opening) varies within the group of species discussed above. The postorbitals

usually approach each other in the midline but they do not always cover the parietals

completely. Nevertheless, the ‘pinched-in’ appearance of the intertemporal bar seems to

be a typical feature of Diictodon, as noted by Cluver & Hotton (1981).

Some features (2.6, 2.11, 8.9) are based on subjective non-quantitative assessments,

e.g. the suggestion that the skull is delicate. Broom (1921) used this feature to distinguish

between Diictodon feliceps and Diictodon sollasi but, in any case, the two skulls are of

different sizes and it might be expected that the larger would be less delicate. I will not

use these more subjective features further in the discussion.

The remaining features are discussed further below. I have used a sample of well-

preserved and well-prepared specimens in the South African Museum (Appendix 1) to

investigate the states of the features given above: to determine whether the features are

identifiable consistently, how often they occur, and how much variation they exhibit.

HOW MANY SPECIES OF DIJICTODON WERE THERE? BUS)

Of the 37 skulls and jaws in this sample, two are from the Tapinocephalus—

Bradysaurus assemblage zone, four from the Pristerognathus—Diictodon, three from the

Cistecephalus—Aulacephalodon, two from the Dicynodon-Theriognathus, and the remain-

der from the Tropidostoma—Endothiodon assemblage zone. These numbers do not repre-

sent the abundance of specimens per stratigraphic zone in the field, nor probably in life,

but reflect the composition of the collections available for study.

Pineal foramen surrounded by a ring of bone (1.1); and pineal foramen situated on an

elevated preparietal (1.2)

Out of the South African Museum sample of 37 skulls, 15 specimens show some

indication of a ring-like structure or bulbous area around the pineal foramen. (The latter

seems to be what is meant by elevated preparietal in previous authors’ descriptions of

Diictodon sesoma and Diictodon psittacops.) The two conditions do not seem to be

independent. In larger specimens the ring around the pineal opening becomes swollen,

giving the bulbous or elevated structure.

It is possible that, in two specimens where the skull roof is slightly weathered, the

ring may have been lost if it had been present but very slightly developed in the first

place.

The presence of this structure does not correlate with any particular assemblage zone,

the 16 specimens coming from the Tapinocephalus—Bradysaurus assemblage zone (1), the

Pristerognathus—Diictodon assemblage zone (2), and Tropidostoma—Endothiodon assem-

blage zone (13). Specimens without the feature are also known from the Tapinocephalus—

Bradysaurus and Tropidostoma—Endothiodon zones.

The feature correlates in some way with size, since the seven largest specimens

(approximately 90-120 mm skull length) in the sample all have it, but in smaller speci-

mens the feature may or may not be present. The smallest skull in the sample (skull length

47 mm) does not have the feature. The ring or bulbous area tends to be better developed

in the larger specimens. Both tusked and tuskless specimens have the feature. Since this

is a feature that may well be correlated with size of skull (as Toerien (1953) also pointed

out), it is not a useful feature for delineating species.

Width of intertemporal region relative to interorbital (2.9)

Comments on the relative intertemporal and interorbital widths are made by previous

authors for several species. This feature is dependent on the measurement of the intertem-

poral width, which is in turn dependent on how much the postorbitals overlie the parietals.

This may be affected by distortion in two ways. The intertemporal region may be pinched

together, forcing the postorbitals to approach each other more closely, forming ridges

above the surface of the parietals. Secondly, it may be possible for post-mortem compres-

sion to cause the postorbitals to slide over the parietals, presumably either medially or

laterally, reducing or increasing the intertemporal distance, respectively. The evidence for

this is that some specimens in the South African Museum sample have a layer of matrix

between the postorbitals and parietals, indicating that there must be space between the two

bones. Because of this it is difficult to measure the true intertemporal width and use of

this ratio as a specific character is not advisable, as Keyser (1975) pointed out.

316 ANNALS OF THE SOUTH AFRICAN MUSEUM

Length of preorbital part of the skull (2.10)

There are two problems with using this feature for taxonomic purposes. The first is

that it is not known how the preorbital part of the skull would be affected by distortion

and whether it would react in the same way to compressive or tensional forces as the rest

of the skull. If, for example, it were more resistant to compression, then preorbital/skull

length ratios would be unreliable.

Secondly, Toerien (1953) gave evidence that the snout length increases relatively

more quickly with increasing skull length, and so this may be another feature attributable

to age difference.

Width of nasals (3.1)

Broom (1912) suggested that the nasals of Diictodon psittacops are so narrow that

the nostrils face almost directly upwards. However, this is an unreliable character, since

dorso-ventral flattening of the skull due to distortion would produce more upwardly-

directed nostrils, and also the antero-dorsal margin of the nostril is very thin and may be

worm away, making the nasals narrower.

Position of nostril relative to maxillary rim (3.3)

This feature will be affected by distortion and preservation. If the skull is subject to

antero-posterior compression, the snout becomes flattened posteriorly and the nostril

appears to lie nearer the anterior surface of the skull. If the maxillary rim of the snout is

worn away, the nostril will appear to be nearer the ventral maxillary rim. Wearing away

of the maxillary rim is not always obvious, since the rim is thin and the bone has an

unfinished appearance and does not always show a break cleanly.

Whether and how far the postorbitals approach each other behind the pineal opening (4.1)

The tendency for the postorbitals to approach each other in the dorsal midline,

covering the parietals in the intertemporal region has been used as a diagnostic character

for Diictodon (Cluver & Hotton 1981; Cluver & King 1983). Although this tendency is

present, the degree to which the postorbitals cover the parietals is variable. As mentioned

under 2.9, the intertemporal region may be affected by distortion such that the configura-

tion of the postorbitals changes. Because of this the degree to which the postorbitals meet

each other over the parietals should not be given undue emphasis.

Shape of the postorbital (4.2)

Broom (1913) noted that the postorbital in Diictodon palustris differs in shape from

anything known in dicynodonts, being so broad in front as to roof over part of the

temporal fenestra, but narrowing rapidly behind. This condition is seen in other specimens

(SAM-K7673, SAM-K7674), where it would appear that the thinner medial part of the

postorbital behind the pineal opening has broken off on both sides. This gives the appear-

ance of the postorbital narrowing posteriorly.

Presence and size of canine tusks (5.2)

Various conditions of the caniniform tusks have been noted by previous authors

(Broom 1905, 1913, 1921, 1935; Broili & Schroder 1937; Toerien 1953). They have been

HOW MANY SPECIES OF DIICTODON WERE THERE? Sry

reported to be large or small, always present, always absent, or variably present within the

same alleged species. In the latter case authors have explained this by sexual dimorphism

and/or ontogenetic variation.

The question of sexual dimorphism in dicynodonts, as evidenced by presence or

absence of tusks, is a long-standing one. It was reviewed by Cluver (1971), but it was not

then possible to draw any conclusions. As far as Diictodon is concerned, data given by

Smith (1989) are highly suggestive that sexual dimorphism is present in this genus, one

sex being tusked, the other tuskless. In a collection of skulls from one locality (Dunedin,

Beaufort West), Smith noted that 94 specimens were tusked, 84 tuskless and the condition

of 18 could not be deduced from the material at that stage.

If several Diictodon skulls were to be reported from one locality and horizon all

either with or without tusks, then presence and absence of tusks could be used as a valid

feature of a species. However, this is not true for any of the hitherto recognized species

of Diictodon. It is far more likely that the condition of the tusks is a sexually dimorphic

or ontogenetic feature and, therefore, it should not be used to distinguish different species.

Whether a boss is present behind the pineal foramen (6.1)

Only one species, Diictodon grimbeeki, is reported to have a boss behind the pineal

region. In his description of the species, Broom (1935) mentioned 19 ‘fairly good skulls’,

but does not say how many of them have a post-pineal boss.

The feature occurs in one specimen in the South African Museum sample, SAM-—

K7132. The latter specimen is tusked, approximately 115 mm in skull length (the largest

in the collection), and is from the locality Leeukloof in the Tropidostoma—Endothiodon

assemblage zone of South Africa. The type of Diictodon grimbeeki is from the same

locality. Both tusked and tuskless forms were reported by Broom (1935). The type skull

is 120 mm in length.

Several other Diictodon specimens in the South African Museum sample are known

from this locality, but none has a post-pineal boss. These specimens are otherwise indis-

tinguishable from the specimen with the post-pineal boss. There is evidence that other

kinds of dicynodont skull bosses (nasal, frontal, pineal) are correlated with size or sexual

dimorphism (Toerien 1953; Tollman et al. 1981) and it is most likely that the occasional

occurrence of a post-pineal boss also falls within the category of intraspecific variation of

some kind.

Whether separate nasal bosses are present (6.2); and whether a median boss over the

nostrils is present (6.3)

Separate nasal bosses are present in Diictodon heunei and a single median boss in

various other species. All specimens in the South African Museum sample have nasal

bosses of some description, but it is often difficult to say whether they are separate or

confluent. For example, they may be separated from each other medially, but confluent

posteriorly. Separation also depends to some extent on how well developed the bosses are,

and on distortion of the snout.

Whether the dentary symphysis is beak-like (7.1)

The degree to which the dentary symphysis appears beak-like depends on how well

the front end of the lower jaw is preserved, and is an unreliable character.

318 ANNALS OF THE SOUTH AFRICAN MUSEUM

Whether the front part of lower jaw is broad and deep (7.2)

In order to be applicable to other specimens, this feature needs quantifying: the

breadth and depth in question need to be related to some other aspect of jaw size, such as

length. However, such measurements are prone to distortion and affected by incomplete

preservation. Therefore in many cases they would simply be estimates, so the feature is

likely to be unreliable.

Size of intramandibular foramen (7.3)

This feature is affected by distortion. The fenestra may appear to be closed-up if the

spur of the dentary, which runs dorsal to the fenestra, is pushed downwards over it. The

fenestra may appear to be of very different sizes on the opposite sides of a single jaw (e.g.

SAM-K7738) if distortion has affected them differently.

Presence, absence or reduction of the septomaxilla (8.2)

It has been suggested that the septomaxilla may be absent or reduced in two species,

Diictodon palustris (Broom 1913) and Diictodon testudirostris (Broom & Haughton

1913). Broom (1913) stated that, “There does not appear to be a septomaxillary. If one is

present it is very small and does not show on the face.’ Broom & Haughton (1913) were

similarly unsure, ‘There is no evidence of a septomaxillary, at least on the outer side of

the skull. Whether there may be one hidden underneath it is impossible to say without

damaging the skulls. We incline, however, to think that the septomaxillary is absent.’

The septomaxilla is a small, fairly loosely-articulated bone, which is probably easily

lost from the skull, as noted by Cluver & Hotton (1981) in Diictodon galeops. Its absence

in the fossil does not necessarily mean it was absent in life. Well-preserved and well-

prepared specimens in the South African Museum sample invariably show a septomaxilla,

although it is often broken and never appears on the face.

Depth of the jugal arch (8.3)

Both Diictodon ictidops and Diictodon jouberti are noted as having a very deep

zygomatic arch beneath the postorbital bars. However, this is a feature that may be

produced by deformation of the skull. Two skulls in the South African Museum sample

illustrate this. SAM—K7738 has been dorso-ventrally compressed and the zygoma lies

more horizontally than usual. In side view this gives the impression of a shallow zygoma.

SAM-K7281 has been laterally compressed and the zygoma lies almost vertically and

appears very deep. The type of Diictodon jouberti is damaged in the zygomatic region but

the skull has been compressed laterally and there is a break between zygoma and post-

orbital bar. The type of Diictodon ictidops is described as narrow so there is also the

possibility that it has been laterally compressed.

This is clearly an unreliable feature on which to base specific distinction.

The prefrontal region of orbital margin is elevated (8.4)

Broom (1912) noted this feature in Diictodon psittacops, referring to thickening of

the antero-dorsal margin of the orbit. A swelling in this position is present in many

specimens and is particularly well developed in SAM-—K7643, one of the largest indivi-

duals known. It is probably size related and unhelpful as a specific character.

HOW MANY SPECIES OF DIICTODON WERE THERE? 319

Size and shape of the postfrontal (8.10)

Various sizes and shapes of the postfrontal have been described in type specimens.

The bone was said to be short and wide in Diictodon haughtonianus, absent in Diictodon

nanus and Diictodon galeops and fairly well-developed in Diictodon pseudojouberti, large

in Diictodon rubidgei, and clearly present in Diictodon huenei. The South African

Museum sample also shows that the bone is very variable both in size and shape. It may

be a fairly well-developed triangular bone that reaches the orbital border, or a mere sliver

of bone confined to the skull roof. Conditions in between these extremes can be seen. It

seems unwise to use this character to separate species because of its great variability.

The palatine reaches relatively far forward on the palate (9.1)

Although there is some variability in the sutural pattern in Diictodon, the typical

pattern (as seen in the South African Museum sample specimens) is for the maxilla to

separate palatine and premaxilla (see Fig. 2). The palatine bone does not usually run

anteriorly to contact the premaxilla as it does in Diictodon grossarthi. This seems then to

be a distinctive feature of Diictodon grossarthi. The type is from the locality La-de-da in

the Pristerognathus—Diictodon assemblage zone, as are the types of Diictodon heunei and

Diictodon broomi, which are otherwise very similar but do not exhibit this particular

condition of the premaxilla. SAM—K7643 from La-de-da also shows a sutural pattern

within the normal range of variation for other Diictodon specimens.

In the South African Museum sample, the degree to which the palatine approaches

the premaxilla is variable but nothing like the condition illustrated in Diictodon grossarthi

occurs. Although it is possible that this might represent a real difference between Diicto-

don grossarthi and other specimens, in view of its occurrence in only one known speci-

men, and the variability of the sutural pattern in other specimens, it should not be regarded

as a reliable specific character.

Presence of an oval depression at the junction of the ectopterygoid, jugal and palatine (9.2)

This feature was noted in Diictodon grossarthi by Broili & Schréder (1937). It

occurs in many, but not all, of the specimens of the South African Museum sample,

although its size is variable. It does not seem to be helpful in distinguishing separate

species, because of its wide variability.

Whether a nodule is present on the anterior end of the palatine (9.5)

This feature is noted by Sun (1973) in Diictodon tienshanensis, a tuskless specimen.

It is present also in specimens of the South African Museum sample, both tusked and

tuskless, where preservation of the palatine is good. It is probably a constant feature of

the genus and not of use in delineation of species.

Whether a small boss is present on the inner posterior side of the maxillary process (9.6)

This is another feature noted by Sun (1973) in Diictodon tienshanensis. Most speci-

mens of the South African Museum sample have a small tubercle behind the canine tusk

or flange, although in some cases this is very indistinct. A similar tubercle was noted by

Cluver (1970) in Diictodon testudirostris. Again this feature is probably present through-

out the genus and not of help in delineation of species.

320 ANNALS OF THE SOUTH AFRICAN MUSEUM

SUMMARY

None of the various features used by previous authors seems acceptable as characters

on which to base specific differences. However, with a sample of fossils that are ade-

quately preserved and prepared, it might prove possible to find other features not noted

by previous authors owing to indifferent specimens or incomplete preparation. The South

African Museum sample was studied from this point of view, but no obvious candidates

were found. Although there is variability in the sample in features such as preparietal

suture, premaxilla suture, bosses, and canine tusks, none of these is useful for delimiting

species, as discussed above.

CONCLUSION

Since no reliable specific characters can be found, it is not possible to justify the

20 different species of Diictodon that exist in the literature. There is no reason why all

Diictodon specimens should not be referred to Diictodon galeops.

Diictodon galeops is distinguished by the generic features of Diictodon: dicynodonts

that are tusked or tuskless, have a narrow intertemporal region with the postorbitals

tending to cover the parietals behind the pineal foramen, although this covering is not

always complete. The pineal foramen is often surrounded by a bony ring or boss, espe-

cially in larger specimens. The intertemporal region is approximately the width of the

interorbital region. There are bosses over the nostrils, which are sometimes confluent in

the midline. The caniniform process (whether bearing a tusk or not) is set obliquely to the

ventral maxillary rim, leaving a sharp-edged notch. The premaxilla and palatine are

separated by the maxilla on the palatal surface. The palatine bears a small boss on its

antero-medial corner. The maxilla bears a small tubercle postero-medial to the caniniform

process. The anterior rami of the pterygoid are straight and built up into strong keels

midway along their length. There are two anterior palatal ridges that meet the anterior

premaxillary margin, a single median premaxillary ridge, and low ridges running along

the maxilla—premaxillary suture.

The lower jaw has distinctive dentary tables that are excavated into shallow troughs

and whose medial edge is higher than the lateral. The postero-medial corner of the dentary

table is drawn out into a distinct angle overhanging the edge of the jaw ramus. The lateral

dentary shelf is not prominent.

As far as can be ascertained from specimens available for study, the genus Diictodon

appears to contain only one justifiable species, Diictodon galeops. This species is thus

known from the Tapinocephalus—Bradysaurus assemblage zone through to the Dicyno-

don-Theriognathus zone of the Late Permian Karoo sediments of South Africa. The genus

is also known from Zambia (Gale 1988), China (Sun 1973), and Russia (pers. obs.).

Although the precise correlation with the South African stratigraphic scheme is uncertain

at present, the non-South African specimens of Diictodon are from rocks probably equiva-

lent to Cistecephalus—Aulacephalodon or Dicynodon—Theriognathus assemblage zones,

(King 1992), and they therefore fall within the known range of the genus in South African

rocks.

One qualification about this range should be noted. Specimens of Diictodon from the

Tapinocephalus—Bradysaurus assemblage zone are rare in museum collections, and often

HOW MANY SPECIES OF DIICTODON WERE THERE? |

very poorly preserved. (Several specimens in the South African Museum collections

which were prepared in the hope that they were Diictodon in fact proved to be Robertia

(King & Rubidge in press). It is possible that Robertia may well be the relatively common

dicynodont in the Tapinocephalus—Bradysaurus assemblage zone, rather than Diictodon.)

Therefore, while this study has attempted to use well-preserved and well-prepared speci-

mens, this has been possible only to a lesser degree with Tapinocephalus—Bradysaurus

assemblage zone specimens. However, specimens at my disposal, even when poorly pre-

served, show no difference in the general suite of characters used to define the genus, and

exhibit no characters which might be used to delineate a species other than Diictodon

galeops.

If Diictodon can be considered to span all but the lowest of the Late Permian assem-

blage zones of the Karoo, this would give it a species longevity of between five and ten

million years, since Rubidge (in press) considers the Tapinocephalus—Bradysaurus assem-

blage zone to be between 258 Ma and 253 Ma and the Dicynodon-Theriognathus assem-

blage zone to be Upper Tatarian (up to 248 Ma).

It appears that the one species, Diictodon galeops, is both long-lived and widespread.

Since few specific revisions of mammal-like reptiles have been carried out so far, few

measurements of specific longevity are available and this information on Diictodon will

therefore make a useful contribution to discussions of species longevity and diversity in

the Late Palaeozoic.

ACKNOWLEDGEMENTS

I would like to thank Drs Michael Cluver and Margaret Avery (South African

Museum) for useful discussion and Bruce Rubidge and Roger Smith for providing

stratigraphic information. I am grateful to Clive Booth for photography and to Janet

Goodall and Sheena Kaal for carrying out the extensive preparation of fossils which this

project necessitated.

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176-183.

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HOW MANY SPECIES OF DIJCTODON WERE THERE? 323

ABBREVIATIONS

Morphology

b nasal boss

cp caniniform process

ds dentary shelf

dt dentary table

Idt lateral edge of dentary table

mdt medial edge of dentary table

mf mandibular fenestra

n notch in maxillary rim

pa palatine

pm _ premaxilla

po postorbital

pp preparietal

r ring of bone around pineal foramen

Institutions

AMNH American Museum of Natural History, New York, USA

BMNH Natural History Museum, London, UK

BPI Bernard Price Institute for Palaeontological Research, Johannesburg, RSA

BSP Bayerische Staatssammlung fiir Palaontologie und historische Geologie, Munich, Germany

IVPP Institute for Vertebrate Paleontology and Paleoanthropology, Beijing, China

RC Rubidge Collection, Wellwood, Graaff-Reinet, RSA

SAM South African Museum, Cape Town, RSA

™ Transvaal Museum, Pretoria, RSA

UMZC University Museum of Zoology, Cambridge, UK

UT Universitat Tiibingen Museum und Institut fiir Geologie und Paldontologie, Germany

APPENDIX

Diictodon specimens in the South African Museum used in this study:

SAM-—2354: Skull from Dunedin (Quaggafontein 82, Beaufort West District). Tropidos-

_ toma—Endothiodon assemblage zone. Type of Dicynodon testudirostris.

SAM-—K6654: Skull (with anterior part of snout missing) and lower jaw, from Willowdene

(Beato 238, Beaufort West District). Tropidostoma—Endothiodon assemblage zone.

SAM-—K6929: Skull and lower jaw, and skeleton from Oukloof Pass (Wilgerbosch-

kloof 449, Fraserburg District). Tropidostoma—Endothiodon assemblage zone.

SAM-K6724: Skull and lower jaw, from Amandelboom (Ryers Valley 401, Fraserburg

District). Tropidostoma—Endothiodon assemblage zone.

SAM-K6588: Skull and lower jaw, from Amandelboom (Ryers Valley 401, Fraserburg

District). Tropidostoma—Endothiodon assemblage zone.

SAM-K5189: Skull and lower jaw, from Dunedin (Quaggafontein 82, Beaufort West

District). Tropidostoma—Endothiodon assemblage zone.

SAM-K7084: Skull and lower jaw, from Leeu Kloof 43, Beaufort West District. Tropi-

dostoma—Endothiodon assemblage zone.

SAM-—10394: Anterior part of skull, and lower jaw, from Highlands (Matjesfontein 220,

Victoria West District). Tropidostoma—Endothiodon assemblage zone.

SAM-K5990: Skull and lower jaw, from Dunedin (Quaggafontein 82, Beaufort West

District). Tropidostoma—Endothiodon assemblage zone.

324 ANNALS OF THE SOUTH AFRICAN MUSEUM

SAM-10394: Anterior part of skull, and lower jaw, from Highlands (Matjesfontein 220,

Victoria West District). Tropidostoma—Endothiodon assemblage zone.

SAM-K5990: Skull and lower jaw, from Dunedin (Quaggafontein 82, Beaufort West

District). Tropidostoma—Endothiodon assemblage zone.

SAM-K7281: Skull, from Karreebosch (Karree Bosch 63, Murraysburg District). Tropi-

dostoma—Endothiodon assemblage zone.

SAM-K5105: Skull and lower jaw, from Dunedin (Quaggafontein 82, Beaufort West

District). Tropidostoma—Endothiodon assemblage zone.

SAM-K6017: Skull and lower jaw, for Dunedin (Quaggafontein 82, Beaufort West Dis-

trict). Tropidostoma—Endothiodon assemblage zone.

SAM-—10078: Skull (with right squamosal, and occipital condyle absent) and lower jaw,

from Dunedin (Quaggafontein 82, Beaufort West District). Tropidostoma—Endothiodon

assemblage zone.

SAM-—10377: Skull in two parts, from Noblesfontein (Nobelsfontein 248, Victoria West

District). Cistecephalus—Aulacephalodon assemblage zone.

SAM-—10341: Skull, from Noblesfontein (Nobelsfontein 248, Victoria West District).

Cistecephalus—Aulacephalodon assemblage zone.

SAM-K5204: Skull, from Hoeksplaas (Hoeksplaas 159, Murraysburg District). Ciste-

cephalus—Aulacephalodon assemblage zone.

SAM-K6991: Skull and lower jaw, from Doornplaats (Rust 126, Graaff-Reinet District).

Dicynodon-Theriognathus assemblage zone.

SAM-K7795: Skull and lower jaw, and some post-crania, including complete fore-limb

and hand, from Meltonwold (Melton Wold 158, Victoria West District). Pristerognathus—

Diictodon assemblage zone.

SAM-K6873: Skull and lower jaw, from Waterval (Brandewyns Gat 214, Beaufort West

District). Tropidostoma—Endothiodon assemblage zone.

SAM-10086: Skull and lower jaw, from Dunedin (Quaggafontein 82, Beaufort West

District). Tropidostoma—Endothiodon assemblage zone.

SAM-K6827: Skull and lower jaw, from Willowdene (Beato 238, Beaufort West Dis-

trict). Tropidostoma—Endothiodon assemblage zone.

SAM-K6668: Skull (with right side of snout absent) and lower jaw, from Willowdene

(Beato 238, Beaufort West District). Tropidostoma—Endothiodon assemblage zone.

SAM-K7036: Skull and lower jaw, from Leeuwe Kloof 402, Fraserburg District. Hoede-

maker M., Tropidostoma—Endothiodon assemblage zone.

SAM-K6921: Skull and lower jaw, from Oukloof Pass (Wilgerboschkloof 449, Fraser-

burg District). Tropidostoma—Endothiodon assemblage zone.

SAM-K6009: Skull and lower jaw, from Dunedin (Quaggafontein 82, Beaufort West

District). Tropidostoma—Endothiodon assemblage zone.

SAM-11851: Skull and lower jaw, from Perdewater (Veldmans River 9, Prince Albert

District). Pristerognathus—Diictodon assemblage zone.

SAM-K7603: Skull and lower jaw, from Leeurivier (Rietfontein 306, Beaufort West

District). Pristerognathus—Diictodon assemblage zone.

HOW MANY SPECIES OF DIICTODON WERE THERE? 325

SAM-K7132: Skull and lower jaw, from Leeukloof (Leeu Kloof 43, Beaufort West District).

Tropidostoma—Endothiodon assemblage zone.

SAM-K7673: Skull and lower jaw, from Leeukloof (Leeu Kloof 43, Beaufort West District).

Tropidostoma—Endothiodon assemblage zone.

SAM-K7674: Skull and lower jaw, from Leeukloof (Leeu Kloof 43, Beaufort West District).

Tropidostoma—Endothiodon assemblage zone.

SAM-K7738: Skull and lower jaw, from Leeukloof (Leeu Kloof 43, Beaufort West District).

Tropidostoma—Endothiodon assemblage zone.

SAM-K6979: Skull and lower jaw, from Doornplaats (Rust 126, Graaff-Reinet District).

Dicynodon-Theriognathus assemblage zone.

SAM-K7675: Skull and lower jaw, from Leeukloof (Leeu Kloof 43, Beaufort West District).

Tropidostoma—Endothiodon assemblage zone.

SAM-K7730: Skull and lower jaw, from Leeukloof (Leeu Kloof 43, Beaufort West District).

Tropidostoma—Endothiodon assemblage zone.

SAM-K7643: Skull and lower jaw, from La-de-da (La-de-da 178, Beaufort West District).

Pristerognathus—Diictodon assemblage zone.

SAM-11563: Skull and lower jaw, from Kroonplaas (Honing Kaps Fontein 321, Beaufort

West District). Tapinocephalus—Bradysaurus assemblage zone.

SAM-11589: Skull, from Koedoeskop (Leeuwkraal 309, Beaufort West District). Tapino-

cephalus—Bradysaurus assemblage zone.

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6. SYSTEMATIC papers must conform to the International 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. Du Toit 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

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‘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.

GILIAN M. KING

HOW MANY SPECIES OF DITICTODON

WERE THERE?

y VOLUME 102 PART 10 FEBRUARY 1993 ISSN 0303-2515

OF THE SOUTH AFRICAN

he MUSEUM

CAPE TOWN

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paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable):

(a) Author’s name and year of publication given in text, e.g.:

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

FIscHER, P. H. 1948. Données sur la résistance et de la vitalité des mollusques. Journal de conchyliologie 88 (3): 100-140.

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

expérimentale et générale 74 (33): 627-634.

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

Magazine of Natural History (13) 2 (17): 309-320.

Koun, A. J. 1960b. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. Bulletin of

the Bingham Oceanographic Collection, Yale University 17 (4): 1-51.

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

pologische Ergebnisse einer Forschungsreise im westlichen und zentralen Stid-Afrika ausgefiihrt in den Jahren

1903-1905 4 (15). Denkschriften der medizinisch-naturwissenschaftlichen Gesellschaft zu Jena 16: 269-270.

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 102 °#Band

February 1993 Februarie

Part 10 Deel

ANNOTATED CATALOGUE OF THE AVES TYPE

SPECIMENS IN THE SOUTH AFRICAN MUSEUM

R. K. BROOKE

Cape Town Kaapstad

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D1842

ANNOTATED CATALOGUE OF THE AVES

TYPE SPECIMENS IN THE SOUTH AFRICAN MUSEUM

R. K. Brooke

Percy FitzPatrick Institute of African Ornithology, University of Cape Town,

and South African Museum

[MS accepted 12 November 1991]

ABSTRACT

There is type material of 50 avian nominal taxa (of which 12 are fossil) in the South African

Museum, as well as those of two further names that were not published. These are listed with comments,

including nomenclatural, as appropriate. Particular attention is paid to where the specimens were obtained.

It is suggested that the type locality of Pterocles gutturalis Smith should be clarified by selection of a

lectotype. The type locality of Agapornis nigrigenis Sclater is established for the first time. The type

locality of Parisoma layardi Hartlaub is corrected. Two synonyms in Pachyptila Illiger are clarified.

Barbatula extoni Layard is shown to be based on two syntypes. The date of publication of Winterbottom’s

proposed names in Chersomanes albofasciata is established. Smith’s proposal of a timaliid jardineii is

clarified. It is argued that Laniarius maraisi Sclater is a colour phase of Telophorus olivaceus (Shaw) and

not just the juvenile plumage.

CONTENTS

PAGE

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ISOMEY NC CSPECLINCNS 1.82. oils wees ine Cd Cae we a be Bee 328

PCM ONE CER CIIICIBES ula ec Ss hen eles) oe a leu es aes ween e AN eee, wena 345

SGUSIETTOSS. EPS Wala SOULE RCICPOE Ree or St EP eter UM es ton av a 345

INTRODUCTION

The South African Museum has already published lists of types in the following

taxa: decapod Crustacea (Kensley 1974), Hydroida (Coelenterata) (Millard 1979), Dino-

cephalia (Therapsida) (Van den Heever & Grine 1981), and marine Mollusca other than

Cephalopoda (Giles & Gosliner 1983); Carabidae (Coleoptera) (Cochrane in press) is to

be published shortly. The avian nominal taxa are listed below, following the sequence

proposed by Wetmore (1960). It is accepted that Wetmore’s system will soon be replaced,

perhaps by that of Sibley & Ahlquist (1990) and Sibley & Monroe (1990), or perhaps by

another that has yet to be published. In the meantime, clarity is best served by using

Wetmore’s system.

Under ordinal and family names, type specimens are listed first under the name

proposed, with bibliographic reference. In the case of fossil species, this is preceded by

a ‘+’. Details of type specimens are then given, with particular reference to where each

specimen was obtained. Dr J. M. Winterbottom did not usually put his name as collector

on specimens labelled in his handwriting. In some cases, the describer of a taxon has

stated that J. M. W. was the collector and it has been assumed that he was the collector

of specimens labelled by him and collected in the 1950s and 1960s. Under ‘Remarks’,

Ann S. Afr. Mus. 102 (10), 1993: 327-349.

328 ANNALS OF THE SOUTH AFRICAN MUSEUM

comments are made on the state of specimens, historical and nomenclatural points, and

any other pertinent matters, including the meaning of the epithet chosen by the proposer.

The current name is then given, usually following Clancey (1980).

In searching for type specimens, I have been guided by the names of ornithologists

who have worked at the South African Museum: in historical order, Edgar Leopold

Layard, Roland Trimen (better known as an entomologist), William Lutley Sclater, Edwin

Leonard Gill, and John Miall Winterbottom (Summers 1975). However, Dr Gill did not

describe any birds as new. If I could not find the type of a taxon proposed by any of the

remaining four workers, I ascertained that the type was said to be in another museum and

searched no further. In addition, Dr Winterbottom used to annotate in red the card

index to the bird skin collection in respect of type specimens and this has been of great

assistance.

Appreciation of ornithologists at the South African Museum by their contemporaries

and successors has never been brought together and is, in any case, very scattered through

the literature. However, it seems desirable to mention the principal appreciations: Layard

(Liversidge 1957); Trimen (Sclater 1916); Sclater (Grant 1945); Gill (Broekhuysen 1956;

Winterbottom 1957a); and Winterbottom (Clancey 1984; Brooke & Siegfried 1985).

LIST OF TYPE SPECIMENS

Order SPHENISCIFORMES Sharpe

Family Spheniscidae Bonaparte

+ Spheniscus predemersus Simpson, 1971: 1145.

Holotype SAM—PQL12887A, left humerus from the late Pliocene, at E Quarry,

Langebaanweg, 32°58’S 18°09’E, south-western Cape Province, South Africa.

Remarks

In fact, the type comes from the Quartzose Sand Member, Varswater Formation,

early Pliocene (Hendey 1981). The epithet indicates that the taxon was the predecessor

of S. demersus, at least in time, if not phylogenetically. Subsequently, Simpson (1975)

proposed a new monotypic genus /nguza for this taxon. Now known as Spheniscus pre-

demersus Simpson (Clancey et al. 1987), following the preliminary advice of Dr S. L.

Olson (in litt. 1985) on the spheniscid taxa proposed by the late Dr George Gaylord

Simpson.

+ ?Palaeospheniscus huxleyorum Simpson, 1973: 343.

Holotype SAM-PQ1882, right humerus with both ends incomplete from the Mio-

cene, at Ysterplaats airforce base, 33°55’S 18°29’E, Cape Town, Cape Province, South

Africa.

Remarks

The deposits at Ysterplaats are now thought to date from the early Pliocene (Olson

1983, 1985b). Named in honour of Thomas Huxley and his grandson, Sir Julian Huxley,

for their contributions to vertebrate systematics and evolutionary theory. Now known as

Spheniscus huxleyorum (Simpson) (Clancey et al. 1987).

AVES TYPE SPECIMENS IN THE SOUTH AFRICAN MUSEUM 329

+ Dege hendeyi Simpson, 1979a: 6.

Holotype SAM—PQL28455, left tarsometatarsus with most of metatarsals II and III

(metatarsal IV found and attached later) and proximal articulation from the Quartzose

Sand Member, Varswater Formation, early Pliocene, at E Quarry, Langebaanweg,

32°58’S 18°09’E, south-western Cape Province, South Africa.

Remarks

Named in honour of Dr Quentin Brett Hendey, then Curator of Tertiary Palaeon-

tology at the South African Museum, who devoted many years to the collection of material

at Langebaanweg and to the study of the mammals represented therein. He is currently

Director of the Durban Natural Science Museum. The palaeoecology of the Langebaanweg

fossil deposits is set out in Hendey (1981). Now known as Spheniscus hendeyi (Simpson)

(Clancey et al. 1987).

+ Nucleornis insolitus Simpson, 1979b: 4.

Holotype SAM—MBD4, right tarsometatarsal lacking the distal ends of the second

and third metatarsals, from ?Miocene deposits, at Koeberg Nuclear Power Station,

Duinefontein, 33°39’S 18°27’E, south-western Cape Province, South Africa.

Remarks

The deposits at Duinefontein are now thought to date from the early Pliocene (Olson

1983, 1985b). The epithet means unusual and is a reference to the presence of a single

medial intermetatarsal foramen. Now known as Spheniscus insolitus (Simpson) (Clancey

et al. 1987).

Order PROCELLARIIFORMES Furbringer

Family Procellariidae Leach

+ Pachyptila salax Olson, 1985a: 130.

Holotype SAM—PQL25 187, left humerus, complete, from the Quartzose Sand Mem-

ber, Varswater Formation, early Pliocene, at E Quarry, Langebaanweg, 32°58’S 18°09’E,

south-western Cape Province, South Africa.

Remarks

The epithet means a sieve and refers to the filtering apparatus in the upper mandibles

of living members of the genus. Now known as Pachyptila salax Olson (Clancey et al.

1987).

Heteroprion desolata peringueyi Mathews, 1912: 223.

Holotype SAM-6792, adult female from Port St Johns, 31°38’S 29°32’E, Transkei,

collected by Mr Guy C. Shortridge from the beach on 4 September 1902.

Remarks

In fair to good condition. Named in honour of Dr Louis Albert Péringuey, then

Director of the South African Museum and a noted coleopterist. Captain Guy C.

Shortridge (as he later became, and still later Director of the Kaffrarian Museum, King

William’s Town) is better known for his work on southern African mammals, particularly

330 ANNALS OF THE SOUTH AFRICAN MUSEUM

those of Namibia. Curiously, the type specimen is not mentioned in Shortridge & Sclater

(1904), although its date of collection falls within their purview. Jouanin & Mougin

(1979) place H. peringueyi as a synonym of Pachyptila desolata banksi Smith but this

taxon is not admitted by either Cox (1980) or Bretagnolle et al. (1990) who all

synonymize it with the nominate race. The type specimen measures in mm: length of

exposed culmen 28,0, maximum width of culmen 12,4, wing > 175, tarsus 30,6. It is in

active moult of the primaries (primary score 30) and the unguis at the tip of the upper

mandible is pale, the rest being black. I place H. peringueyi as a junior synonym of

P. desolata desolata (Gmelin). The taxonomy of the genus Pachyptila Wliger is still

strongly disputed. I have followed Bretagnolle et al. (1990) without believing that they

have said the last word.

Prion vittata keyteli Mathews, 1912: 212.

Holotype SAM-—10316, unsexed adult from Tristan da Cunha, South Atlantic,

collected by Mr P. C. Keytel on an unstated date.

Remarks

In poor condition. Named in honour of Mr P. C. Keytel, the collector, who visited

Tristan da Cunha and Inaccessible Islands in 1908 and 1909 (Phillips 1913; Winterbottom

1976) for somewhat less than one year. Mathews (1932) added that the type was collected

with its egg on Inaccessible Island, 37°19’S 12°44’W, on 17 September 1908. The type

specimen measures in mm: length of exposed culmen 32,1, maximum width of culmen

21,4, wing length 200, tarsus 36,2. There is no moult of the primaries and the bill is

wholly black. Jouanin & Mougin (1979) place Prion keyteli as a synonym of Pachyptila

vittata vittata (Forster). The status of Prion keyteli was studied by Clancey (1981) who

also found that P. keyteli is a junior synonym of Pachyptila v. vittata (Forster).

Family Hydrobatidae Mathews

+ Oceanites zaloscarthmus Olson, 1985a: 126.

Holotype SAM—PQL25214, right humerus, complete, from the Quartzose Sand

Member, Varswater Formation, early Pliocene, at E Quarry, Langebaanweg, south-

western Cape Province, South Africa.

Remarks

The epithet means skipping over the surging sea, a common foraging behaviour

pattern in the living members of the family. I have used the family-group name Hydro-

batidae Mathews (1912), which he created as a replacement name for Thalassidromidae

Miiller—Thalassidroma Vigors being a junior synonym of Hydrobates Boie—instead of

Oceanitidae Forbes as used by Clancey (1980) and other authors listed by Olson (1987).

In this I follow Prof. Walter J. Bock MS of an application to the International Commis-

sion on Zoological Nomenclature seeking a general ruling conserving all family-group

names currently widely used in the Aves. This will probably be accepted in due course,

despite the trenchant objections of Olson (1987), and therefore I use the name in the

interests of long-term clarity. Now known as Oceanites zaloscarthmus Olson (Clancey et

GEMS):

AVES TYPE SPECIMENS IN THE SOUTH AFRICAN MUSEUM Sie

Family Pelecanoididae Gray

+ Pelecanoides cymatotrypetes Olson, 1985a: 139.

Holotype SAM—PQL 14564, left humerus, complete, from the Quartzose Sand Mem-

ber, Varswater Formation, early Pliocene, at E Quarry, Langebaanweg, south-western

Cape Province, South Africa.

Remarks

The epithet means a wave-borer, from the habit of living members of the family of

flying straight through the crests of waves. The family Pelecanoididae is not known from

South African waters, even as vagrants, in the historical period (Clancey 1980). However,

this taxon was a breeding species on offshore islands near Langebaanweg (Olson 1985a).

Now known as Pelecanoides cymatotrypetes Olson (Clancey et al. 1987).

Order CICONIIFORMES Bonaparte

Family Ciconiidae Sundevall

+ Ciconia kahli Haarhoff, 1988: 300.

Holotype SAM—PQL22164, partial associated skeleton from the Quartzose Sand

Member, Varswater Formation, early Pliocene, at E Quarry, Langebaanweg, south-

western Cape Province, South Africa.

Remarks

Named in honour of Dr Marvin Philip Kahl who has written extensively on the

systematics and behaviour of ciconiiform birds (e.g. Kahl 1979).

Family Scopidae Bonaparte

+ Scopus xenopus Olson, 1984b: 737.

Holotype SAM—PQL43396, distal end of left tarsometatarsus with posterior parts of

inner and outer trochleae abraded from the Quartzose Sand Member, Varswater Forma-

tion, early Pliocene, at E Quarry, Langebaanweg, south-western Cape Province, South

Africa.

Remarks

The epithet means strange foot and is a reference to the unusual morphology of the

tarsometatarsus. Xenopus Wagler is also the generic name of some toads which the living

member of the family captures and eats. Now known as Scopus xenopus Olson (Clancey

et al. 1987).

Family Threskiornithidae Poche

+ Geronticus apelex Olson, 1985c: 59.

Holotype SAM-—PQL20692, partial associated skeleton from the Quartzose Sand

Member, Varswater Formation, early Pliocene, at E Quarry, Langebaanweg, south-

western Cape Province, South Africa.

552 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

The epithet means lacking a helmet and is an allusion to the lack of an expanded

bony occipital crest found in the two living species of this genus. The family name used

is that recommended by Prof. Walter J. Bock—see under Oceanites zaloscarthmus above.

Now known as Geronticus apelex Olson (Clancey et al. 1987).

Order GALLIFORMES Temminck

Family Phasianidae Horsfield

Scleroptila jugularis cunenensis Roberts, 1932: 22.

Holotype SAM-—16508a, adult male from the Cunene River, Namibia, collected by

Mr Archibald Gerald White, on 14 March 1923.

Remarks

In good condition. The epithet refers to the type locality, subsequently elucidated as

Otjimbombe (Macdonald 1953). The site may now be covered by the waters of the Rua

Cana Dam (see map in Tarboton (1967) and discussion of Lamprotornis mevesii violacior

below). The label contains an MS note in Austin Roberts’s handwriting that he intended

to name this subspecies S. j. gilli in honour of Dr E. L. Gill but he did not do so,

preferring to convey that honour by way of Pternistis swainsoni gilli (see below). The

collector, Mr White, was then the junior taxidermist at the South African Museum. He

soon moved to the post of taxidermist at the Transvaal Museum, Pretoria. Now known as

Francolinus levaillantoides jugularis Biittikofer (Clancey 1980).

Pternistis afer cunenensis Roberts, 1932: 22.

Holotype SAM-16511c, adult male from the Cunene River, Namibia, collected

by Mr A. G. White on 15 March 1923.

Remarks

In very good condition. The epithet refers to the type locality. For notice of the type

locality and collector see the preceding taxon. Now known as Francolinus afer afer

Statius Miller (Clancey 1980).

Since S. j. cunenensis and P. a. cunenensis are both now placed in Francolinus, a

minor problem in homonymy arises. Both names were proposed by Roberts (1932) on

p. 22 and Scleroptila jugularis cunenensis has line priority. The matter is of little conse-

quence unless the proposed subspecies are later thought worthy of recognition (unlikely).

White (1958) proposed Francolinus afer palliditectus as a replacement name for Pternistis

afer cunenensis Roberts.

Pternistis swainsoni gilli Roberts, 1932: 23.

Holotype SAM-—16532, adult male from Ondonga (more correctly Ondangwa),

17°58’S 16°01’E, collected by Mr A. G. White on 22 February 1923.

Remarks

In very good condition. Named in honour of Dr Edwin Leonard Gill, then Director

of the South African Museum. Now known as Francolinus swainsonii gilli (Roberts)

(Clancey 1980).

AVES TYPE SPECIMENS IN THE SOUTH AFRICAN MUSEUM 333

Order CHARADRIIFORMES Huxley

Family Rostratulidae Mathews

+ Rostratula minator Olson & Eller, 1989: 118.

Holotype SAM—PQL25552, nearly complete left humerus from the Quartzose Sand

Member, Varswater Formation, early Pliocene, at E Quarry, Langebaanweg, 32°58’S

18°09’E, south-western Cape Province, South Africa.

Remarks

Type not seen: on loan to Dr Storrs L. Olson in September 1991. The epithet means

one who threatens and is an allusion to the distinctive threat display of the living old

world member of this family.

Order PTEROCLEIFORMES Boucard

Family Pterocleidae Bonaparte

Pterocles burchelli delabati Winterbottom, 1964: 30.

Holotype SAM-—54727a, adult male from Onguma, 18°44’S 17°03’E, Namutoni,

Etosha National Park, Namibia, collected by Mr P. J. Buys on 27 August 1963.

Remarks

In fair condition though the head is in poor condition. Named in honour of Mr Bernabé

de la Bat, Chief Game Warden, Game Department of South West Africa (now Namibia).

For Mr Buys see Campethera bennettii buysi below. Now known as Pterocles burchelli

burchelli Sclater (Clancey 1980).

The spelling of the family and ordinal names based on the genus Pterocles is vari-

able, depending on estimates of the stem of Greek nouns ending in KLES: some have

~ stems KL-, some KLE- and some KLED-. Prof. Walter J. Bock in MS recommends

Pteroclidae and, by implication, Pterocliformes. However, after discussion with Dr M. R.

Mezzabotta of the Classics Department of the University of Cape Town, it appears that

the correct view is that the stem of Pterocles is KLE-, giving Pterocleidae and Pteroclei-

formes, as used here, cf. KLEOPATRA (= Cleopatra).

Pterocles gutturalis Smith, 1836: 56.

Syntype SAM—20890, adult female, labelled Latakoo [= Kuruman, 27°28’S 23°26'E]

in which case it would have been collected in 1835 (Kirby 1965).

Remarks

Formerly mounted; in good condition. Presented to the Museum in 1855 by the

South African Literary and Scientific Institution. Mr W. L. Sclater in MS on the label

suggested that it is one of Andrew Smith’s syntypes, apparently correctly. The type locality

is Kurrichane, 25°21’S 26°11’E, near Zeerust in the western Transvaal, following Smith

(1836). The British Museum (Natural History) has a male syntype from 80 miles east of

Latakoo as well as a female syntype (Warren 1966). It appears from the map in Kirby

(1965) that Smith was never due east of Latakoo but that his route took him north-east of

there. This means that the place of collection of the male syntype is just west of

Vryburg, 26°57’S 24°44’E. Since Smith (1836) only described the male, the male syntype

334 ANNALS OF THE SOUTH AFRICAN MUSEUM

could be declared the lectotype, requiring an adjustment of the type locality. This might

be desirable since Smith (1836) gave the vague ‘Inhabits the country north and south of

Kurrichaine’. The epithet refers to the more distinctive throat pattern found in this species

compared with other southern African sandgrouse. Now known as Pterocles gutturalis

gutturalis Smith (Clancey 1980).

Order PSITTACIFORMES Wagler

Family Psittacidae Rafinesque

Agapornis nigrigenis W. L. Sclater, 1906: 61.

Holotype SAM-—8485, ?adult male from the Muguazi River some 25 km above its

confluence with the Zambezi River, Southern Province, Zambia, collected by Dr A. H. B.

Kirkman in September 1904.

Remarks

A poor specimen not properly stuffed or sewn up afterwards. The Muguazi River is

not a known name (Benson ef al. 1971) but I think it is a misspelling of the name of the

Ngwezi River shown on map A of Pitman (1934) and map 210 of the 1984 Reader’s

‘ Digest Atlas of Southern Africa and which joins the Zambezi at 17°40’S 25°06’E, about

where the Muguazi would (Benson et al. 1971). The collector, Dr Kirkman, made two

game-hunting trips into southern Zambia (Sclater 1906) and subsequently practised in

Queenstown, eastern Cape Province (deduced from specimen material in the South

African Museum). The epithet refers to the most distinctive feature (blackish cheeks and

throat but not black on the crown as in A. personata) distinguishing this species from

other members of the genus. Agapornis nigrigenis is treated as a full species by Clancey

(1980) but it seems to be an isolated melanistic population of A. lilianae Shelley and

should be referred to as A. I. nigrigenis as White (1965) and Benson et al. (1971) have done.

Order COLIIFORMES Murie

Family Coliidae Sundevall

+ Colius hendeyi Rich & Haarhoff, 1985: 23.

Holotype SAM—PQL28858, left tarsometatarsus from the Quartzose Sand Member,

Varswater Formation, early Pliocene, at E Quarry, Langebaanweg, 32°58’S 18°09’E,

south-western Cape Province, South Africa.

Remarks

Named in honour of Dr Q. B. Hendey: see Dege hendeyi above. Now known as

Colius hendeyi Rich & Haarhoff (Clancey et al. 1987).

Order CORACIIFORMES Forbes

Family Coraciidae Rafinesque

Coracias spatulata Trimen, 1880a: 31.

Holotype SAM-—20898, adult male from Leshumo Valley, 17°50’S 25°15’E, near the

Victoria Falls, Zimbabwe, collected by Dr Benjamin Frederick Bradshaw on 23 May 1878.

AVES TYPE SPECIMENS IN THE SOUTH AFRICAN MUSEUM aa5

Remarks

Formerly mounted; in very good condition. Dr Bradshaw was a police surgeon when

he died in 1883 (Sharpe 1884; Oates 1889). Dr Bradshaw’s bird collections were never

written up and the specimens were dispersed to various public and private collections,

often by sale (Tabler 1966). The epithet describes the unusual shape of the outermost

rectrix. The genitive of the Greek noun Coracias is KORAKIADIS and in the Latin

alphabet the stem would be CORACIAD.-, giving Coraciadidae and Coraciadiformes.

However, the International Commission on Zoological Nomenclature has placed the fam-

ily name Coraciidae on the Official List of Family-Group Names in Zoology (Direction

58 of 20 December 1956) and it would be unbearably pedantic then to spell the ordinal

name as Coraciadiformes. Now known as Coracias spatulata spatulata Trimen (Clancey

1980).

Order PICIFORMES Meyer & Wolf

Family Capitonidae Bonaparte

Barbatula extoni Layard, 1871: 226.

Syntype SAM-19684, adult female from Kanye, 24°59’S 25°21’E, Botswana, pre-

sumably collected by Dr Hugh Exton on an unknown date, presumably in 1869 (Roberts

i935);

Remarks

Formerly mounted (still mounted in 1934 (Roberts 1935)); in good condition. Named

in honour of Dr Hugh Exton, a medical practitioner, whose ornithological work was

noticed by Roberts (1935), including a short biography. The status of this specimen is

uncertain though it is marked as the type by Mr W. L. Sclater. Roberts (1935) regarded

the type as being held in the British Museum (Natural History), then in London, now at

Tring. This is supported by Warren (1966) who stated that that museum holds the male

holotype. However, Layard (1871) did not nominate a type nor indicate which sex it

might be in this sexually monomorphic species. I believe that the species is based on two

syntypes, the male in the B. M. (N. H.) and the female in the South African Museum.

Exton in Roberts (1935) added that he obtained male specimens at Kanye in September

and October 1869. Now known as Pogoniulus chrysoconus extoni (Layard) (Clancey

1980).

Family Picidae Leach

Dendrobates striatus Layard, 1871: 227.

Syntypes SAM—21162a and SAM-21162b, adult male and female (the latter by

plumage), respectively, the male from Kanye, 24°59’S 25°21’E, Botswana, and the female

from Bechuanaland (presumably near Kanye), both collected by Dr Hugh Exton.

Remarks

Both formerly mounted; in fair to good condition. The epithet refers to the heavily

streaked underparts. For Dr Exton see the preceding entry. Dendrobates striatus was

proposed provisionally and does not appear to have been used since, though it is listed as

336 ANNALS OF THE SOUTH AFRICAN MUSEUM

a synonym by Reichenow (1902). Now known as Campethera abingoni abingoni Smith

(Clancey 1980).

Campethera bennettii buysi Winterbottom, 1966: 39.

Holotype SAM-—55390, adult male from Swartboois Drift, 17°20’S 13°52’E, Cunene

River, Namibia, collected by Mr Pieter J. Buys of the State Museum, Windhoek, on

21 October 1965.

Remarks

In good condition. Named in honour of the collector, a technical officer of the State

Museum of Namibia, Windhoek. Now known as Campethera bennettii buysi Winter-

bottom (Clancey 1980).

Order PASSERIFORMES Linnaeus

Family Alaudidae Vigors

Mirafra apiata marjoriae Winterbottom, 1956: 156.

Holotype SAM-19594, adult male from Soetendalsvlei, 34°43’S 19°59’E, near

Bredasdorp, western Cape Province, South Africa, collected by Dr Austin Roberts on

27 October 1940.

Remarks

In good condition. Named in honour of his wife, Marjorie G. Winterbottom. Now

known as Mirafra apiata marjoriae Winterbottom (Clancey 1980).

Certhilauda albofasciata macdonaldi Winterbottom, 1958a: 59.

Holotype SAM-—20340, adult male from 36 km north-east of Karoopoort, 33°15’S

19°43’E, western Cape Province, South Africa, collected by Dr J. M. Winterbottom on

23 May 1956.

Remarks

In fair to good condition. Named in honour of Mr James David Macdonald, then

head of the Bird Department at the B. M. (N. H.), London, who led an expedition to

collect and study the birds of western southern Africa (Macdonald 1957). The date of

publication of the name is May 1958. Some copies were issued with the date October

1957 and some with an overprinted sticker giving the date as May 1958. The latter is

correct: the South African Museum Library’s copy was receipted on 30 May 1958 and a

copy in the South African Library was receipted on 9 June. The date of publication also

affects the status of the name C. a. bathoeni Winterbottom, proposed on p. 64 of the same

issue, and attributed by Winterbottom to R. H. N. Smithers and M. L. Paterson in press.

The name was subsequently reproposed by Paterson alone (1958: 125) issued in October

1958, not January as stated by Clancey (1980). The type locality given by Paterson (1958)

is c. 90 km east of Kakia, 24°16’S 23°24’E, Botswana. Now known as Chersomanes

albofasciata macdonaldi (Winterbottom) and Chersomanes albofasciata bathoeni (Winter-

bottom) (Clancey 1980).

AVES TYPE SPECIMENS IN THE SOUTH AFRICAN MUSEUM 337

Anacorys africanoides omaruru Roberts, 1936: 263.

Holotype SAM—21005, adult male from Omaruru, 21°26’S 15°56’E, Namibia, appar-

ently collected by Mr A. W. Eriksson on 5 December 1878.

Remarks

In good condition. The earliest writing on the label is in Mr A. W. Eriksson’s handwrit-

ing: see Cinnyris erikssoni below, p. 343. The label is endorsed in Dr Austin Roberts’s

handwriting that he intended to name the taxon ‘namibensis’. The epithet is, in fact, the name

of the type locality. Now known as Mirafra africanoides omaruru (Roberts) (Clancey 1980).

Calandrella sclateri capensis Ogilvie-Grant, 1913: 41.

Syntypes SAM-—11635a and SAM-11635b, adult male and female respectively from

Philipstown, 30°26’S 24°28’E, Cape Province, collected by Mr H. Leighton Hare on

unknown dates.

Remarks

In fair to good condition, though the abdomens are not sewn up. Named for the Cape

Province as opposed to Namibia, the type locality of the species. Now known as Spizo-

corys sclateri capensis (Ogilvie-Grant) (Clancey et al. 1987).

Calandrella hamiltoni W. L. Sclater unpublished.

Syntypes SAM-—5873 and SAM-—5874, adult male and female respectively from

Christiania (correctly Christiana, 27°55’S 25°10’E) District, south-western Transvaal,

collected by Captain B. Hamilton on 29 and 30 July 1903.

Remarks

The male specimen is in fair to good condition and the female specimen in good

condition. Mr Sclater intended to name a new species after the collector, a British Army

officer, but realized, before publication, that the species had been described by Shelley

(1902) as Calandrella starki on material from Namibia. However, the specimens are

labelled in red with his intention to describe them. Now known as Eremalauda starki

gregaria (Clancey) (Clancey et al. 1991).

Family Remizidae Olphe-Galliard

Anthoscopus minutus gigi Winterbottom, 1959b: 152.

Holotype SAM-—20323a, adult male from Oudtshoorn, 33°35’S 22°12E, Little

Karoo, Cape Province, South Africa, collected by Dr J. M. Winterbottom on 10 April

1956.

Remarks

In poor condition. Named in honour of Mr Gerald Graham (known as G-G) Smith,

Chairman of the Board of the East London Museum, a museum that had built up a large

collection of Cape Province specimens under his guidance. Now known as Anthoscopus

minutus gigi Winterbottom (Clancey 1980).

338 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Timaliidae Vigors & Horsfield

Cratopus jardineii Smith, 1836: 45.

Syntypes SAM—58324 and SAM-—58325, without locality or sex.

Remarks

Formerly mounted; in good condition. The labels are endorsed by Mr W. L. Sclater

as syntypes, probably correctly. There are no syntypes in the British Museum (Natural

History) (Warren & Harrison 1971) where the majority of Dr Andrew Smith’s type

material eventually ended up. However, there was formerly a syntype in the Royal Scot-

tish Museum, Edinburgh (Stenhouse 1930). The generic name Cratopus used by Smith in

the Willughby Society reprint of his (1836) report, as well as in the original, appears to

be a lapsus for Crateropus Swainson. Named in honour of Sir William Jardine Bt, a

distinguished Scottish naturalist who wrote on birds and arranged for the publication of

books on birds and other major taxa. The type locality is ‘banks of rivers beyond

Kurrichaine’ (Smith 1836), i.e. near Zeerust, 25°32’S 26°05’E, in the western Transvaal.

Now known as Turdoides jardineii jardineii (Smith) (Clancey 1980).

Chaetops aurantius Layard, 1867: 126.

Syntypes SAM—15071a and SAM-—15071b, adult male and female respectively from

near Graaff-Reinet, 32°15’S 24°33’E, eastern Cape Province, South Africa, collected

by Mr A. O’Reilly on unknown dates.

Remarks

Formerly mounted; in good condition. The collector is named as A. O’Reilly on the

label but as Mr J. J. O’Reilly by Layard (1867) who added that Mr O’Reilly was a

magistrate stationed at various towns in the eastern Cape Province. The epithet refers to

the orange, not deep red colour of the abdomen. Described by Layard in the Turdidae but

now believed to be a member of the Timaliidae (Olson 1984a; Clancey et al. 1987).

Irrespective of its family allocation, the taxon is still known as Chaetops aurantius Layard

(Clancey 1980), though some consider that it is a subspecies of C. frenatus (Temminck).

Family Pycnonotidae Gray

Xenocichla debilis W. L. Sclater, 1899b: 284.

Holotype SAM-—3116, adult male from north of Inhambane, Inhambane Province,

Mozambique, collected by Mr H. F. Francis on 10 October 1898.

Remarks

In fair to good condition. The epithet refers to the slender build of the species

compared with its close relative Phyllastrephus flavostriatus Sharpe. For the Francis

brothers, see Erythrocercus francisi below. Sclater (1899b) gave the type locality as

‘about twenty miles north of the Bay of Inhambane, near the coast’. This would place it

close to Massinga, 23°20’S 35°22’E, as noted by Clancey (1971, 1980). Now known as

Phyllastrephus debilis debilis (Sclater) (Clancey 1980).

AVES TYPE SPECIMENS IN THE SOUTH AFRICAN MUSEUM 339

Family Turdidae Rafinesque

Saxicola spectabilis Hartlaub, 1865a: 428.

Syntypes SAM-—15115a and SAM-—15115b, adult male and female respectively from

Windvogelberg, 32°18’S 27°04’E, Cathcart, eastern Cape Province, South Africa, col-

lected by Captain George Ernest Bulger on unknown dates.

Remarks

Formerly mounted; the male is in good condition and the female in fair condition.

The epithet refers to the strikingly patterned plumage, unlike that of so many species from

South Africa that Dr K. J. Gustav Hartlaub of the Bremen Museum had to deal with.

Captain Bulger, whose botanical and military careers are briefly summarized in Gunn &

Codd (1981), was clearly interested in chats, since he published on the behaviour of

Myrmecocichla formicivora Vieillot (Bulger 1869a) and noted that S. spectabilis had been

synonymized with S. bifasciata Temminck (Bulger 1869b). Layard (1865) provided data

on the breeding and behaviour of the ‘new’ species. Now known as Oenanthe bifasciata

(Temminck) (Clancey 1980). Subsequently, Tye (1989) argued that it was better placed

in Saxicola Bechstein, and Clancey (1990) recommended resuscitating the genus Campi-

coloides Roberts of which it is the genotype, in the combination Campicoloides bifas-

ciatus (Temminck).

Saxicola pollux Hartlaub, 1865b: 747.

Holotype SAM—15134, unsexed bird from the Traka Karoo collected by Mr W.

Atmore on an unknown date.

Remarks

Formerly mounted; in good condition but somewhat foxed from exposure in the

public galleries. The epithets castor (see the next taxon) and pollux were the names of

the heavenly twins, whose sister was Helen of Troy, and refer to the similarity of the two

types. The Traka River is a right bank affluent of the Olifants River, which itself runs into

the Gourits River. Most of the course of the Traka River lies in the Great Karoo. Curi-

ously, neither this nor the next taxon is included in Layard’s (1867) book. Now known

as Cercomela schlegelii pollux (Hartlaub) (Clancey 1980).

Saxicola castor Hartlaub, 1865b: 747.

Syntypes SAM—15125a and SAM-15125b, unsexed birds (males by plumage) from

the Karoo collected by Mr W. Atmore on unknown date/s.

Remarks

Formerly mounted; in good condition. Both syntypes belong to the morph with least

white in the plumage. The epithet is explained above in discussing S. pollux. Now known

as Oenanthe monticola monticola Vieillot (Clancey 1980).

Myrmecocichla stoehri W. L. Sclater, unpublished.

Holotype SAM-9272, adult female (male by plumage: the opinion also of Mr Sclater

in Stoehr & Sclater 1906) from Lavushi, 12°23’S 30°52’E, Mpika District, Northern

Province, Zambia, collected by Dr Frederick Otto Stéhr in early September 1905.

340 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

In fair condition. Named in honour of the collector who was primarily a botanist

(Brooke 1970; Crook 1970) but was medical officer and naturalist to a geodetic survey

at the time (Benson et al. 1970). Now known as Myrmecocichla nigra Vieillot (Benson

et al. 1971).

Family Sylviidae Leach

Parisoma layardi Hartlaub, 1862: 147.

Holotype SAM-—21623, adult unsexed from Clanwilliam, 32°11’S 18°54’E, western

Cape Province, South Africa, collected by Mr J. Russouw.

Remarks

Formerly mounted; in fair condition but somewhat foxed from exposure in the public

galleries. Named in honour of Mr Edgar Leopold Layard, then Curator of the South

African Museum, who sent the specimen to Dr K. J. G. Hartlaub for study as a possibly

new taxon. The collection locality given on the label does not correspond with that which

Hartlaub (1862) quoted from Layard’s letter accompanying the specimen, viz. ‘received

from Zwartland, in the Malmesbury [33°27’S 18°44’E] division’. Layard (1867) reiterated

these data in his book. Winterbottom (1957b) drew attention to the conflict without

resolving it but obviously considered that the Swartland was where the specimen came

from since Winterbottom (1968, map 27) believed that Clanwilliam birds were aridicola

(see the next taxon below). However, Clancey (1963, 1980) placed the Cedarberg Moun-

tains, against which Clanwilliam lies, within the range of the nominate race. Hockey et

al. (1989) in their Atlas showed that in the 1980s Clanwilliam quarter-degree square

(3218 BB) was one of the few places where the species was regularly reported. They also

show that it is now scarce to absent in the Swartland and that the only place where it

occurs regularly is on the Piketberg massif, 32°42’S 18°33’E, largely in Moravia quarter-

degree square (3218 DC). Of course, one does not know by how much further the

grainlands of the Swartland have been transformed since 1860. Nonetheless, I believe that

Clanwilliam is the real type locality. The type shows most of the characters laid down by

Clancey (1963) for the nominate subspecies. Dr P. A. Clancey has examined both types

of this species for me and advises that no change in nomenclature is required, even if the

type locality is accepted as Clanwilliam. Now known as Parisoma layardi layardi Hart-

laub (Clancey 1980).

Parisoma layardi aridicola Winterbottom, 1958b: 148.

Holotype SAM—21565, adult male from Noisabis, 28°27’S 17°11’E, Richtersveld,

north-western Cape Province, South Africa, collected by Dr J. M. Winterbottom on

25 March 1958.

Remarks

In good condition. The epithet refers to the aridity of the Richtersveld where the type

was collected. Now known as Parisoma layardi aridicola Winterbottom (Clancey 1980).

AVES TYPE SPECIMENS IN THE SOUTH AFRICAN MUSEUM 341

Calamoherpe gracilirostris Hartlaub in Gurney, 1864: 348.

Holotype SAM—15093, unsexed adult from the Liesbeeck River, 33°56’S 18°28’E,

Cape Town, Cape Province, South Africa, collected by Mr E. L. Layard or his son

(Layard 1867) on an unknown date.

Remarks

Formerly mounted; in good condition. The epithet draws attention to the slender bill

vis-a-vis that of other species then placed in Calamoherpe. Now known as Acrocephalus

gracilirostris gracilirostris (Hartlaub) (Clancey 1980).

Prinia pectoralis etoshae Winterbottom, 1964: 59.

Holotype SAM-54841, adult male from Leeubron, 19°04’S 15°50’E, Okaukuejo,

Etosha National Park, Namibia, collected by Mr Miles Berkeley Markus on 20 August

1963.

Remarks

In good condition. The epithet refers to the Etosha National Park, where the type was

collected. Dr Markus is now a professor of zoology at the University of the Witwaters-

rand. The species is not a Prinia (Maclean 1974; Brooke & Dean 1990). Now known as

Malcorus pectoralis etoshae (Winterbottom) (Clancey 1980).

Family Muscicapidae Fleming

Erythrocercus francisi W. L. Sclater, 1898: 60.

Holotype SAM-2077, adult female (no locality given) collected by Mr H. F. Francis

on 18 November 1897.

Remarks

In good condition. Named in honour of the collector who sent his Mozambican and

eastern Transvaal bird specimens to the South African Museum. Some of the Mozambi-

can specimens were obtained by his brother, Mr W. Francis. The specimen came from

near Inhambane, 23°52’S 35°23’E, Inhambane Province, Mozambique (Sclater 1899a).

Now known as Erythrocercus livingstonei francisi Sclater (Clancey 1980).

Family Malaconotidae Swainson

Laniarius atrocroceus Trimen, 1880b: 623.

Holotype SAM-—58323, adult unsexed from the Crocodile [= Limpopo] River in the

north-western Transvaal, South Africa, collected by Dr B. F. Bradshaw on 25 November

879.

Remarks

Formerly mounted; in very good condition. The specimen was collected at Stock-

poort, 23°24’S 27°20’E, on the Limpopo River where the Tropic of Capricorn crosses it.

This is 100 km more or less south-east of Shoshong, 22°57’S 26°29’E, Botswana, as

stated by Trimen (1880b). The epithet (black and yellow) parallels that of L. atrococ-

cineus (Burchell) (black and red) of which it is an aberrant specimen with the deep red

342 ANNALS OF THE SOUTH AFRICAN MUSEUM

replaced by rich yellow (Stark & Sclater 1901), as Trimen (1880b) suspected. A similar

yellow aberration in the normally red L. barbarus (Linnaeus) has recently been reported

from Senegal (Ndao 1989). For Dr Bradshaw see Coracias spatulata above (p. 334). Now

known as Laniarius atrococcineus (Burchell) (Clancey 1980).

Laniarius maraisi W. L. Sclater, 1901: 183.

Syntypes SAM-4866, adult male from Craddock’s Bush collected on 3 November

1898, and SAM-—4867, adult female from Salt River, 34°02’S 23°02’E, collected on

14 May 1898, both by Johann van Oosterzee Marais.

Remarks

The male is in good condition but in worn plumage. The female is in fair to good

condition but in fresh plumage. Both localities are near Knysna, 34°02’S 23°02’E, south-

ern Cape Province, South Africa. Named in honour of the collector, then in the Forestry

Department of the Cape Colony (Anon. 1905). Curiously, Sclater (1930) did not mention

Laniarius maraisi in his Afrotropical bird list.

Sclater (1901) pointed out that some birds shot in Laniarius maraisi plumage were

in breeding condition. The state of wear of the primaries of both syntypes is consistent

with their being adults which moult after breeding, as do nearly all passerines. Laniarius

maraisi 1s now held to be the juvenile/immature plumage of Telophorus olivaceus

olivaceus (Shaw) (Clancey 1980). It would appear that in Telophorus olivaceus reproduc-

tive maturity does not correlate with plumage maturity, as Harris (1988) has remarked in

respect of females, and that a juvenile-type plumage may be retained for a long time,

perhaps throughout life. It is a pity that Hall et al. (1966) did not consider L. maraisi in

their study of plumage polymorphism in this and related bush-shrikes. I believe that

L. maraisi is yet another colour phase in this polymorphic species. Whichever view is

correct, L. maraisi Sclater is not a zoological taxon.

Laniarius maraisi is found around Knysna. Extreme southern and south-eastern

Africa is the base for a number of geographically restricted polymorphisms in plumage:

Accipiter melanoleucus melanoleucus Smith (Accipitridae) (melanistic morphs) (Malan

1988; A. J. van Zyl, pers. comm. on the Tarkastad District); Clamator jacobinus serratus

(Sparrman) (Cuculidae) (melanistic morph) (Rowan 1983); Alcedo semitorquata semitor-

quata Swainson (Alcedinidae) (whitish belly morph) (Clancey 1978); Hirundo paludicola

paludicola (Vieillot) (Hirundinidae) (brown belly morph) (Maclean 1985; pers. obs.);

Campephaga flava (Vieillot) (Campephagidae) (yellow shoulder morph) (Maclean 1985);

Telophorus olivaceus olivaceus (olive-yellow and ‘immature’ or maraisi morphs) (Hall et

al. 1966; Clancey 1967, 1969); and perhaps others. All these appear to be subspecies in

the throes of being absorbed by more northerly ones with larger ranges.

The morphs probably originated as subspecies during Pleistocene glaciations, when

the fall in sea-level markedly extended the land area of the extreme south of Africa

(Dingle & Rogers 1972, particularly their figs 3a and 4a). Much of the exposed land was

grassland (Avery 1982) but forests would have extended southwards along the major

rivers, many of which had exceedingly shallow gradients leading to extensive marshy

conditions (Dingle & Rogers 1972). It will be noted that the species listed above as

showing geographically limited polymorphism are mostly species of forest and forest

edge, or marshland in the case of R. paludicola. There is so little natural grassland left in

AVES TYPE SPECIMENS IN THE SOUTH AFRICAN MUSEUM 343

the southern Cape Province that subspecific taxa that developed on the grassland areas to

the south of the present coastline, if any, have probably become extinct without leaving

a trace, unlike the grazing antelope Hippotragus leucophaeus (Pallas) (Klein 1974).

Family Sturnidae Rafinesque

Lamprotornis mevesii violacior Clancey, 1973: 279.

Holotype SAM-—16308a, adult male from Otjibombe (= Otjumbumbe), Nankwale

Rapids on the Cunene River between Rua Cana and Eriksson’s Drift, Namibia, collected

by Mr A. G. White on 23 March 1923.

Remarks

In very good condition. Otjumbumbe lies at 17°24’S 14°25’E (cf. Tarboton 1967).

The site is probably covered by the waters of the Rua Cana Dam. The epithet indicates

the more purplish tinge on the foreparts compared with the nominate race. Now known

as Lamprotornis mevesii violacior Clancey (Clancey 1980).

Family Nectariniidae Vigors

Cinnyris erikssoni Trimen, 1882: 451.

Syntypes SAM-22045a, SAM-22045c and SAM-—22045d, adult male and two

females, respectively, from Shella, Angola, collected by Axel Wilhelm Eriksson in 1883,

obviously 1882 as stated by Trimen (1882).

Remarks

SAM-—22045c was formerly mounted; all syntypes in good condition. Unlike the

other two syntypes, the former mount is stated to have been collected in 1882. Shella is

an anglicized spelling of Chela, an escarpment lying west of Lubango and separating the

plateau from the coastal lowlands. It is also the boundary between the Huila and Namibe

Provinces of Angola in that sector. Named in honour of the collector who did much to

make the ornithology of Namibia and adjacent countries known. Rudebeck (1955) gave

a detailed account of Eriksson’s ornithological activities but noted that there was no data

for the period 1881 to 1883. This was the period of his residence in south-western Angola

when he sent some, at least, of his material to the South African Museum. He had moved

to Angola to avoid the troubles created by the Second Nama—Herero War (Tabler 1973)

and to open a new route for driving cattle and other produce to the sea at Mocamedes

(now Namibe) (Brinck 1955). Now known as Nectarinia ludovicensis ludovicensis du

Bocage (Clancey & Irwin 1978).

Family Ploceidae Sundevall

Hyphantornis nigriceps Layard, 1867: 180.

Holotype SAM-1968, adult male in nuptial plumage from Kuruman, 27°28’S

23°26’E, northern Cape Province, South Africa, collected by Rev. Robert Moffat at an

unknown date.

344 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

Formerly mounted; in fair condition but lacking a tail. Grant & Mackworth-Praed

(1957) corrected the type locality to Bulawayo, 20°10’S 28°43’E, Zimbabwe, on the

adequate grounds that H. nigriceps is not known from anywhere near Kuruman, Rev.

Moffat’s missionary base from which he travelled widely, including to Bulawayo. Named

for its black as opposed to yellow crown. Now known as Ploceus cucullatus nigriceps

(Layard) (Clancey 1980).

Ploceus velatus inustus Clancey, 1959: 173.

Holotype SAM—20427, adult male in nuptial plumage from Lokenburg, 31°41’S

19°11’E, western Cape Province, South Africa, collected by Dr J. M. Winterbottom on

21 October 1956.

Remarks

In good condition. The epithet refers to the relative absence of the pectoral fiery

reddish brown wash in the nuptial plumage of this subspecies. Now known as Ploceus

velatus velatus Vieillot (Clancey 1980; Brooke 1985).

Euplectes orix turgida Clancey, 1958: 96.

Holotype SAM-—20218, adult male in nuptial plumage from Citrusdal, 32°35’S 19°O1’E,

western Cape Province, South Africa, collected by Dr J. M. Winterbottom on 20 October

1955.

Remarks

In good condition. The epithet refers to the increased size of this extra-tropical

subspecies compared with tropical ones. Now known as Euplectes orix turgidus Clancey

(Clancey 1980).

Family Carduelidae Vigors

Serinus flaviventris quintoni Winterbottom, 1959a: 318.

Holotype SAM-—20188, adult male from Hillmore Farm, 32°30’S 22°49’E, Beaufort

West, Great Karoo, South Africa, collected by Dr J. M. Winterbottom on 22 September

IDS.

Remarks

In fair to good condition. Named in honour of Mr William F. Quinton, a local

ornithologist and owner of the farm on which the type was collected (cf. Quinton &

Winterbottom 1968). Now known as Serinus flaviventris quintoni Winterbottom (Clancey

1980).

Serinus atrogularis lwenarum White, 1944: 40.

Holotype SAM—20098, adult male from Balovale, 13°33’S 23°07’E, North-western

Province, Zambia, collected presumably by Mr Charles Matthew Newton White, on

27 May 1943.

AVES TYPE SPECIMENS IN THE SOUTH AFRICAN MUSEUM 345

Remarks

In good condition. The epithet refers to the Lwena people in whose territory the type

was collected. Now known as Serinus atrogularis lwenarum White (Benson et al. 1971).

Family Emberizidae Vigors

Fringillaria tahapisi nivenorum Winterbottom, 1964: 73.

Holotype SAM-—54934a, adult male from Otjivasando (correctly Otjovasandu),

19°09’S 14°30°E, Kaokoveld, north-western Namibia, collected by Dr J. M. Winter-

bottom on 9 September 1963.

Remarks

In poor condition. Named in honour of the Niven family of Amanzi in the eastern

Cape Province and chiefly in honour of Mr and Mrs J. P. Mackie Niven. Mrs Cecily

Kathleen (J. P. M.) Niven, daughter of Sir Percy FitzPatrick, founded the Percy

FitzPatrick Institute of African Ornithology, which started work in 1960 and of which

Dr Winterbottom was the first Director. Now known as Emberiza tahapisi nivenorum

(Winterbottom) (Clancey 1980).

ACKNOWLEDGEMENTS

I am much obliged to Dr P. A. Clancey for some invaluable comments on a draft of

this catalogue and for examining the two types of Parisoma layardi for me. I am obliged

to Miss P. J. Haarhoff for help with the fossil types.

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aay) ras

- 4 7 7 - >

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6. SYSTEMATIC papers must conform to the International 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.,

Cle.

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. Du Toit 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.

R. K. BROOKE _

ANNOTATED CATALOGUE OF —

THE AVES TYPE SPECIMENS —

IN THE SOUTH AFRICAN MUSEUM

3 9088 01206 701

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