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ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 104 Band

October 1994 Oktober

Part 5 Deel

OSTRACODA FROM QUATERNARY

COASTAL SEQUENCES

IN THE SOUTH-WESTERN CAPE

R. V. DINGLE

A. HONIGSTEIN

Cape Town Kaapstad

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D3294

OSTRACODA FROM QUATERNARY COASTAL SEQUENCES

IN THE SOUTH-WESTERN CAPE

R. V. DINGLE

Micropalaeontology Research Unit, South African Museum, Cape Town

A. HONIGSTEIN

Oil & Gas Section, Ministry of Energy & Infrastructure, Jerusalem, Israel

(With 20 figures and 3 tables)

[MS accepted 18 April 1994]

ABSTRACT

Quaternary sediments from the south-western Cape coastal plain were examined for their

ostracod faunas. Seventeen species were recovered, four of which are new: Paranesidea

verlorevleiensis, Cytheromorpha milleri, Cyprideis draaihoekensis and Caudites tankardi. A

new generic name (Garciaella) is proposed for the pre-occupied Bensonia.

Two sedimentary successions (at Draaihoek, and in the Gypsum Quarry, south of Cape

Deseada) contain relatively diverse ostracod faunas and a preliminary palaeoecological

assessment is made for each, using a combination of empirical and multivariate (Q-mode

factor) analyses based on the known, modern ecological preferences of various species. Both

indicate initial sedimentation under normal marine conditions, followed progressively by

high-salinity marine and freshwater deposition. At the Gypsum Quarry, the upper part of the

sequence consists of a gypsiferous evaporite with evidence of periodic freshwater influxes,

whereas at Draaihoek there is a reversion to a high-salinity marine environment, followed by

a second period of deposition under freshwater conditions. Strontium isotope analysis of

marine molluscs from the base of the Gypsum Quarry indicate an age of < 500 kyr (late

Pleistocene-Holocene).

CONTENTS

PAGE

BSLEROSHIC ELON ice ys ste ye eee Oe ST TaN is ITE IA La RIA EE 63

@Callectmpslocalities'andi sedimentary SEQUENCES 2. s-e ance sncncoesceaescacer- seca estiacme meen 64

S SIRITSTIVE LITE AR eR S OEE OC EERE OCAOCECE TON OBESE EES ia Smart etiam aia mn ce Hr BNA A 67

FRESU STATO IGISCUSSION ES rates occa a TOS oS ee aes 97

ESTIV EON Meri tatyiNGI CALOTS Meee Gasset Toe corer ec sith eT es Ee Lae 97

BAlACOeNVATONINEN UES acces cece rt omer rerio RO etc lezen CA Uno Re eRe TE 102

PRC KATES LED CENICTIES ade rare see et soe soe oes eR erect TaN See or cO RCo ees ator EN Soa se 109

CLC CH CES eam etre ee ey Ae Cha enue Ha Rea H ODL AY AGMoeN exten NON ein as ounce 109

PRBS PIC HOLY Ra esercto es Sato ts ciselas em va Cercd jercte os RS OeTe re wis sie re eeaba ro ale WEI REC SE Rae eal econ Mele ates 113

|. VEIN Poe COCR BPEN GCE GAOT HBOS ES ROCCE CRO TEER CSAC USED HOST AE CSE er CET ene entin enna en amr 114

INTRODUCTION

Tankard (1976, table 2) recorded 23 species of Ostracoda from Quaternary

sediments in the coastal region of the south-western Cape. As part of an investi-

gation of Quaternary palaeoenvironments, undertaken in collaboration with the

Department of Archaeology at the University of Cape Town (UCT), we have

re-examined several of Tankard’s original localities, as well as excavating new

sections, to verify the previous faunal lists, and to better establish the geo-

graphical and temporal ranges of the various species.

Ann. S. Afr. Mus. 104 (5), 1994: 63-114, 20 figs. 3 tables.

64 ANNALS OF THE SOUTH AFRICAN MUSEUM

Here, we present taxonomic notes on the species that we recovered,

including the description of four new species of ostracods. A comprehensive

palaeoenvironmental assessment of the faunas, including sedimentological

and stratigraphical data, will be presented elsewhere but, in the meantime,

summary data on sites, sections and some environmental implications are

included herein.

Previous relevant studies on coastal, inner-shelf and continental Ostracoda

from south and south-western Africa have been made by Brady (1880), Miller

(1908), Sars (1924), Klie (1940), Benson & Maddocks (1964), McKenzie

(1971, 1977), Hartmann (1974), Keeler (1981), Boomer (1985) and Dingle

(1992, 1993, 1994).

COLLECTING LOCALITIES AND SEDIMENTARY SEQUENCES

We collected sediment samples from the south-western Cape coastal area at

10 sites between Churchhaven and the Olifants River mouth (Fig. 1). Ostracods

were present at most sites, but only at Draaihoek and the Gypsum Quarry (south

of Cape Deseada) were there sufficiently long sequences for an evolving palaeo-

ecology to be interpreted from the ostracod faunas. Nevertheless, material from

the other sites allows us to extend our regional interpretations and supplement

previous palaeoenvironmental analyses. Figure 1 also includes the localities of

offshore samples utilized by Dingle (1992, 1993, 1994), which contained

species recorded in the present study. Sections were excavated, but found to be

lacking in ostracods at Malkop Pan (32°8.2’S 18°18.4’E), Soutkloof (32°25.0’S

18°20.0’E), and Churchhaven (c. 33°10.0’S 18°04.0’E). The last site, in

particular, was disappointing: Tankard (1976, sampling sites 21, 22, table 2)

had recorded an extensive ostracod fauna from Churchhaven, but despite taking

16 samples from a variety of horizons and lithologies, no specimens were found

in the present study.

Olifants River mouth (31°42.5’S 18°12.5’E)

A 4.08-m section was excavated in clays and sandy clays 1 300 m south of

Papendorp, on the eastern side of a large salt pan. Only sample 64 (soft, light

greenish-grey sandy clay with abundant small gastropods and Solen, 3.41 m

above the quarry floor) contained ostracods.

Verlorevlei (Quarry: 32°19.5’S 18°22.25’E; VMS-9: 32°19.0’S 18°20.0’E)

Outcrops of palaeo-estuarine deposits have been described from the southern

side of the modern vlei by Tankard (1976: 98-99), Miller (1987) and Miller er

al. (1993). We collected from a small disused quarry immediately south of the

road at Verlorevlei settlement, where bedrock with attached oysters at elev-

ations from 4.47 m to 6.78 m above present mean sea-level (Miller 1987: 53) is

overlain by a basal shelly, sandy rudite, and shelly coarse sand with occasional

rounded quartzite pebbles. These sediments lie on a steeply inclined bedrock

surface, and the succession in this quarry has been accurately surveyed by

members of the Department of Archaeology at UCT, but remains unpublished

OSTRACODA FROM QUATERNARY COASTAL SEQUENCES 65

(Miller pers. comm. 1993). Ostracods were recovered from the coarse basal

unit (sample 24 at 2.45 m above sea-level) and the overlying sands (sample 26

at c. 2 m above sea-level). We also recorded ostracods from shelly sand

horizons of samples 7 and 9 (c. 4 m above sea-level) in the VMS-9 hole sunk

by Miller et al. (1993).

Gypsum Quarry (site 1—North end: 32°21.15’S 18°19.69’E; site 2—South end:

a2521-207S 18°19 TE)

This is a locality originally described by Tankard (1976: 88), c. 4.2 km

south of Cape Deseada. It lies between 4.5 and 5 m above sea-level, immedi-

ately to the east of the iron-ore railway, and is an abandoned opencast

excavation. Tankard (1976: 88) recorded a sequence of shelly calcareous sands

overlain by up to 75 cm of ‘rhythmically laminated . . . carbonate-gypsum-

halite units’. We dug two pits into the sands beneath the gypsiferous horizon

and revealed 1.14 m of whitish and yellow shelly sands at site 1 (north),

and 1.0 m of greenish sands at site 2 (200 m farther south). The sands in

both sequences rest on greenish, nodular clays. Practically the whole sequence

at site 1 is ostracodiferous, and a strontium isotope ratio date on whole

Choromytilus shells from the lowermost sand (sample 78) gives a date of

< 500 kyr (Late Pleistocene-Holocene: Lavelle & Armstrong 1993; Lavelle in

prep.).

Draaihoek (32°29.0’S 18°20.25’E)

Situated 18.5 km south of Cape Deseada on the farm Draaihoek, immedi-

ately west of the coast road. Here we excavated two small pits on the west side

of the farm dam and about 200 m apart. Assuming that the local sequences are

essentially horizontal, the two reveal a composite section approximately 2.23 m

thick consisting of c. 98 cm of whitish clays and sandy clays, overlying 1.25 m

of white and green sands with a lower layer of shelly, more clayey sands with

abundant paired bivalves (primarily Venerupis corrugata and Choromytilus

meridionalis—J. Pether pers. comm.). Practically the whole of this sequence is

ostracodiferous.

Velddrif (32°47.0’S 18°10.0’E)

This is the section described by Tankard (1976, localities 9-10) as the inner,

or Velddrif bar. The shelly sands lie in banked units 15-30 cm thick and

contained ostracods at two levels (samples 11, 14; 80 cm and 170 cm above the

base, respectively).

Laaiplek (Refuse Pit: 32°45.0’S 18°10.5’E; Parrot Grit Quarry: 32°44.5’S

18°10.5’E)

These two sites lie on the eastern edge of Tankard’s (1976, fig. 10) outer, or

Laaiplek bar. They expose fine-medium sands with abundant comminuted

shells. Ostracods were recovered from one sample in each quarry: Refuse Pit

(sample 15; 50 cm above base) and Parrot Grit Quarry (sample 19; 165 cm

above base).

66 ANNALS OF THE SOUTH AFRICAN MUSEUM

Olifants River

____—. Papendorp

TBD 3892 x

LAMBERTS BAY

Verlorevlei Quarry

C. DESEADA

Gypsum Quarry

ST HELENA

BAY

TBD 3089 |

C. COLUMBINE

Berg River

SALDANHA BAY

Churchhaven

TBD 2224

Fig. 1. Ostracodiferous sampling sites in Quaternary sediments, south-western Cape.

Coordinates of localities and summaries of the sedimentary sequences are given in the text.

TBD sites are offshore samples from the Thomas B. Davie dataset used by Dingle (1992,

1993, 1994).

OSTRACODA FROM QUATERNARY COASTAL SEQUENCES 67

SYSTEMATICS

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; 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; RPC = radial pore canal; RV = right valve; SCT = sub-

central tubercle; TE = terminal element; VM = ventral margin.

Type and illustrated specimens are housed at the South African Museum

under catalogue numbers prefixed SAM-PQ-MF-.

Notation for sites: DH = Draaihoek; GQ = Gypsum Quarry (1); VMS =

Vlei Mouth Sounding (at Verlorevlei); VQ = Verlorevlei Quarry.

Class CRUSTACEA Pennant, 1777

Subclass OSTRACODA Latreille, 1806

Order PODOCOPIDA Miller, 1894

Suborder PODOCOPIDA Sars, 1866

Superfamily CYPRIDACEA Baird, 1845

Family Cyprididae Baird, 1845

Subfamily Cypridinae Baird, 1845

Genus Heterocypris Claus, 1893

Heterocypris capensis (Miller, 1908)

Fig. 2A-F

Cyprinotus capensis Muller, 1908: 162-163, text-figs 1-7.

Heterocypris capensis (Miller, 1908) Sars, 1924: 118-119, pl. 4 (figs 5-20).

Illustrated material

length height

MF-1667, RV, DH sample 43 1.19 0.69

MF-1668, LV, DH sample 43 1.43 0.80

MF-1669, C, DH sample 43 1.30 0.59

MF-1670, LV, DH sample 43 1.20 0.69

MF-1671, RV, DH sample 43 a3) 0.72

Remarks

Adult valves of this relatively large, thin-shelled species occurred mostly

fragmented in our samples. As shown by illustrations in both Miller (1908) and

Sars (1924), it is characterized by an upcurved AM in the RV, which in dorsal

view is wrapped around by the AM of the larger LV. We present SEM micro-

graphs of the MS and hinge of this species for the first time.

68 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 2. A-F. Heterocypris capensis (Miller, 1908), Draaihoek, sample 43. A. MF-1667,

RV, SEM 4414. B. MF-1668, LV, SEM 4482. C. MF-1669, C, dorsal view,

SEM AH261. D. MF-1670, LV, internal view, SEM AH253. E-F. MF-1671, RV,

internal view. E. SEM 4453. F. Detail, anteroventral area, SEM 4454.

G-H. Sarscypridopsis ?aculeata (Costa, 1847). G. MF-1672, LV, GQ, sample 80.

H. MF-1673, LV, DH, sample 39. Scale bars: A-E, G-H = 100 p; F = 10 uz.

OSTRACODA FROM QUATERNARY COASTAL SEQUENCES 69

Distribution

McKenzie (1971) summarized the localities from which this species had

previously been reported, and these were confined to the freshwater ponds and

lakes in the region between Saldanha Bay and Fish Hoek: Zeekoevlei,

Plumstead, Fish Hoek, Bergvliet and ‘near Cape Town’ on the Cape Peninsula,

and near the old whaling station in Saldanha Bay. In the present study, Hetero-

cypris capensis was restricted to Draaihoek, where it forms an important

element of the fauna, particularly in the upper section (up to 46%).

Genus Sarscypridopsis McKenzie, 1977

Sars (1924) reported 17 species of this genus (as Cypridopsis) from the Cape

Province, 12 of which he recognized in the south-western Cape. Of these, nine

have generally rounded sub-triangular lateral and elliptical dorsal outlines (Sars-

cypridopsis gregaria, S. spinifera, S. aculeata, S. reniformis, S. tonsa,

S. ochracea, S. echinata, S. trigonella and S. brevis), so that subdivision into

different species, particularly in juvenile forms, is an uncertain exercise. We

have tentatively identified two of the species recorded by Sars (1924), one with

a distinctly triangular outline (S. aculeata), and the other with a distinctly reni-

form outline (S. renifermis). We feel that it would be inappropriate to attempt to

isolate some of the more intermediate types (e.g. S. tonsa).

Sarscypridopsis ?aculeata (Costa, 1847)

Figs 2G-H, 3A-E

Compare:

Cypris aculeata Costa, 1847: 11, pl. 3 (fig. 5).

Cypridopsis aculeata (Costa, 1847) Brady, 1867: 117. Sars, 1924: 160-161,

pl. 14 (figs 3-4).

‘Cypridopsis’ aculeata (Costa, 1847) McKenzie, 1971: 167.

Sarscypridopsis aculeata (Costa, 1847) McKenzie, 1977: 49. De Deckker,

1981: 81-84, figs 27-28.

Illustrated material

length height width

MF-1672, LV, GQ sample 80 0.46 0.30 —

MF-1673, LV, DH sample 39 0.52 0.36 —

MF-1674, RV, DH sample 39 0.49 0.34 —

MF-1675, C, DH sample 41 0.48 — 0.24

MF-1676, RV, DH sample 42 0.53 0.38 —~

MF-1677, LV, DH sample 47 0.49 0.31 _

Remarks

Sars (1924) distinguished this species from two of his own (Sarscypridopsis

gregaria (Sars, 1895) and S. spinifera (Sars, 1924)) on the basis of its ‘some-

what more steeply’ sloping posterodorsal margin, and its shorter and stouter

spines (which, however, are not preserved fossilized). From his illustrations

70 ANNALS OF THE SOUTH AFRICAN MUSEUM

(1924, pls 13-14), however, the three species appear to be very similar in

outline, and we are not confident of our choice between them. De Deckker

(1981) considers S. spinifera to be synonymous with S. aculeata, and that

5. aculeata includes both spinose and non-spinose forms (such as Cypridopsis

obstinata Barclay, 1968, from New Zealand).

Distribution

Sarscypridopsis aculeata has been recognized throughout Europe, Iceland,

central Asia and North Africa (Sars 1924), Australia and New Zealand (De

Deckker 1981), and in South Africa from the vicinity of the Cape Peninsula,

and at one site in the Transvaal (McKenzie 1971). Sars himself recorded it only

from a pool on the Cape Flats, whereas he noted S. gregaria from Knysna,

Bergvliet and Saldanha Bay, and S. ochracea from the Cape Flats and near

Cape Town.

De Deckker (1981) remarked that this cosmopolitan species is commonly

found in temporary pools in Australia. He quoted a salinity range of fresh to

11.2%o (one exceptional record of 21.3%c), and noted that these values are

much higher than recorded from Europe; he suggested that the species has

adapted to more arid conditions in Australia. Presumably, a similar logic could

be applied to the southern African populations.

In the present study, we recorded this species at three localities: at Draai-

hoek it occurs throughout the upper section, and is the dominant species in the

top part (30-74%); as a minor component at four levels scattered throughout the

Gypsum exposure (maximum 8%); and at site 11 at Velddrif.

Sarscypridopsis ?reniformis (Sars, 1924)

Figs 3F-H, 4A-C

Compare:

Cypridopsis reniformis Sars, 1924: 161-162, pl. 14 (figs 7-8).

Illustrated material

length height width

MF-1678, LV, DH sample 39 0.69 0.41 —

MF-1679, RV, GQ sample 86 0.55 0.32 —

MF-1680, C, GQ sample 86 0.60 — 0.29

MF-1681, RV, DH sample 39 0.63 0.40 —

MF-1682, LV, DH sample 39 0.65 0.37 —

Remarks

This species is recognized on the basis of its reniform outline, where Sars

(1924) emphasized that the highest point of the valve lies at about mid-length. It

is distinguished from Sarscypridopsis aculeata by its more elongate outline, and

from the similar §. ochracea, which has its highest point consistently in the

anterior half of the valve.

OSTRACODA FROM QUATERNARY COASTAL SEQUENCES UM

Fig. 3. A-E. Sarscypridopsis ?aculeata (Costa, 1847), Draaihoek. A. MF-1674, RV,

sample 39, SEM 4412. B. MF-1675, C, dorsal view, sample 41, SEM 4485. C. MF-1676,

RV, internal view, sample 42, SEM AH243. D-E. MF-1677, LV, internal view,

sample 47. D. Detail, anteroventral area, SEM 4431. E. MS, SEM 4429. F-H. Sars-

cypridopsis ?reniformis (Sars, 1924). F. MF-1678, LV, Draaihoek, sample 39, SEM 4408.

G-H. Gypsum Quarry, sample 86. G. MF-1680, C, dorsal view, SEM 4388. H. MF-1679,

RV, SEM 4386. Scale bars: A-C, E-H = 100 pn; D = 10 np.

fz ANNALS OF THE SOUTH AFRICAN MUSEUM

Distribution

Sars (1924) recorded this species only from near Fish Hoek station on the

False Bay coast. In the present study, we found it at three localities: it is most

common at Draaihoek, where the species occurs at almost every level, being

most abundant in the lower part of the upper section; in the Gypsum Quarry,

where it is moderately abundant at various levels throughout the sequence; and

in sample 11 from Velddrif.

Tankard (1976) recorded one species of Sarscypridopsis (as Cypridopsis

ochracea) from the Gypsum Quarry. We suspect that this is most likely to have

been §. reniformis.

Family Paracyprididae Sars, 1923

Genus Aglaiella Daday, 1910

Aglaiella railbridgensis Benson & Maddocks, 1964

Fig. 4D-H

Aglaiella railbridgensis Benson & Maddocks, 1964: 16-17, pl. 1 (figs 7, 9-10), text-fig. 7.

Hartmann, 1974: 357-358, pl. 138 (figs 952-961).

Illustrated material

length height

MF-1683, LV, GQ sample 85 0.90 0.40

MF-1684, RV, GQ sample 85 0.85 0.35

MF-1685, LV, GQ sample 85 0.90 0.42

MF-1686, RV, GQ sample 85 0.84 0.35

Remarks

Hartmann (1974) described a further species of Aglaiella (A. kenmckenziei)

from the coast of Angola, which is distinguished from Benson & Maddocks’s

species by having a less strongly arched DM and a slightly different arrange-

ment of the MS. Our material is comparable with A. railbridgensis in both these

aspects.

Distribution

Previously recorded only from the railbridge and Leisure Island sites in

Knysna Lagoon by Benson & Maddocks (1964) and Hartmann (1974), where

salinities range 30-31%o and 33-35%, respectively, and the substrates are

muddy sand and fine sands (Benson & Maddocks 1964).

In the present study, Aglaiella railbridgensis is relatively rare, being found

only as a single valve in the Verlorevlei Quarry (sample 24), and in small

numbers (up to 4%) at the top of the Gypsum Quarry section (samples 85-86).

Tankard (1976) listed this species as a minor component at Kruispad

(sample 12).

OSTRACODA FROM QUATERNARY COASTAL SEQUENCES 13,

Fig. 4. A-C. Sarscypridopsis ?reniformis (Sars, 1924), Draaihoek, sample 39.

A. MF-1681, RV, internal view, SEM AH233. B. MF-1682, LV, internal view,

SEM 4409. C. MS, SEM 4411. D-H. Aglaiella railbridgensis Benson & Maddocks, 1964,

Gypsum Quarry, sample 85. D. MF-1683, LV, SEM 4396. E. MF-1684, RV,

SEM AH166. F. MF-1685, LV, internal view, SEM AH167. G. MF-1686, RV, internal

view, SEM AH172. H. MF-1685, LV, MS, SEM AHI171. Scale bars: A-B, D-G =

100 n; C, H = 10n.

74 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus Paracypris Sars, 1866

Paracypris westfordensis Benson & Maddocks, 1964

Fig. 5A-G

Paracypris westfordensis Benson & Maddocks, 1964: 15-16, pl. 1 (figs 4, 11-12), text-

fig. 6.

Illustrated material

length height width

MF-1687, LV, GQ sample 86 0.91 0.45 —

MF-1688, RV, GQ sample 86 0.92 0.40 —

MF-1689, C, GQ sample 86 0.96 — 0.40

MF-1690, LV, GQ sample 85 1.00 0.49 —

MF-1691, RV, DH sample 49 0.92 0.40 —_

Remarks

The MS of our specimens are identical with those illustrated by Benson &

Maddocks (1964), in particular the large ‘cap’ adductor scar.

Distribution

Previously, this species had been recorded unequivocally only from the

Knysna estuary, where it was confined to the upper reaches at the Westford

Bridge and Ashford (one, probably transported valve, had been recorded farther

downstream at the railbridge) (Benson & Maddocks 1964). Salinity ranges at

these two sites are 16.5-22%o0 and 18.0-24.0%o0, respectively, and the

substrates are soft black mud (Benson & Maddocks 1964). Hartmann (1974) did

not record it from any of his coastal sites along the southern and south-western

coasts of southern Africa, and Dingle (1992, 1993) did not report it offshore.

This distribution suggests a strictly brackish water habitat.

In the present study, Paracypris westfordensis is never an abundant species,

but we have recorded it from three localities. There is a single valve from the

Verlorevlei Quarry (sample 24), whereas it occurs in small numbers throughout

the Draaihoek section, reaching a maximum of 16 per cent in sample 40 in the

middle part. At the Gypsum Quarry, the species also occurs sparsely in the

middle and upper parts of the section (maximum of 3% near the top).

Tankard (1976) recorded this species at Laaiplek (5%).

Superfamily BAIRDIACEA Sars, 1887

Family Bairdiidae Sars, 1887

Genus Paranesidea Maddocks, 1969

Paranesidea verlorevleiensis sp. nov.

Figs 5H, 6A-F

Derivation of name

The name of this species is derived from Verlorevlei, the locality of the type

material.

OSTRACODA FROM QUATERNARY COASTAL SEQUENCES 75

Fig. 5. A-G. Paracypris westfordensis Benson & Maddocks, 1964. A-D. Gypsum Quarry.

A-C. Sample 86. A. MF-1687, LV, SEM AH064. B. MF-1688, RV, SEM AHO066.

C. MF-1689, C, dorsal view, SEM 4385. D. MF-1690, LV, internal view, sample 85,

SEM AH068. E-G. MF-1691, RV, internal view, Draaihoek, sample 49. E. SEM 4432.

F. MS, SEM 4434. G. Hinge and dorsal scars, SEM 4435. H. Paranesidea verlorevleiensis

sp. nov., holotype, MF-1692, RV, Verlorevlei Quarry, sample 24, SEM 4488.

Scale bars: A-E, H = 100 »; F-G = 50 yp.

76 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 6. A-F. Paranesidea verlorevieiensis sp. nov., Verlorevlei Quarry, sample 24.

A. Paratype, MF-1695, C, dorsal view, SEM 4446. B. Paratype, MF-1694, C, ventral

view, SEM 4444. C. Paratype, MF-1696, LV, internal view, SEM 4441. D. Holotype,

MF-1692, RV, SEM 4437. E. Paratype, MF-1693, LV, SEM AH437. F. MF-1745, RV,

MS, SEM 4460. G-H. Indeterminate bairdiid. G. MF-1697, LV, Gypsum Quarry,

sample 78, SEM AH161. H. MF-1698, C, right view, Verlorevlei Quarry, sample 24,

SEM 4462. Scale bars: A-E, G-H = 100 yw; F = 50 pu.

OSTRACODA FROM QUATERNARY COASTAL SEQUENCES dil

Holotype

length height width

MF-1692, RV, VQ sample 24 0.78 0.38 =

Paratypes

MF-1693, LV, VQ sample 24 0.91 0.48 —

MF-1694, C, VQ sample 24 0.89 — 0.35

MF-1695, C, VQ sample 24 0.85 _ 0.34

MF-1696, LV, VQ sample 24 0.86 0.48 =

MF-1745, RV, VQ sample 24 1.05 0.57 —

Diagnosis

Finely punctate species of Paranesidea with denticulate posterior and

anterior margins and selvages.

Description

External features. Elongate bairdiid outline, RV and LV differ in shape,

particularly the DM, which is straight in RV and gently convex in LV. VM in

RV is concave, almost straight in LV. AM outline in both valves is truncated

dorsally, PM in RV is more drawn out. AM and PM in both valves are denticu-

late. Carapace in dorsal and ventral views is extended ovate. In ventral view

AM and PM both show gapes, with contact along selvage. Overall, valve

surface is delicately punctate.

Internal features. Hinge straight, smooth. MS consist of eight round—-ovate

scars set in tight spiral. In both valves (but particularly in RV) there is a promi-

nent selvage which, anteriorly and posteriorly, is denticulate.

Remarks

Paranesidea verlorevleiensis has all the attributes of the genus as specified

by Maddocks (1969), although the DM of the LV is less strongly arched than

the type species (P. fracticorallicola Maddocks, 1969), and the surface orna-

mentation is less pronounced than in other species. Our new species is quite dis-

tinctive amongst other southern African bairdiid ostracods, with its prominently

denticulate PM and AM. Only Hartmann’s species Bairdoppilata mocamedes-

ensis and ?Bairdia problematica (which Maddocks (1991) tentatively placed in

Aponesidea) exhibit prominent marginal denticulation. The former is denticulate

only along the PM and does not have a straight DM in the RV, whereas the

latter has an upturned AM outline in the RV and a different MS pattern to our

new species. The genus is represented by several species in the Tertiary of

Tanzania (Ahmad ef al. 1991), but none are as elongate in lateral view as the

new taxon.

Distribution

Although bairdiid ostracods have been widely reported from the shelf and

coastal areas of southern Africa by various workers (e.g. Brady 1880; Benson

& Maddocks 1964; Hartmann 1974; Dingle 1992, 1993), none have recorded a

78 ANNALS OF THE SOUTH AFRICAN MUSEUM

species similar to Paranesidea verlorevleiensis sp. nov., which is confined to

one outcrop (Verlorevlei Quarry, sample 24). The only previous record of

bairdiids from coastal sediments of the region was by Tankard (1976) of Bairdia

cf. B. villosa from Churchhaven.

Maddocks (1969) suggested that the genus is typical of tropical conditions,

and that it is characteristic of very shallow sublittoral waters, and unable to

tolerate variable salinities.

Indeterminate bairdiid

Fig. 6G-H

Illustrated material

length height

MF-1697, LV, GQ sample 78 0.62 0.39

MF-1698, C, VQ sample 24 ORS 0.42

Remarks

Twenty-nine valves were recovered from sample 24 in the Verlorevlei

Quarry; one juvenile valve was recovered from sample 78 in the middle of the

Gypsum Quarry section. The latter displays a strongly tapering PM outline,

strong posteriorly directed PM denticles, and widely spaced NPC. Bairdiids are

generally regarded as marine species.

Superfamily CYTHERACEA Baird, 1850

Family Loxoconchidae Sars, 1925

Genus Palmoconcha Swain & Gilby, 1974

Palmoconcha? cf. P. peterseni (Hartmann, 1974)

Fig. 7A-B

Compare:

Loxoconcha peterseni Hartmann, 1974: 296-297, pl. 67 (figs 477-487), pl. 151

(figs 3-4).

Illustrated material

length height

MF-1699, RV, GQ sample 78 0.64 0.48

Remarks

The one specimen available to us compares favourably with Hartmann’s

species in outline, particularly the prominent posteroventral keel and the drawn-

out AM outline. Our specimen has somewhat more distinct concentric ribbing in

the central part of the valve, and the surface punctation is stronger.

OSTRACODA FROM QUATERNARY COASTAL SEQUENCES 79

Distribution

Hartmann (1974) recorded Palmoconcha peterseni from sandy boulder

beaches in Angola, and speculated that the species is currently confined to the

tropical—-subtropical sectors of the Benguela system. We recovered one corroded

specimen of this species from sample 78 in the Gypsum Quarry, and Tankard

(1976) reported Hartmann’s species from Laaiplek and Churchhaven. We

cannot confirm Tankard’s identifications, and there were no negatives of loxo-

conchid ostracods in Tankard’s collection of the Rhodes University SEM

archive.

Genus Cytheromorpha Hirschmann, 1909

Cytheromorpha milleri sp. nov.

Figs 7C-H, 8A-D, 9A

Indet. sp. 3412 Dingle, 1993: 149, fig. 84B.

Derivation of name

This species is named for Dr D. Miller, Department of Archaeology,

University of Cape Town, for his close association with, and assistance and

encouragement during, this study.

Holotype

length height width

MF-1700, RV, DH sample 36 0.60 0.29 —

Paratypes

MF-1701, RV, GQ sample 80 0.60 0.29 _—

MF-1702, LV, DH sample JP 0.58 0.28

MF-1703, RV, DH sample 37 0.58 0.28 —

MF-1704, LV, DH sample 37 0.59 0.29 a

MF-1705, C, DH sample JP 0.60 — 0.27

MF-1706, C, DH sample JP 0.58 _ 0.26

MF-1707, LV, DH sample JP 0.60 0.27 —

Diagnosis

Thin-shelled, sighted species of Cytheromorpha with three prominent ribs

adjacent to the AM, and a smooth, puncta-free band adjacent to the postero-

dorsal valve margin. The hinge ME is finely crenulate.

Description

External features. Elongate-ovate in lateral outline, AM asymmetric, with

extended anterodorsal section. PM rounded in RV, more quadrate in LV,

with distinct posterodorsal truncation. DM and VM straight, the latter slightly

obscured by overhang of inflated central part of valve. In dorsal and ventral

views, valves are almond-shaped, tapering anteriorly. There is a shallow sulcus,

just anterior to mid-length, which extends obliquely from the dorsal margin

80 ANNALS OF THE SOUTH AFRICAN MUSEUM

anteriorly to about mid-valve. Surface is overall ornamented with fine puncta

and narrow ribs that run sub-parallel to the valve margins. In the central area of

the valve, the ornamentation is somewhat coarser, and verges on finely reticu-

late. The three ribs that extend from the weak eye tubercle adjacent to the AM

are particularly prominent, the posterior-most of which crosses ribs running

ventrally and extends to the anteroventral margin. Intercostal areas are deli-

cately punctate. The valve margin in the posterodorsal area is free of puncta

and, in dorsal view, the valves display a distinctively smooth marginal band.

There are numerous prominent normal sieve pores. The MS area is indicated on

the lateral surface by a region of elongated, radiating puncta.

Internal features. Typical for the genus. The anterior MA is relatively wide

(Fig. 9A), with a prominent vestibulum, and the line of concrescence close to

the AM. There are at least 20 very short, fine MPC. The MS show a tick-

shaped anterior scar, a prominent fulchral point, and two ventrally lying scars

below the adductors. The hinge has a prominently denticulate ME, and the TE

are bilobate: the RV PTE and LV ATE consisting of elongate, strongly sub-

divided teeth.

Remarks

This species is the same as that recovered as a damaged juvenile valve from

TBD 3892 on the inner continental shelf off Namaqualand (depth 72 m, north-

west of Lamberts Bay) by Dingle (1993), and designated Indeterminate

sp. 3412. It was accompanied by specimens of Propontocypris (P.) cf.

P. subreniformis (Brady, 1880). The genus was recorded by neither Klie (1940)

nor Hartmann (1974) in their surveys of coastal sites off south-western Africa,

but Tankard (1976) listed abundant Cytheromorpha sp. at Verlorevlei

(sample 3), Laaiplek (sample 9) and Churchhaven (samples 21, 22). Because he

did not illustrate this material, it is not possible to speculate whether any of

these are conspecific with C. milleri sp. nov.

Our new species differs from Hirschmann’s (1909) type species (Cythere

fuscata Brady, 1869) in having a crenulate (as opposed to a smooth) ME, and in

the MS, which in C. milleri sp. nov. have a small, wedge-shaped third adductor

(see Fig. 8D).

Cytheromorpha knikensis Forester & Brouwers, 1985, has a more evenly

fine reticulate ornamentation, is not sighted, and has a smooth hinge ME. Two

species have very similar ornamentation to C. milleri sp. nov.: C. suffolkensis

Hazel, 1983 (Pliocene, North Carolina) and C. acupunctata (Brady, 1880) (for

example, as illustrated by Ikeya & Ueda (1988), and Yajima & Lord (1990)

from the Quaternary of the Sea of Japan, eastern China, Korea and eastern

Japan), but both differ in having significantly more tapered posterior outlines,

and somewhat coarser ornamentation in the posteroventral region. In addition,

the MA of the North American species is much wider than in C. milleri sp. nov.

The presence of an eyespot, and a crenulate hinge ME may cast some doubt

upon the placement of our species in the genus Cytheromorpha, but the typical

outline, ornamentation, MS and hingement (other than the ME crenulation)

suggest that the creation of a new genus would be premature without additional

comparative species.

OSTRACODA FROM QUATERNARY COASTAL SEQUENCES 81

Fig. 7. A-B. Palmoconcha? cf. P. peterseni (Hartmann, 1974), MF-1699, RV, Gypsum

Quarry, sample 78. A. Internal view, SEM 4463. B. SEM AH159. C-H. Cytheromorpha

milleri sp. nov. C. Holotype, MF-1700, RV, Draaihoek, sample 36, SEM AH331.

D-H. Paratypes. D. MF-1701, RV, Gypsum Quarry, sample 80, SEM AH218.

E. MF-1702, LV, Draaihoek, sample JP, SEM AH329. F-H. Draaihoek, sample 37.

F. MF-1703, RV, internal view, SEM AH283. G. MF-1704, LV, internal view,

SEM 4419. H, MF-1703, ATM and PTM, SEM 4426, 4427.

Scale bars: A-G = 100 »; H = 50 uy.

82 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 8. A-D. Paratypes, Cytheromorpha milleri sp. nov. A. MF-1704, LV, ATE and PTE,

Draaihoek, sample 37, SEM 4422, 4421. B. MF-1706, C, ventral view, Draaihoek,

sample JP, SEM AH291. C. MF-1705, dorsal view, Draaihoek, sample JP, SEM AH338.

D. MF-1704, LV, MS, Draaihoek, sample 37, SEM 4423. E-F. Garciaella knysnaensis

knysnaensis (Benson & Maddocks, 1964), Gypsum Quarry, sample 78. E. MF-1708, RV,

SEM AH153. F. MF-1709, LV, juvenile, SEM AH158. G-H. Cyprideis remanei Klie,

1940. G. MF-1710, LV, Gypsum Quarry, sample 83, SEM AH199. H. MF-1711, RV,

Gypsum Quarry, sample 84, SEM AH203. Scale bars: B-C, E-H = 100 yp; A = 50 p;

D = 10un.

OSTRACODA FROM QUATERNARY COASTAL SEQUENCES 83

Distribution

Cytheromorpha milleri sp. nov. has been recovered in large numbers from

the lower part of the Draaihoek exposure (samples 37-35 and JP), where it

constitutes 5-94 per cent (mean: 70%) of the fauna, and a single valve from the

middle part of the Gypsum Quarry exposure (sample 80). The single valve

recovered from the inner continental shelf by Dingle (1993—TBD 3892, depth

72 m) lies off. the Olifants River mouth.

Fig. 9. Anterior marginal areas. A. Cytheromorpha milleri sp. nov.,

paratype, MF-1707, LV, Draaihoek, sample JP. B. Cyprideis remanei

Klie, 1940, MF-1718, RV, Gypsum Quarry, sample 86. C. Cyprideis

draaihoekensis sp. nov., paratype, MF-1726, RV, Gypsum Quarry,

sample 86. Scale bar: 200 p.

Family Cytherettidae Triebel, 1952

Subfamily Cytherettinae Howe, 1961

Genus Garciaella gen. nov.

Bensonia Rossi de Garcia, 1969: 218-219, pl. 1 (figs la-c, 3).

non Bensonia (Cantor MS) Gray, 1847: 150.

non Bensonia Pfeiffer, 1855: 119.

non Bensonia Malaise, 1935: 166.

non Argenticytheretta Rossi de Garcia, 1969, emend. Sanguinetti, de Ornellas & Coimbra,

1991: 139.

Type species: Cytheretta argentinensis Rossi de Garcia, 1966.

84 ANNALS OF THE SOUTH AFRICAN MUSEUM

Derivation of name

This taxon is named for Dr Elsa Rossi de Garcia, who first recognized the

new genus.

Remarks

Rossi de Garcia (1969) erected the ostracod genus Bensonia to accommodate

her species Cytheretta argentinensis. In the same publication, she also erected

two related cytherettid taxa (Grekoffiana and Argenticytheretta). The non-

availability of the name Bensonia, pre-occupied for molluscan taxa (Gray 1847;

Pfeiffer 1855) led Sanguinetti et al. (1991) to transfer species previously

assigned to Bensonia to Argenticytheretta, because they assumed that the two

genera were synonymous (along with Grekoffiana: see Sanguinetti 1979).

Correspondence with Dr E. Bertels (pers. comm. 1993) on the subject of the

type material, along with our own interpretation of Rossi de Garcia’s original

descriptions, lead us to believe that Cytheretta argentinensis Rossi de Garcia

does indeed belong in a genus distinct from the types of both Argenticytheretta

and Grekoffiana. Hence, we propose the new name Garciaella to replace

Bensonia.

Garciaella knysnaensis knysnaensis (Benson & Maddocks, 1964)

Fig. 8E-F

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

Bensonia knysnaensis (Benson & Maddocks, 1964) Rossi de Garcia, 1969: 219. Keeler,

1981: 43-45, pl. 2 (figs 2-4).

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

Bensonia knysnaensis knysnaensis (Benson & Maddocks, 1964) Dingle, 1992: 32-34,

fig. 18A-F.

Illustrated material

length height

MF-1708, RV, GQ sample 78 0.80 0.41

MF-1709, LV, juv., GQ sample 78 0.68 0.35

Distribution

This species has previously been recorded from both estuarine (Leisure

Island in Knysna Lagoon) and inner continental shelf environments (between

19°S on the west coast and 24°E on the south coast) (Benson & Maddocks

1964; Hartmann 1974; Keeler 1981; Dingle 1992). Benson & Maddocks (1964)

suggested that it can tolerate salinities between 33 and 35%c, and from his study

of the environmental parameters tolerated by individual species, Dingle (1994)

suggested the following ranges and means for Garciaella knysnaensis knysna-

ensis on the continental shelf: temperature 7.1-13.28°C (10.31°C); salinity

34.6-35.18%0 (34.82 %c); dissolved oxygen 0.9-4.3 ml/I (2.53 ml/l); sand

content 47.3-95.9 per cent (68.8%); mud content 2.1-46.1 per cent (23%); total

organic matter content 0.2-8.8 per cent (3.8%).

In the present study, G. k. knysnaensis occurs in small numbers in the

middle section of the Gypsum Quarry (samples 78-79) and the lower part of the

OSTRACODA FROM QUATERNARY COASTAL SEQUENCES 85

Draaihoek sequence (sample 37, JP), whereas it is an important component of

the small assemblages recovered from the two quarries at Laaiplek (samples 15,

19). In addition, Tankard (1976) recorded the species at his outcrop sample 9 at

Velddrif.

Family Cytherideidae Sars, 1925

Subfamily Cytherideinae Sars, 1925

Genus Cyprideis Jones, 1857

Hartmann (1974) distinguished three species of Cyprideis, each occupying

particular sections along the west coast of southern Africa: C. nigeriensis

Omatsola, 1970, confined to the tropical region; C. limbocostata Hartmann,

1974, occupying the Angolan—Namibian coast as far south as Sandwich

Harbour; and C. remanei Klie, 1940, occurring in the vicinity of Liideritz. He

did not recover any species from the Cape Province or east coast of South

Africa, where the relevant ecological niches were considered to be occupied by

species of Sulcostocythere. Furthermore, he suggested that the three species of

Cyprideis were very closely related and possibly derived from a common

ancestor. Environmental data supplied by Klie (1940) and Hartmann (1974)

suggest that both C. remanei and C. limbocostata prefer coastal habitats with

relatively high salinities (see pp. 87, 89).

Our studies show that in the late Quaternary, C. remanei extended its distri-

bution at least as far south as the south-western Cape, and that it co-habitated

with a further species, C. draaihoekensis sp. nov.

Cyprideis remanei Klie, 1940

Figs 8G-H, 9B, 10A-H

Cyprideis remanei Klie, 1940: 412-415, text-figs 11-17.

Illustrated material

length height width

MF-1710, LV, GQ sample 83 Ovi 0.44 —

MF-1711, RV, GQ sample 84 0.79 0.45 —

MF-1712, LV, DH sample 35 0.80 0.45 -

MF-1713, RV, DH sample 40 0.79 0.45 —

MF-1714, LV, GQ sample 86 0.79 0.45 —

MF-1715, RV, GQ sample 86 0.80 0.45 —

MF-1716, C, GQ sample 86 0.90 — 0.42

MF-1717, C, GQ sample 86 0.80 — 0.42

MF-1718, RV, GQ sample 86 0.90 0.51 —

Remarks

Klie’s species has a distinctly quadrate outline, in contrast to the more

elongate Cyprideis limbocostata, and the intermediate shape of our new species

C. draaihoekensis. All the specimens available to us are smooth or covered in

86 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 10. A-H. Cyprideis remanei Klie, 1940. A. MF-1712, LV, Draaihoek, sample 35,

SEM AH220. B. MF-1715, RV, internal view, Gypsum Quarry, sample 86, SEM 4373.

C. MF-1714, LV, internal view, Gypsum Quarry, sample 86, SEM 4369. D. MF-1713,

RV, Draaihoek, sample 40, SEM AH221. E-F. MF-1714, LV, Gypsum Quarry,

sample 86. E. ATE and PTE, SEM 4372, 4371. F. MS, SEM 4370. G-H. Gypsum

quarry, sample 86. G. MF-1716, C, dorsal view, SEM 4475. H. MF-1717, C, ventral

view, SEM AHO028. Scale bars: A-D, G-H = 100 pp; E-F = 50 pz.

OSTRACODA FROM QUATERNARY COASTAL SEQUENCES 87

relatively small puncta, which lie in a swirl-like pattern in the sub-central region

above the MS and adjacent to the dorsomedian sulcus. Females are distinctly

bulbous in the posterior part of the valve.

The MS pattern of C. remanei is complex, with a loop on the anterior side

of the anterior scar (similar to that of C. stenopora Triebel, 1952, illustrated by

Van Morkhoven (1963: 291)), at least three dorsal scars, a fulchral point, and

an elongate mandibular scar.

Distribution

Cyprideis remanei was recorded by Klie (1940) from the lagoon at the

southern end of Liideritz Bay, where Hartmann (1974: 272) noted the tempera-

ture and salinity as 9.5°C and 35.00%o, respectively.

This is the most abundant of the two species of Cyprideis in our samples,

and occurs throughout the Draaihoek and most of the Gypsum Quarry sections.

In both, it is frequently the dominant ostracod taxon. We recovered one valve

from sample 24 in the Verlorevlei Quarry, but it is absent from Laaiplek and

Velddrif.

Cyprideis draaihoekensis sp. nov.

Figs 9C, 11A-H, 12A-B

Derivation of name

The species is named for Draaihoek farm, locality of type specimens.

Holotype

length height width

MF-1719, RV, DH sample 41 1.00 0.49 —

Paratypes

MF-1720, RV, GQ sample 83 0.95 0.47 os

MF-1721, LV, GQ sample 86 0.91 0.46 -—

MF-1722, LV, GQ sample 86 0.90 0.45 —

MF-1723, RV, GQ sample 86 0.90 0.44 —

MF-1724, C, GQ sample 86 0.91 0.38

MF-1725, C, GQ sample 86 0.90 — 0.39

MF-1726, RV, GQ sample 86 1.00 0.45 —

Diagnosis

Large, ovate, laterally compressed species of the genus Cyprideis, in which

the valve surface ranges from smooth to delicately punctate.

Description

External features. Large, ovate, thin-shelled species, laterally compressed.

Presumed females somewhat inflated posterodorsally. DM broadly arched, VM

almost straight. AM symmetrically rounded, PM slightly asymmetric, truncated

dorsally. LV tend to be slightly more quadrate posterodorsally. There is only a

88 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 11. A-H. Cyprideis draaihoekensis sp. nov. A. Holotype, MF-1719, RV, Draaihoek,

sample 41, SEM AH227. B-H. Paratypes, Gypsum Quarry. B. MF-1720, RV, sample 83,

SEM AH062. C-H. Sample 86. C. MF-1721, LV, SEM AH031. D-E. MF-1722, LV,

internal views. D. SEM 4375. E. MS, SEM 4377. F. MF-1723, RV, internal view,

SEM 4381. G. MF-1722, ATE and PTE, SEM 4379, 4378. H. MF-1723, RV, ATE and

PTE, SEM 4382, 4383. Scale bars: A-D, F = 100 n; E, G-H = S50 p.

OSTRACODA FROM QUATERNARY COASTAL SEQUENCES 89

weak dorsolateral sulcus, and this lies below the indistinct anterodorsal cardinal

angle. NPC small, widely spaced. Valve surface generally smooth, occasionally

finely and faintly punctate.

Internal features. Anterior MA narrow, avestibulate, with 30-40 fine MPC.

There is a prominent selvage. Hinge long with long, curved, denticulate TEs in

RV which taper medianly with only a very short ME groove. MS complex, with

prominent dorsal scars. Adductors lie in a short, curved row. The anterior scar

is small and U-shaped. The most prominent scar is the single, composite man-

dibular scar.

Remarks

Cyprideis draaihoekensis sp. nov. is closest to C. limbocostata, but differs

in being less acuminate posteriorly, and having a less prominent sulcus. It is

easily distinguished from C. remanei, which is plumper overall and more quad-

rate posteriorly. Both species have similar hinge structures, although the TE of

Klie’s species tends to be stronger and in the LV the TE is angled. The anterior

MA of the two species is very similar (Fig. 9B-C), although that of C. remanei

is slightly wider, and has a succession of more prominent RPC (with thickened

proximal portions) interspersed with finer, shorter canals that do not always

extend to the AM.

Hartmann’s (1974) suggestion that the various species of Cyprideis along the

west coast of Africa could be descendants of a common ancestor applies equally

to C. draaihoekensis, which is so far known only from the late Pleistocene-

Holocene.

Distribution

Cyprideis draaihoekensis is much more restricted in its distribution than

C. remanei. It occurs in small numbers (up to 5%) in the upper part of the

Draaihoek section (samples 40-46) and moderate numbers (up to 9%) in the

upper part of the Gypsum Quarry (samples 83-87). It is locally more abundant

at two horizons in the lower part of the Gypsum Quarry (samples 49, 51). We

did not find this species at either Verlorevlei or the southern sections (Laaiplek

and Velddrif).

Hartmann (1974: 271) recorded the temperature and salinity ranges for his

species C. limbocostata as 17-29.3°C and 40-43 %o, respectively. Tankard

(1976) recorded Cyprideis cf. C. limbocostata from the Verlorevlei Quarry.

Family Hemicytheridae Puri, 1953

Genus Ambostracon Hazel, 1962

Subgenus Ambostracon (Ambostracon) Hazel, 1962

Ambostracon (Ambostracon) levetzovi (Klie, 1940)

Fig. 12C-E

Eucythereis levetzovi Klie, 1940: 419-421, figs 23-29.

Aurila levetzovi (Klie, 1940) Hartmann, 1974: 284, pl. 149 (fig. 7).

Ambostracon (Ambostracon) levetzovi (Klie, 1940) Dingle, 1992: 46, figs 28E-F, 29B, E,

34A-C.

90 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 12. A-B. Cyprideis draaihoekensis sp. nov., Gypsum Quarry, sample 86.

A. MF-1724, C, dorsal view, SEM AH045. B. MF-1725, C, ventral view, SEM AH047.

C-E. Ambostracon (A.) levetzovi (Klie, 1940). C. MF-1727, LV, VMS, sample 9,

SEM AH462. D. MF-1728, RV, VMS, sample 9, SEM AH466. E. MF-1729, RV,

Gypsum Quarry, sample 80, SEM AH210. F-H. Aurila kliei Hartmann, 1974.

F. MF-1730, RV, VMS, sample 9, SEM AH451. G. MF-1731, LV, Gypsum Quarry,

sample 78, SEM AH148. H. MF-1732, LV, internal view, Gypsum Quarry, sample 78,

SEM AH134. Scale bars: all 100 p.

OSTRACODA FROM QUATERNARY COASTAL SEQUENCES 91

Illustrated material

length height

MF-1727, LV, VMS-9 0.80 0.43

MF-1728, RV, VMS-9 0.79 0.42

MF-1729, RV, GQ sample 80 0.78 zB

Distribution

This species has previously been recorded from coastal sites in Liideritz Bay

(Klie 1940; Hartmann 1974), and from a water depth of 18 m in St Helena Bay

(Dingle 1992, TBD 3089). In the present study, Ambostracon (Ambostracon)

levetzovi occurs in small numbers at Verlorevlei (VMS-9), the Gypsum Quarry

(sample 80), Velddrif (sample 11) and the two quarries at Laaiplek (samples 15,

19). Preservation of specimens from Verlorevlei and Velddrif is good, but from

Laaiplek and the Gypsum Quarry the valves are fragmented and cracked (e.g.

see Fig. 12E).

Genus Aurila Pokorny, 1955

Aurila kliei Hartmann, 1974

Figs 12F-H, 13A-B

Hemicythere? sp. Benson & Maddocks, 1964: 27-29, pl. 5 (figs 3-4, 6, 8-9), text-fig. 16.

Aurila kliei Hartmann, 1974: 286-288, pl. 54 (figs 402-411), pl. 55 (figs 412-416), pl. 149

(fig. 10). Dingle, 1993: 98-99, figs 54F, SSA-D, 56A.

Illustrated material

length height

MF-1730, RV, VMS-9 Omi 0.40

MF-1731, LV, GQ sample 78 0.77 0.44

MF-1732, LV, GQ sample 78 0.74 0.46

MF-1733, RV, GQ sample 79 0.75 0.44

Distribution

This species has previously been recorded from coastal sites between

Liideritz and Knysna Lagoon, and nearshore sites (15-160 m water depth)

between Liideritz and the Cape Peninsula. In the present study, Aurila kliei is an

important component of the middle part of the Gypsum Quarry section

(samples 77-80) and the Verlorevlei borehole (VMS-9; samples 7-8), and

occurs in small numbers at Velddrif (samples 11, 14) and Laaiplek (sample 15).

Aurila dayii Benson & Maddocks, 1964

Fig. 13C-F

Aurila dayii Benson & Maddocks, 1964: 31-32, pl. 5 (figs 10-12), text-fig. 19. Hartmann,

1974: 282-284, pls 51-53 (figs 385-401), pl. 150 (fig. 2).

92 ANNALS OF THE SOUTH AFRICAN MUSEUM

Illustrated material

length height

MF-1734, LV, GQ sample 79 0.70 0.44

MF-1735, RV, GQ sample 78 0.70 0.40

MF-1736, RV, GQ sample 85 0.63 O37,

MF-1737, LV, GQ sample 85 0.60 0.38

Distribution

This species has previously been recorded from Leisure Island in Knysna

Lagoon (Benson & Maddocks 1964), and from coastal sites between Cacuaco

and Mocamedes in Angola (Hartmann 1974). Dingle (1992, 1993) did not

record Aurila dayii from any of the continental shelf nearshore sites, so we can

be confident that this species inhabits only lagoonal and estuarine habitats. In the

present study, A. dayii was found in small numbers in the Verlorevlei Quarry

(sample 24), but as an important component (< 1-38%, mean: 14%) of the

fauna throughout the upper part of the Gypsum Quarry section (samples 77-86),

and as a minor element in one sample in the lower part of the same section

(sample 51). Tankard (1976) listed A. dayii from Verlorevlei, Velddrif and

Churchhaven.

Genus Caudites Coryell & Fields, 1937

Caudites tankardi sp. nov.

Figs 13G-H, 14A-E

Caudites sp. 3329 Dingle, 1993: 122, fig. 67F

Derivation of name

This species is named for Dr A. J. Tankard, formerly of the South African

Museum, for his pioneering studies on the Quaternary sediments of the south-

western Cape.

Holotype

length height width

MF-1738, RV, DH sample 37 0.55 0.29 —

Paratype

MF-0717, C, TBD 2224, 58 m 059 0.29 0.18

Diagnosis

Species of Caudites with narrow, prominent ridge sub-parallel to AM.

Description

External features. Valve sub-triangular in lateral outline, with an asymmet-

ric, drawn-out AM. Anterodorsally, the outline straight. DM strongly arched,

OSTRACODA FROM QUATERNARY COASTAL SEQUENCES 93

Fig. 13. A-B. Aurila kliei Hartmann, 1974, MF-1733, internal views, Gypsum Quarry,

sample 79. A. SEM AH138. B. MS, SEM AH141. C-F. Aurila dayii Benson &

Maddocks, 1964, Gypsum Quarry. C. MF-1734, LV, sample 79, SEM AH128.

D. MF-1735, RV, sample 78, SEM AH117. E. MF-1736, RV, internal view, sample 85,

SEM 4391. F. MF-1737, LV, internal view, sample 85, SEM 4394. G-H. Caudites

tankardi sp. nov. G. Holotype, MF-1738, RV, Draaihoek, sample 37, SEM 4418.

H. Paratype, MF-0717, C, right view, TBD 2224, SEM 3329.

Scale bars: A, C-H = 100 p; B = 10 wp.

94 ANNALS OF THE SOUTH AFRICAN MUSEUM

VM gently concave. The strongest surface feature is a sharp, narrow ridge that

extends from the anterodorsal margin, sub-parallel to the AM, and is continuous

with a narrow ridge that runs parallel, and very close to, the VM. There is an

indistinct SCT, from which a low curved ridge extends almost to the AM ridge.

Further distinctive features are a short longitudinal ridge close to the VM,

immediately anterior of mid-length, and a step-shaped ridge that runs from the

posterodorsal corner to just below mid-height at the PM caudal process. Overall

the valve surface is smooth, with prominent, widely spaced NPC.

Internal features. Details poorly seen. Anterior MA moderately wide. There

is a prominent ocular sinus. Hinge holamphidont, slightly bowed dorsally. MS

typical of hemicytherids, with the middle anterior scar subdivided.

Remarks

Hartmann (1974) recorded three species of Caudites from around southern

Africa (C. knysnaensis—Knysna, C. dacunhai—northern Mozambique, and

C. algicola—Natal to Mozambique), but none of these appear to be close to our

species. Tankard (1976) included C. knysnaensis in his species list from Veld-

drif. As these specimens cannot be located, we have been unable to verify their

identity. However, we suspect that they are in reality our new species C. tank-

ardi, because negatives of material photographed by Tankard and now in the

Rhodes University SEM collection, are only of C. tankardi. It is likely, there-

fore, that C. knysnaensis does not extend as far west as the south-western Cape.

The closest species to C. tankardi is C. africana Omatsola, 1972, which is

the only species of the genus recorded from Nigeria. The two species have a

very similar lateral outline and overall ornamentation, but differ in the latter

lacking the prominent AM ridge and the curved ridge that extends anteriorly

from the SCT. Caudites tankardi has more prominent NPC. The two species

can also be distinguished by differences in MS patterns (compare Fig. 14E with

Omatsola 1972, pl. 31 (fig. 6)).

Distribution

This species has previously been recorded from a nearshore site

(TBD 2224, 58 m) off Saldanha Bay (Dingle 1993). In the present study, a

single valve was recovered from the lowermost level of the section at Draaihoek

(sample 37). For reasons given above, we suspect that Tankard’s (1976) record

of Caudites knysnaensis Hartmann, 1974, from outcrop 9 at Velddrif, is in fact

C. tankardi sp. nov.

Family Xestoleberididae Sars, 1928

Genus Xestoleberis Sars, 1866

Xestoleberis capensis Miller, 1908

Figs 14F, 15A-F

Xestoleberis capensis Miller, 1908: 127-128; 1912: 300. Stebbing, 1910: 505. Benson &

Maddocks, 1964: 26-27, pl. 2 (fig. 12), text-fig. 15. Dingle, 1993: 144-145,

fig. 81E-F.

Xestoleberis ramosa Miller, 1908. Hartmann, 1974 (part.—Knysna specimens only).

OSTRACODA FROM QUATERNARY COASTAL SEQUENCES 95

Fig. 14. A-E. Caudites tankardi sp. nov. A-B. Paratype, MF-0717, C, TBD 2224.

A. Left view, SEM 4467. B. Dorsal view, SEM 4477. C-E. Holotype, MF-1738, RV,

internal views, Draaihoek, sample 37. C. SEM 4471. D. ATE and PTE, SEM 4473, 4472.

E. MS, SEM 4479. F. Xestoleberis capensis Miller, 1908, MF-1739, LV, Gypsum Quarry,

sample 78, SEM AH198. Scale bars: A-C, F = 100 n; D = 50 y; E= 10un.

96 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 15. A-F. Xestoleberis capensis Miller, 1908. A-B. Gypsum Quarry, sample 85.

A. MF-1742, RV, internal view, SEM AH184. B. MF-1743, LV, SEM 4398.

C. MF-1744, C, dorsal view, Verlorevlei Quarry, sample 24, SEM AH398. D. MF-1743,

LV, MS, Gypsum Quarry, sample 85, SEM 4399. E. MF-1740, RV, Gypsum Quarry,

sample 78, SEM AH196. F. MF-1741, LV, Verlorevlei Quarry, sample 24, SEM AH395.

Scale bars: A-C, E-F = 100 yn; D = 10 x.

OSTRACODA FROM QUATERNARY COASTAL SEQUENCES 97

Illustrated material

length height width

MF-1739, LV, GQ sample 78 0.57 O35 —

MF-1740, RV, GQ sample 78 0.54 0.30 —

MF-1741, LV, VQ sample 24 0.55 0.30 —

MF-1742, RV, GQ sample 85 0.49 0.30 —

MF-1743, LV, GQ sample 85 0.49 0.31 —

MF-1744, C, VQ sample 24 0.59 a 0.33

Remarks

We have distinguished this species from the similar taxon Xestoleberis

ramosa Miller, 1908, on the basis of its more prominently arched DM,

straighter VM, and more acutely rounded AM. Our material allows good illus-

tration of the hinge and MS of X. capensis for the first time.

Distribution

This species has previously been recorded from Knysna Lagoon (Benson &

Maddocks 1964; Hartmann 1974), False Bay (Miller 1908), and two inshore

sites west of the Cape Peninsula (15-90 m) (Dingle 1993). In the present study,

X. capensis is most abundant in the Gypsum Quarry, where it occurs throughout

the sequence, particularly in the middle section (samples 51, 77-80, 84-86). It

is also of moderate importance at one level in the Verlorevlei Quarry

(sample 24). Tankard (1976) included the species in his list from Verlore-

vlei Quarry, Laaiplek, Kruispad (sample 12), Saldanha (sample 18) and

Churchhaven.

RESULTS AND DISCUSSION

A total of 17 species of Ostracoda was recovered from 88 samples collected

at 10 sites between Langebaan Lagoon and the Olifants River mouth. Table 1

shows the distribution and abundance of the taxa.

ENVIRONMENTAL INDICATORS

Because the majority of species have been described previously and some

information on their modern habitats is available, estimates can be made for the

environmental preferences of the bulk of the fauna, and the tolerances of the

four new species can be assessed from their association with the known taxa.

These habitat preferences are presented in Table 2, where we can group the

species into five broad categories of habitat: marine, normal salinity (< 35%o),

inner shelf; marine, normal salinity, lagoonal; marine, high salinity (> 35%o),

lagoonal; estuarine, hyposaline (< 30%o); and freshwater.

To test these empirically derived categories against quantitative data, we

performed a Q-mode factor analysis on the species distribution and abundance

matrix using a version of Oregon State University’s CLIMAP/CABFAC program

(Imbrie & Kipp 1971). The technique has been described and applied previously

to ostracod faunas from the west-coast continental margin by Dingle &

ANNALS OF THE SOUTH AFRICAN MUSEUM

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102

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OSTRACODA FROM QUATERNARY COASTAL SEQUENCES

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100 ANNALS OF THE SOUTH AFRICAN MUSEUM

Giraudeau (1993). The varimax factor component and varimax factor score

matrices that resulted from one rotation of the original matrix are presented in

Appendices 1 and 2, respectively. We elected to analyse for six factors, and in

Table 3 list the species that comprise the factor associations (FA). The six

factors account for 79.5 per cent of the variance of the data set but, as can be

seen from the cumulative variance in Appendix 2, a mere 2 per cent increase

was achieved by adding the sixth factor, and the addition of several more factors

to account for, perhaps 90 per cent of the variance, would not be helpful for the

analysis. The reason for this can be seen in the low communality values

(< 0.800) for the minority of sites from which only isolated samples contained

ostracods (usually in low numbers, and representing environments with no ana-

logues in the richer assemblages). Seventy-eight per cent of the samples have a

communality > 0.800, suggesting that the factor analysis can be used

successfully to help interpret the depositional environments of the samples using

the FA.

Basing estimates for each of the FA on previously described species habitats

(Table 2), the statistical analysis suggests that the faunas in our samples can

TABLE 2

Habitats of south-western Cape Quaternary ostracods.

MARINE (INNER SHELF): NORMAL MARINE

Ambostracon (A.) levetzovi inner shelf (St Helena Bay, 18 m), coastal sites

(Lideritz).

inner—mid shelf (19°S-24°E), coastal sites

(Liideritz—Leisure Island).

inner shelf (Ltideritz—Cape Peninsula), coastal

sites (Liideritz, Kommetjie, Leisure Island).

inner shelf (western Cape Peninsula, False Bay),

coastal sites (Hout Bay, Leisure Island).

inner shelf (off Saldanha Bay, 58 m).

inner shelf (off Olifants River mouth, 72 m).

Garciaella k. knysnaensis

Aurila kliei

Xestoleberis capensis

Caudites tankardi sp. nov.

Cytheromorpha milleri sp. nov.

MARINE (LAGOONAL): NORMAL MARINE

Palmoconcha? cf. P. peterseni

Aurila dayii

Aglaiella railbridgensis

Paranesidea verlorevleiensis sp. nov.

Indeterminate bairdiid

MARINE (LAGOONAL): HIGH SALINITY

Cyprideis remanei

Cyprideis draaihoekensis sp. nov.

ESTUARINE: BRACKISH (HYPOSALINE)

Paracypris westfordensis

VLEIS: FRESHWATER

Heterocypris capensis

Sarscypridopsis ?reniformis

Sarscypridopsis ?aculeata

coastal sites (Angola).

coastal sites (Angola, Leisure Island)

coastal site (Leisure Island), estuarine site (upper

Knysna Lagoon).

lagoonal (Liideritz).

upper estuary (Knysna).

ponds and vleis, south-western Cape.

OSTRACODA FROM QUATERNARY COASTAL SEQUENCES 101

be described in terms of three normal salinity marine associations (factors 3,

4, 6), two freshwater associations (factors 2, 5), and a high-salinity marine

association (factor 1).

Normal salinity marine environments

The only major difference between the empirical and factor analyses of the

marine taxa is the inclusion by the latter of Aurila dayii in the same factor

association (3) as A. kliei and Xestoleberis capensis. Aurila dayii has been

reported previously only from coastal sites, whereas the other two species are

known to occur on the inner shelf (in addition to coastal sites). On this

evidence, we are inclined to interpret FA 3 as indicating coastal marine, in

contrast to inner-shelf environments. This would accommodate Palmoconcha cf.

P. peterseni, which has been recorded previously only from coastal sites

(although its inclusion in FA 3 is based on a very low score, which is statisti-

cally of no significance). We have insufficient evidence from modern

assemblages to differentiate the environments identified by FA 4 and 6, except

that we suspect that they are both indicative of the innermost shelf. It should be

pointed out that little confidence can be placed on the inclusion within them of

the species linked by ‘weak’ factor scores. It is possible that if a larger number

of factors had been specified in the analysis, then some at least would have

fallen into different categories. This is particularly the case with Aglaiella

railbridgensis, which has previously been reported only from two environments

in Knysna Lagoon (range, 30-35 %o). The latter may be an allochthonous

element in an otherwise inner-shelf association. The two bairdiid categories

(Paranesidea verlorevleiensis and the indeterminate bairdiid) are effectively

linked to the other taxa only via one sampling site, so here we may be dealing

with an environment that has no analogues in the rest of the data set.

High-salinity marine environment

In our empirical grouping, the two species of Cyprideis were placed

together and the factor analysis also weakly links them. However, the factor

analysis also suggested that Paracypris westfordensis has its strongest link

(albeit weak) with Cyprideis remanei. On the evidence of previous reports on

the habitats of these two taxa, such a linkage seems unlikely in an autochthonous

assemblage, with Paracypris westfordensis having only been found in

hyposaline environments, whereas Cyprideis remanei occurs in high-salinity

coastal sites. The most likely explanation is that the assemblages we have

analysed are mixed populations, i.e. FA 1 represents a high-salinity marine

setting into which hyposaline elements have been washed.

Freshwater environments

The three species listed under this heading in Table 2 have been recorded

previously only from freshwater sites in the south-western Cape, and worldwide

exceptionally in very low-salinity conditions (Sarscypridopsis aculeata, 11.2 %o

in Australia—De Deckker 1981, 1983). The factor analysis identified two

freshwater FA (2, 5), differentiating the combination of S. aculeata and

Heterocypris capensis from S. reniformis.

102 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 3

Factor Associations, factor scores and variance of Quaternary ostracod faunas, south-western Cape.

HIGH SCORES LOW SCORES

MARINE: NORMAL SALINITY

COASTAL

Factor 3 (12%)

Xestoleberis capensis 0.634 Palmoconcha? cf. P. peterseni 0.006

Aurila kliei 0.624

Aurila dayii 0.425

?INNER SHELF

Factor 4 (8%)

Cytheromorpha milleri 0.999 Caudites tankardi 0.003

?INNER SHELF (?WITH ALLOCHTHONOUS LAGOONAL ELEMENTS)

Factor 6 (2%)

Ambostracon levetzovi 0.617 Garciaella k. knysnaensis -0.297

Paranesidea verlorevleiensis 0.185

Indeterminate bairdiid 0.069

Aglaiella railbridgensis 0.017

MARINE: HIGH SALINITY

LAGOONAL (?WITH ALLOCHTHONOUS HYPOSALINE ELEMENTS*)

Factor 1 (25%)

Cyprideis remanei 0.987 *Paracypris westfordensis 0.133

Cyprideis draaihoekensis 0.075

FRESHWATER

Factor 2 (16%)

Sarscypridopsis ?aculeata 0.907

Heterocypris capensis 0.358

Factor 5 (14%)

Sarscypridopsis ?reniformis 0.948

PALAEOENVIRONMENTS

Applying the empirical and statistical estimates to individual samples allows

us to suggest depositional environments for the various sites, and to present an

analysis of evolving palaeoenvironments for the Draaihoek and Gypsum Quarry

sequences.

Draaihoek and Gypsum Quarry (Figs 16-20)

‘The ostracod faunas at Draaihoek indicate two cycles of salinity change. At

the base, the fauna is dominated by Cytheromorpha milleri, indicating a normal

Factor scores

Percentage total fauna

OSTRACODA FROM QUATERNARY COASTAL SEQUENCES

DRAAIHOEK

Peay

[4]

0.84—

—

0.6

0.4

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48 47 46 45 44 43 42 41 40 39 34 35 36 JP 37

Samples

fee Base

++ F1: Cyprideis remaneilParacypris westfordensis/Cyprideis draaihoekensis

>< F2: Sarscypridopsis ?aculeata/Heterocypris capensis

HE F3: Xestoleberis capensis/Aurila klieilAurila dayii

—t— F4: Cytheromorpha milleri

i F5: Sarscypridopsis ?reniformis

>< F6: Ambostracon (Ambostracon) levetzovi A

100—

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Samples

Top Base

+ Cyprideis remaneilParacypris westfordensis/Cyprideis draaihoekensis

> Sarscypridopsis ?aculeata/Heterocypris capensis

Sarscypridopsis ?reniformis

103

Fig. 16. Ostracod faunas and palaeo-salinity interpretations in the Draaihoek sequence. The

significance of the faunal changes between samples 47 and 48 (top of sequence) is unclear

because of low valve numbers. In the top panel, M:h = marine, high salinity. A. Varimax

factor scores for samples. See Appendix 1 for a listing of factor scores and communalities.

B. Variations in percentages of the high-salinity marine and freshwater ostracod species.

See Table 1 for data.

104 ANNALS OF THE SOUTH AFRICAN MUSEUM

salinity, coastal marine environment (FA 4). This passes upward via a

high-salinity marine fauna (Cyprideis remanei with allochthonous Paracypris

westfordensis, FA 1) into a freshwater assemblage with Sarscypridopsis reni-

formis and Heterocypris capensis (FA 5 and FA 2, respectively). Immediately

above (in sample 40), the fauna indicates a reversion to more marine conditions

with a flood of Cyprideis remanei, rare C. draaihoekensis and abundant,

presumed allochthonous, Paracypris westfordensis (FA 1).

Overlying sediments indicate a second transition to a freshwater environ-

ment. Sample 41 has a mixture of high-salinity marine and freshwater faunas

(FA 1, 2, 5), suggesting deposition in a vlei subjected to significant marine infil-

tration, and this passes upwards into 0.5 m of clays and sands with freshwater

assemblages dominated by FA 2 (Heterocypris capensis and Sarscypridopsis

aculeata) throughout, but with FA 5 (8. reniformis) relatively more important in

the lower part. The low number of valves (2) in the uppermost sample

(sample 48) precludes any confident interpretation of the fauna.

The salinity fluctuations are schematically shown on the curve in Figure 17.

Ostracod abundances fluctuate considerably throughout these faunal changes

(Fig. 18). Generally we found < 100 valves/10 g, but at two horizons with

freshwater assemblages (samples 39 (S. reniformis) and 43 (Heterocypris

capensis), > 200 valves/10 g), and in sample 40 with a high-salinity marine

fauna (Cyprideis remanei, > 500 valves/10 g), there are sudden increases in the

numbers of a particular species. Only in sample 41 (mixed marine and fresh-

water) did we recover > 100 valves/10 g, with no domination by one species.

OSTRACOD

ASSEMBLAGES

GYPSUM QUARRY DRAAIHOEK

~ periodic marine flooding,

evaporation to dryness

and freshwater input

Freshwater

Marine

Normal

——-> Time

Fig. 17. Schematic representation of cyclical salinity changes at Draaihoek (solid line)

and the Gypsum Quarry (dashed line). Dots indicate presence of samples with a

particular ostracod fauna indicative of the suggested palaeoenvironment. The degree of

contemporaneity of the two sequences is unknown.

In contrast, the ostracod faunas from the Gypsum Quarry (Fig. 19) indicate

only a partial cycle of salinity change from normal marine at the base

(samples 77-80), through sediments with a high-salinity marine fauna

(samples 82-84), via a mixed assemblage (sample 85), to freshwater sands and

OSTRACODA FROM QUATERNARY COASTAL SEQUENCES 105

gypsiferous beds at the top. We have indicated these changes in Figure 17, but

can only speculate that they were contemporaneous with the same part of the

cycle at Draaihoek. Ostracod abundances (Fig. 20) are generally lower than

found at Draaihoek (< 50 valves/10 g), with the notable exception of the fresh-

water sands immediately below the gypsiferous beds (500 valves/10 g), where

there is a marked influx in numbers of Sarscypridopsis reniformis Kens abun-

dant Cyprideis remanei and C. draaihoekensis).

DRAAIHOEK |" Emcee] (ore) Gem) (mete mama

600

300+ all

Number valves per 10 g

T T al

43 42 41 40 39 34 35 36 JP 37

Samples Base

Fig. 18. Abundance of ostracods (expressed as number of valves per 10 g of

> 250 p residue) per sediment sample at Draaihoek.

In detail there are significant differences between the successions at Draai-

hoek and the Gypsum Quarry.

Firstly, the normal salinity marine faunas are not the same. At the beginning

of the first cycle at Draaihoek, these are dominated by FA 4 (Cytheromorpha

milleri), whereas at the Gypsum Quarry they are dominated by the more diver-

sified FA 3 (Xestoleberis capensis, Aurila kliei, A. dayii), which we suspect

may represent a less open-water, more coastal assemblage.

Secondly, Paracypris westfordensis comprises a larger component of FA 1

at Draaihoek, compared to that at the Gypsum Quarry, suggesting that at the

former site there was a more persistent inflow of brackish water (which intro-

duced allochthonous valves of this taxon) into the high-salinity marine lagoon

with its autochthonous Cyprideis remanei and C. draaihoekensis assemblage.

Finally, although the freshwater component of the ostracod assemblages

throughout the succession at the Gypsum Quarry is almost exclusively Sars-

cypridopsis reniformis (FA 5), and Heterocypris capensis does not occur at this

106 ANNALS OF THE SOUTH AFRICAN MUSEUM

) — = GYPSUM

0.8

0. .) as

—

fo}

77)

6 «(04

8 q

: |

0.2 =

0 —

Se alr aie T T Te Taal aso i ea |

87 86 85 84 83 82 80 79 78 ta

Samples

++ F1: Cyprideis remanei/Paracypris westtordensis/Cyprideis draaihoekensis

>< F2: Sarscypridopsis ?aculeata

+H- F3: Xestoleberis capensis/Aurila kliei/Aurila dayit

—t— F4: Cytheromorpha milleri

ta F5: Sarscypridopsis ?reniformis A

>< F6: Ambostracon (Ambostracon) levetzovi

Percentage total fauna

Samples

To oy ns - E Base

P += Cyprideis remanei|Paracypris westfordensis/Cyprideis draaihoekensis

>< Sarscypridopsis ?aculeata B

th Sarscypridopsis ?reniformis

Fig. 19. Ostracod faunas and palaeo-salinity interpretations in the Gypsum Quarry (1—North

end) sequence. A. Varimax factor scores for samples. See Appendix 1 for a listing of factor

scores and communalities. B. Variations in percentages of the high-salinity marine and

freshwater ostracod species. See Table 1 for data.

OSTRACODA FROM QUATERNARY COASTAL SEQUENCES 107

locality, there is more variation at Draaihoek. At the end of the first cycle at

Draaihoek, FA 5 (Sarscypridopsis reniformis) dominates but, during the course

of the second cycle, FA 2 (mixed Heterocypris capensis and Sarscypridopsis

aculeata) becomes progressively more important.

All the samples show a high communality in the factor components matrix,

except sample 43 at Draaihoek. This has a particularly high percentage of

Heterocypris capensis, in conjunction with Cyprideis remanei, suggesting that

the former may be a largely allochthonous element.

=) re

Freshwater

600——

Number valves per 10 g

ee ee oe ie

83 82 80 79 78 ale

| Top Samples Base

Fig. 20. Abundance of ostracods (expressed as number of valves per 10 g of

> 250 pu residue) per sediment sample at Gypsum Quarry (1—North end).

In his summary of the distribution of ostracods in Australian lakes, De

Deckker (1983) noted that Sarscypridopsis aculeata and Heterocypris spp. are

confined to temporary pools, which can occasionally attain a salinity of up to

5%o (although he also noted that the former can ‘withstand slightly saline waters

up to 20%o’ (De Deckker 1982: 260)). A totally different halobiotant fauna

occurs in salt lakes, with salinities of up to c. 180%0 (De Deckker 1983,

figs 1-3). Tankard (1976) estimated that salinities must have exceeded 117%o

during the precipitation of the gypsiferous layers at the top of the Gypsum

Quarry section, and that the most likely mechanism was the periodic flooding by

the sea of a back barrier depression, which then evaporated to dryness.

He described the gypsiferous sediment as typically consisting of 2-mm thick

calcium carbonate-calcium sulphate-halite cycles. It is not clear from his

descriptions which layers at the Gypsum Quarry contained the ostracods listed

in his table 2 (1976), but in our examination we recovered only three species

from within the gypsiferous sequence (sample 87): Sarscypridopsis reniformis,

108 ANNALS OF THE SOUTH AFRICAN MUSEUM

Cyprideis remanei and C. draaihoekensis. There is no evidence in the literature

that any of these species can withstand hypersaline conditions and, since no

specialized high-salinity species were found (e.g. similar to those described

from Australia by De Deckker 1983), we conclude that the assemblage

represents a freshwater fauna that re-established itself after the lagoon had evap-

orated to dryness, and mixed with a marine component brought in during the

next invasion of the lagoon by the sea. In the sample immediately below the

gypsiferous layer (sample 86), there is a similar faunal combination, only here

the freshwater ostracod element is more important, suggesting that during this

earlier phase, the inflow of fresh water to the lagoon was on a more substantial

scale, and that marine incursions were less frequent. An alternative explanation

is that the fauna in the gypsiferous layer represents contamination from the

immediately underlying horizon (either naturally as reworking, or during

sampling).

A more comprehensive palaeoenvironmental interpretation will be attempted

in a later publication, when sedimentological evidence will also be considered.

Other sites

From the other sites, we have data only from isolated horizons, and in the

factor analysis communalities for these samples are often low, suggesting that

the faunas do not have strong analogues within the data set as a whole.

Olifants River mouth. The fauna in sample 64 has a high communality and

falls within FA 1, suggesting a high-salinity marine habitat (Cyprideis remaneéi),

with allochthonous hyposaline elements.

Velddrif. The two samples at this site indicate a lower, normal salinity

coastal marine fauna (sample 11, FA 3) and an upper freshwater fauna

(sample 14, FA 2). Both faunas occur in sedimentologically similar samples, but

the communality of sample 11 is low, containing only two valves of marine taxa

Ambostracon levetzovi and Aurila kliei, which the factor analysis as a whole

places in separate categories. The upper fauna, on the other hand, has a high

communality, is relatively abundant, and the thin valves are well preserved.

Clearly, the freshwater fauna is either autochthonous (which does not accord

with the shelly sand matrix of the sample), or it is a short-travelled allochthon-

ous assemblage (there was one valve of A. kliei in the sample).

Laaiplek. Faunas are sparse, and communalities low for both ostracod-

bearing samples in the Laaiplek area. The species present suggest normal

salinity, coastal-inner-shelf marine environments (FA 3, 6), with no estuarine

or freshwater elements. The closest analogue amongst the other sampling sites

is sample 11 at Velddrif.

Verlorevlei. With the exception of VMS-9 sample 7, the factor analysis

communality is very low for the ostracod-bearing samples from the Verlorevlei

valley. This is not surprising in the case of the samples from the quarry,

because they contain a high proportion of bairdiid species that have no ana-

logues in the other samples. In fact, the two species present have not been

recorded previously, which makes comparison difficult. Nevertheless,

sample 24 contains a moderately diverse fauna, and of the elements for which

OSTRACODA FROM QUATERNARY COASTAL SEQUENCES 109

we do have previous environmental data, all, bar one valve of Paracypris west-

fordensis, suggest a normal marine, probably lagoonal environment (Table 2:

Xestoleberis capensis has been found at Leisure Island in Knysna Lagoon, even

though it is more usually found on the inner shelf, and the higher salinity Cypri-

deis remanei is restricted to one valve). There are no freshwater indicators.

Sample H1 contained only valves of Paranesidea verlorevleiensis which, by

deduction, is probably a normal marine, lagoonal species.

The two samples from Miller et al.’s (1993) borehole VMS-9 also contain

only marine species (particularly Ambostracon levetzovi and Aurila kliei), but

the fauna has no elements common with the quarry site (VQ 24, H1) farther up

the vlei, and suggests a more open coastal or inner-shelf environment.

ACKNOWLEDGEMENTS

We are grateful to the Foundation for Research Development (FRD) and the

South African Museum for funds to cover field and laboratory work. AH

completed this study while on sabbatical leave at the South African Museum

from the Ministry of Energy & Infrastructure, Israel. We especially thank Linda

Bisset (SAM) for processing all the sediment samples, for assisting with the

SEM work and for making the photographic prints and plates. Various

colleagues assisted with the field work, in particular D. Miller and J. Parkington

(Archaeology Department, UCT), and J. Pether and J. Giraudeau (SAM).

M. Lavelle (in collaboration with R. Armstrong) (Geological Sciences, UCT)

kindly undertook and permitted us to quote the Sr isotope date on the molluscs

from the base of the Gypsum Quarry section. J. Giraudeau helped with the

factor analysis, using a program supplied by Professor H. Schrader (University

of Bergen). We also thank R. Cross, Director of the Electron Microscope Unit,

Rhodes University, for kindly allowing us access to the archive of Tankard’s

SEM photographic negatives.

We gratefully acknowledge the help given by Dr E. Bertels (Buenos Aires)

in connection with our investigation of Rossi de Garcia’s original descriptions of

Bensonia and related taxa.

Finally, we thank Professor A. R. Lord (London), Dr P. de Deckker

(Canberra) and Dr D. Miller (Cape Town) for their helpful reviews of the

manuscript.

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APPENDIX 1

Varimax Factor Components Matrix.

Factors

Sites Samples Communality

D, 3 4 5 6

GYPSUM QUARRY

1—North end 87 0.964 0.537 0.188 0.025 0.009 0.799 —0.011

86 0.960 0.396 0.209 0.030 0.006 0.871 —0.012

85 0.901 0.656 0.080 0.498 0.008 0.368 0.285

84 0.980 0.979 0.033 0.065 0.021 0.117 0.038

83 0.980 0.969 0.051 0.021 0.020 0.191 0.008

82 0.968 0.892 0.097 0.055 0.017 0.400 0.020

80 0.867 —0.003 0.126 0.886 0.160 —0.046 0.196

79 0.891 0.008 —0.028 0.942 —0.014 0.000 0.054

78 0.943 0.115 0.002 0.957 —0.010 0.103 —0.056

77 0.904 0.330 0.057 0.811 0.005 0.326 0.167

2—South end 51 0.963 0.733 0.159 0.031 0.013 0.632 —0.001

50 0.976 0.987 0.013 —0.001 0.021 0.016 0.002

49 0.924 0.960 0.014 0.001 0.017 0.036 0.004

DRAAIHOEK 48 0.944 0.696 0.650 0.029 0.015 —0.185 0.035

47 0.942 0.022 0.966 0.039 0.002 —0.072 0.034

46 0.988 0.016 0.991 0.031 0.003 —0.062 0.025

45 0.938 0.137 0.804 0.041 0.004 0.520 0.011

44 0.886 0.078 0.856 —0.002 0.007 0.382 —0.025

43 0.623 0.198 0.699 —0.028 0.012 0.304 —0.045

42 0.925 0.095 0.851 0.003 0.008 0.437 —0.024

4] 0.847 0.549 0.459 —0.018 0.017 0.576 —0.045

40 0.990 0.991 0.035 —0.004 0.022 0.078 —0.005

39 0.958 0.004 0.269 0.029 0.000 0.940 —0.018

34 0.986 0.131 0.319 0.009 0.005 0.930 —0.037

35 0.814 0.855 0.130 —0.020 0.124 0.221 —0.029

36 0.999 0.044 —0.003 0.011 0.998 0.001 0.006

JP 0.990 0.077 0.001 0.013 0.996 0.017 —0.005

37 0.999 0.076 —0.003 0.012 0.997 0.002 0.001

VELDDRIF 14 0.858 —0.013 0.890 0.067 —0.001 0.248 0.019

11 0.163 —0.019 —0.004 0.398 —0.012 —0.013 0.061

LAAIPLEK 15 0.337 —0.013 —0.004 0.495 —0.005 —0.024 —0.302

19 0.052 —0.005 —0.005 —0.025 0.008 0.011 0.226

OLIFANTS RIVER 64 0.889 0.943 0.015 —0.014 0.022 0.006 —0.011

VERLOREVLEI

Quarry 24 0.110 0.016 0.003 0.220 —0.010 —0.022 0.247

H1 0.048 —0.007 0.007 0.115 —0.010 —0.025 0.185

Soundings 7 0.716 —0.010 —0.001 0.617 —0.009 —0.036 —0.578

9 0.189 —0.019 —0.004 0.434 —0.012 —0.016 —0.001

VARIANCE 25.999 16.946 12.295 8.184 14.072 2.008

CUMULATIVE VARIANCE 25.999 42.945 55.240 63.423 77.495 79.504

* top of sedimentary succession lies towards the top of list for each site.

114 ANNALS OF THE SOUTH AFRICAN MUSEUM

Species

Ambostracon levetzovi

Caudites tankardi

Garciaella k. knysnaensis

Aurila kliei

Cytheromorpha milleri

Xestoleberis capensis

Palmoconcha? cf. P. peterseni

Indeterminate bairdiid

Paranesidea verlorevieiensis

Aurila dayii

Aglaiella railbridgensis

Paracypris westfordensis

Cyprideis remanei

Cyprideis draaihoekensis

Heterocypris capensis

Sarscypridopsis ?reniformis

Sarscypridopsis ?aculeata

APPENDIX 2

Varimax Factor Score Matrix.

—0.061

0.000

0.026

0.624

0.011

0.634

0.006

0.051

0.115

0.425

0.014

—0.030

—0.001

0.004

—0.061

0.030

0.042

Factors

—0.005

0.003

0.016

-0.012

0.999

0.009

0.000

—0.004

—0.010

—0.018

—0.001

0.007

0.021

—0.008

0.010

—0.001

0.000

0.018

0.000

—0.003

—0.037

0.000

0.014

0.000

—0.008

—0.025

0.020

0.005

0.019

0.016

0.067

0.127

0.948

—0.278

0.617

0.000

—0.297

—0.530

0.005

0.376

0.002

0.069

0.185

0.248

0.017

—0.028

0.002

0.008

—0.069

—0.016

0.047

6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature (particu-

larly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be followed

by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb. nov., syn. nov.,

etc.

An author’s name when cited must follow the name of the taxon without intervening punctuation

and not be abbreviated; if the year is added, a comma must separate author’s name and year. The

author’s name (and date, if cited) must be placed in parentheses if a species or subspecies is trans-

ferred from its original genus. The name of a subsequent user of a scientific name must be separated

from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published scientific

names by which the species previously has been designated are listed in chronological order, with all

references to that name following in chronological order, e.g.:

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).

Nucula largillierti Philippi, 1861: 87.

Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:

comma separates author’s name and year

semicolon separates more than one reference by the same author

full stop separates references by different authors

figures of plates are enclosed in parentheses to distinguish them from text-figures

dash, not comma, separates consecutive numbers.

Synonymy arrangement according to chronology of bibliographic references, whereby the year is

placed in front of each entry, and the synonym repeated in full for each entry, is not acceptable.

In describing new species, one specimen must be designated as the holotype; other specimens

mentioned in the original description are to be designated paratypes; additional material not regarded

as paratypes should be listed separately. The complete data (registration number, depository, descrip-

tion of specimen, locality, collector, date) of the holotype and paratypes must be recorded, e.g.:

Holotype

SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach, Port Eliza-

beth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

Note standard form of writing South African Museum registration numbers and date.

7. SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text

e.g... . the Figure depicting C. namacolus...’: ‘*. . . in C. namacolus (Fig. 10)...’

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded by

initials or full names

e.g. DuToit but A.L. du Toit; Von Huene but F. von Huene

e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should preferably be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a book or

article, such as

‘Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial

capital letter, provided the same generic name is used consecutively. The generic name should

not be abbreviated at the beginning of a sentence or paragraph.

Name of new genus or species is not to be included in the title; it should be included in the abstract,

counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts.