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ANNALS OF THE ANNALE VAN DIE

SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

VOLUME 99 BAND 99

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

VOLUME 99 BAND

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S B:8.B:B ev

tw) \

7 E

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LIQ, at

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THE TRUSTEES OF THE DIE TRUSTEES VAN DIE

SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

CAPE TOWN KAAPSTAD

1990-1991

SET, PRINTED AND BOUND IN THE REPUBLIC OF SOUTH AFRICA

BY THE RUSTICA PRESS (PTY) LTD, NDABENI, CAPE

D2109

LIST OF CONTENTS

Boomer, I. D. see DINGLE, R. V. et al.

Cooper, M. R.

A new genus of Rutitrigoniinae (Bivalvia, Trigoniacea) from the Lower Cretaceous

(Aptian) of Zululand? (PublishediFebnuary 19902) syns) oe eae

DINGLE, R. V., Lorp, A. R. & Boomer, I. D.

Deep-water Quaternary Ostracoda from the continental margin off south-western

Africa (SE Atlantic Ocean). (Published September 1990.)....................

DuRAND, J. F.

A revised description of the skull of Moschorhinus (Therapsida, Therocephalia).

(Rublishe he DintanyelOONs) ie cere ye tate aeie acne iaciae Ve coe setae ate ap Rse eee a

GERNEKE, D. A. see WILSON, M. L. et al.

Gow, C. E.

A tooth-bearing maxilla referable to Lycorhinus angustidens Haughton, 1924

(Dinosauria, Ornithischia). (Published December 1990.).....................

HILxer, N.

Benthic communities and sedimentary facies in the Lower Witteberg Group

(evonianySouthAtnica)s(zubliswed uly: 19903) eee sodas oee ace aoe

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

Cretaceous faunas from Zululand and Natal, South Africa. Hatchericeras Stanton,

1901 (Cephalopoda, Ammonoidea), from the Barremian of Zululand. (Pub-

hishve GRU RIOOU Retry cee ian gerne stem neue bieteh ger bat hacte 2 eas Sag ae tua

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

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

Cretaceous faunas from Zululand and Natal, South Africa. A Koloceras (Cephalo-

poda, Ammonoidea) from the Mzinene Formation (Albian). (Published

| SSL DYRUET EN SED Nes ceehcre cach 6 ecnumioy Se ane a ae atte ean ace ea fre ee nee a

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

Lorp, A. R. see DINGLE, R. V. et al.

McMILLan, I. K.

Foraminifera from the late Pleistocene (latest Eemian to earliest Weichselian) shelly

sands of Cape Town City Centre, South Africa. (Published May 1990.).........

PETHER, J.

A new Austromegabalanus (Cirrepedia, Balanidae) from the Pliocene of Namaqua-

land, Cape Province, South Africa. (Published February 1990.) ...............

VAN RIJSSEN, W. J. J. see WILSON, M. L. et al.

VISSER, J. N. J.

Geography and climatology of the late Carboniferous to Jurassic Karoo Basin in

south-western Gondwana. (Published March 1991.) .....................005-

WILLIAMS, G. C.

The Pennatulacea of southern Africa (Coelenterata, Anthozoa). (Published May

ISIS) 5) Stes eae oe acter ty cada ba ake o's oe COR Ota ee CC

Witson, M. L., VAN RUSSEN, W. J. J. & GERNEKE, D. A.

An investigation of the ‘Coldstream Stone’. (Published May 1990.) ................

Volume 99 is complete in 12 parts.

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245

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367

Mal)

231

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415

187

rae

f .

on od

¢ i

1S | aiey

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ey,

NEW GENERIC NAMES PROPOSED IN THIS VOLUME

Aulniriconial Cooper, 1990 i eee ene

Rugocythereis Dingle, Lord & Boomer, 1990

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

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

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

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(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 99 Band

February 1990 Februarie

Part 1 Deel

és

A NEW AUSTROMEGABALANUS

(CIRREPEDIA, BALANIDAE)

FROM THE PLIOCENE OF NAMAQUALAND,

CAPE PROVINCE, SOUTH AFRICA

JOHN PETHER

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

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Kopieregnavrae aan die Suid-Afrikaanse Museum

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

A NEW AUSTROMEGABALANUS (CIRREPEDIA, BALANIDAE)

FROM THE PLIOCENE OF NAMAQUALAND,

CARE PROVINCE, SOUITE APRICA

JOHN PETHER

Department of Cenozoic Palaeontology, South African Museum, Cape Town

(With 3 figures and 1 table)

[MS accepted 14 November 1988]

ABSTRACT

A preliminary investigation of fossil barnacles from shallow marine coastal plain deposits

exposed at Hondeklip, Namaqualand coast, South Africa, revealed the presence of a new species

of austral megabalanine, Austromegabalanus (Notomegabalanus) kensleyi sp. nov., in an Upper

Pliocene formation informally termed the 50-metre Package. This is the first extinct barnacle

described from South Africa and, with the possibility of additional new species being recognized

in future work, the biostratigraphic utility of barnacle assemblages for nearshore deposits of the

South African margin is promising.

CONTENTS

PAGE

HERI ROCIO TON oak ws (eurrsai aes ON NEL aleoN et ace Aree OneCare tt, a aes Sins oe 1

i DESSTIOUMIOMN, & aso. © ain een ala note cigt RC ok feat aie nace ae eee Ta mare Screaes Z

| DISCUSSIOMs. J'5 h 0.5\6 6:5 ons iso.6 aro enon Ee oe area RRS otro c/a centr ae ere 7

GEOlOPICAESE UM Eee Err ett aI ioc Mee a Aare gens ercrmee eset Siete snc i cee 10

| PRS OSTROW TNE I Bere ckoio HIE eee Cet, ceo oats Sua rons Cena cera il

Conichucimesne nails wien Oe ee ao eee ne a we eed LS Nau arene eee 11

ANCHO ECUSMNETUS: ag eee Coats emery guts oA Sia aS Ie org ae ae & 12

IN@UEREMCGOS ‘ose, Sosa 9. been MOTE cee Ree asec a rch CTE tho cid herb Gopeacein is 1

INTRODUCTION

Shallow marine deposits, ranging in age from Early Pliocene to Late

Pleistocene, underlie the terrestrial cover sands of the coastal plain along the

Atlantic margin of South Africa. Mining of diamonds in these marine deposits

adjacent to the Namaqualand coast creates temporary exposures that provide

Opportunities to examine the sedimentology and sample the fossil content.

Hitherto the fossil molluscs have received attention, the most recent work being

that of Kensley & Pether (1986), which presents a list of taxa and discusses the

composition and biogeographic affinities of the fauna. This paper describes the

first new species of barnacle recognized from the coastal plain marine deposits of

South Africa. The specimens were obtained from the diamond mine at Hondeklip

Bay (Fig. 1). The stratigraphy, sedimentary geometry, depositional environ-

ments, and evidence for the age of the deposits in this area are summarized in

Pether (1986).

Ann. S. Afr. Mus. 99 (1), 1990: 1-13, 3 figs, 1 table.

D. ANNALS OF THE SOUTH AFRICAN MUSEUM

SOUTH

ATLANTIC

Fig. 1. Provenance of Austromegabalanus (Notomegabalanus) kensleyi sp. nov. Broken lines in

left insert show inland (subsurface) limits (transgressive maxima) of main marine formations

underlying the coastal plain. Informal formation labels record elevation of transgressive

maximum, e.g. 50-metre Package. Asterisks indicate localities where the new species was

obtained.

DESCRIPTION

Family Balanidae Leach, 1817

Subfamily Megabalaninae Newman, 1979

Genus Austromegabalanus Newman, 1979

Subgenus Notomegabalanus Newman, 1979

Austromegabalanus (Notomegabalanus) kensleyi sp. nov.

Figs 2-3

Diagnosis

Relatively small, smooth shell with single row of parietal pores and porous

radii; no transverse septa in either. Thick scutum of high relief, no exterior radial

sculpture; small, triangular, deeply excavated, lateral depressor pit, deep

adductor muscle pit, no definite adductor ridge. Tergum not beaked and with

short, broad spur at less than two-thirds its own width from basi-scutal angle;

broad, very shallow spur furrow; no definite depressor crests.

Distribution

Upper Pliocene of the Namaqualand coastal plain, west coast of South

Africa.

AUSTROMEGABALANUS FROM THE PLIOCENE OF SOUTH AFRICA 3

Material

Holotype. SAM—PQ—-HB337a, one complete shell with opercula removed

from cluster on boulder, Hondeklip Zone 12A (30°19'29"S 17°18'21"E).

Paratypes. SAM—PQ-HB337b-—df, 110 complete shells from same cluster as

holotype, many with opercula and bases. SAM—PQ-HB 1261, cluster of approxi-

mately 30 shells on Striostrea margaritacea (Lamarck, 1819) valve, one of which

contained an articulating opercular pair, Hondeklip Zone 4A.

Additional material. SAM—PQ-HB321, unattached cluster of 9 shells with-

out opercula, Hondeklip Zone 12A. SAM—POQ-HB180, disarticulated shells and

scuta, Hondeklip Zone 12A. SAM—POQ-AV984, 4 shells without opercula, Avon-

tuur A, T2/3. SAM—PQ—AV1067, 3 shells without opercula, Avontuur A, T2/3.

Measurements

Holotype. Height 4,2 mm, carino-rostral diameter 11,1 mm, scutum occlu-

dent margin 3,7 mm, tergum scutal margin 2,6 mm.

Largest shell. SAM—PQ-AV984a, height 6,3 mm, diameter 14,7 mm.

Largest scutum. SAM—PQ-HB180, occludent margin 6,2 mm.

Description

Shell relatively small (~1 cm basal diameter). Specimens from smooth, un-

crowded substrate (Fig. 2C) are low-conical with regular and smooth exteriors,

have pentagonal to pointed-ovate, moderately toothed orifices about one-third

carino-rostral diameter, and thin, flat bases. Those from uneven and crowded

substrata (Fig. 3C) are more globulo-conical with relatively larger orifices, rugose

exteriors and have cup-shaped bases (up to | cm deep), often with irregularly

cellular underlayers.

Parietes with single row of open longitudinal pores, ovoid in section near

apex, subrectangular near basis; internally ribbed, ribs usually on lower portion

but may extend almost to sheath, short secondary ribs sometimes on larger speci-

mens; basal denticles conspicuous with correspondingly small tapered ridges on

lower ribs; sheath about half the length of plate, lower margin usually separated

from wall by moderate to very shallow concavity. Many specimens still have a

rose-pink hue colouring the sheath. Darker radial lines on exteriors of parietes

correspond to the underlying septa. Radii inset from parietes, transversely porous

although upper pores may be infilled; sutural edges regularly and densely septate

with denticles on lower edges of septa (Fig. 3D), articulating surfaces on adjacent

parietes with corresponding low, upward denticulations. Alae thin and broad,

crenate sutural edges sometimes visible. In unabraded specimens, such as the

SAM-—PQ-HB337, the radii summits are obliquely truncated and the alae have

steeply oblique summits, usually projecting above the opposite sloping radu. In

more abraded specimens the summits are approximately parallel to the base.

Basis radially porous, with shallow furrows corresponding to the underlying

pores.

4 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 2. Holotype of Austromegabalanus (Notomegabalanus) kensleyi sp. nov., SAM—PQ-

HB337a. A. Interior and exterior views of scutum. B. Interior and exterior views of tergum.

C. Apertural and lateral views of shell. Scales = 1 mm.

AUSTROMEGABALANUS FROM THE PLIOCENE OF SOUTH AFRICA 5

Fig. 3. Austromegabalanus (Notomegabalanus) kensleyi sp. nov. A-B. Large paired scutum

and tergum from paratypes, SAM—PQ-HB1261. Scale = 1mm. C. Shell cluster from same

paratypes, set on Striostrea margaritacea valve. Scale = 5mm. D. Sutural edge of radius

showing austral megabalanine denticulation. Scale = 1 mm.

6 ANNALS OF THE SOUTH AFRICAN MUSEUM

Scutum thick, of high relief; concave exterior with prominent growth ridges,

every alternate one denticulating occludent margin; no radial sculpture; basal

margin convex and about two-thirds length of occludent margin; tergal margin

about three-quarters length of occludent margin and on exterior raised as a low

ridge from apex to basi-tergal angle before being strongly but quite narrowly

reflexed, the ridge enhancing the concavity of the exterior; a shallow groove is

often present along the reflexed tergal segment. Articular ridge high, about

four-fifths of tergal margin in length and reflexed over relatively narrow articular

furrow; articular ridge thins basally, ending in a closely rounded-off corner above

concave terminal edge; outer face shallowly concave and faint grooves are

generally visible. Adductor muscle scar large, ovate and conspicuously deep,

usually being visible without magnification, even in small scuta. No obvious

adductor ridge but a low, rounded swelling is usually present beside adductor

tergal margin and laps on to the basal flank of articular ridge. Lateral depressor

pit small, narrowly triangular and deeply excavated, usually with slightly

overhanging sides. Occludent margin interiorly reflexed, producing marginal

ridge that basally overhangs rostral depressor insertion.

Tergum carinal and basal margins convex, scutal margin straight; exterior

with low growth ridges, more pronounced on scutal portion; no radial sculpture.

Spur furrow broad, open and very shallow to flush; margins slightly incised.

Exterior carinal portion gently convex, exterior scutal portion usually slightly

outwardly deflected. Articular margin quite narrowly reflexed at approximately

right angle, rugose, forming ridge along scutal margin of articular furrow;

articular furrow broad, moderately deep; articular ridge relatively thin,

moderately high and partially overhanging furrow near apex. Apex not beaked,

area of transverse ridges small; interior of valve below apex with several irregular

grooves; carinal depressor crests effectively absent but faint grooving sometimes

visible. Spur short and wide, approximately as long as broad; occupying from

quarter to third of basal margin, situated about half to two-thirds own width from

basi-scutal angle; basal margin segments slope towards spur, basi-scutal margin

concave. Scutal end of spur closely rounded off, carinal corner very widely

rounded off and almost a continuous curve from basal margin to scutal end. On

interior of valve spur is only little raised and has a flat to slightly concave inner

face. On the largest tergum available (Fig. 3B) the spur is seen to be relatively

- more developed than is the case in smaller specimens and the upper scutal margin

is eroded.

Etymology

The species is named for Dr Brian Kensley of the Smithsonian Institution,

Washington, D.C., in recognition of his contributions to the knowledge of the

southern African fossil molluscan fauna.

AUSTROMEGABALANUS FROM THE PLIOCENE OF SOUTH AFRICA 7

DISCUSSION

The porous radii establish the Namaqualand specimens as belonging to a

megabalanine genus. The classification system followed here is that of Newman

(1979), with the modifications proposed by Buckeridge (1983). In the former

work, Balanidae with porous radii were allocated to a new subfamily Megabalani-

nae. Taxa possessing radii with secondary denticles on the lower sides of primary

denticles only were recognized as pan-austral Southern Hemisphere endemics and

assigned to two new genera, Austromegabalanus and Notomegabalanus. The

former genus 1s distinguished by beaked terga and closed spur furrows, the latter

by non-beaked terga and open furrows. However, Buckeridge (1983), in his

monograph on the fossil barnacles of Australia and New Zealand, expressed the

opinion that the differences in tergal morphology, with respect to spur furrow and

beak, between Austromegabalanus and Notomegabalanus are not sufficiently

strong to justify full generic separation. He thus proposed that Notomegabalanus

be regarded as a subgenus of Austromegabalanus. Furthermore, the taxa with

porous outer parietal laminae were removed from Notomegabalanus and placed

in a new genus Fosterella. With secondary denticles on the lower sides of the

primary radii denticles only, this species from Namaqualand is confirmed as an

austral megabalanine. The open spur furrow and lack of a strongly beaked tergum

place it in the subgenus Notomegabalanus.

Listed in Table 1 are the fossil and extant species of Notomegabalanus,

together with their approximate geological ages. Austromegabalanus s.s. and

Fosterella have been included to provide a complete list of these related austral

genera.

The austral megabalanines have not always been confined to the Southern

Hemisphere. Fossil occurrences of Notomegabalanus in Tertiary deposits on both

the Pacific and Atlantic coasts of North America have been recorded by Zullo &

Guruswami-Naidu (1982) and Zullo (1986). As discussed by Newman & Foster

(1987), this distribution suggests that Notomegabalanus has had a Tethyan and an

amphitropical history, the present austral endemism being due to Northern Hemi-

sphere extinction and Neogene dispersal via the West Wind Drift. Notomega-

balanus is considered more generalized than Austromegabalanus s.s., whilst

Megabalanus s.s. is the most highly evolved of the megabalanines (Newman 1979).

The presence of an undescribed species of Notomegabalanus in the Oligocene of

Oregon (Zullo 1986) is consistent with such a relationship. The Neogene radiation

of these related taxa and the extinctions due to Pleistocene glaciations (which

particularly affected Fosterella and Notomegabalanus), are evident in Table 1.

Austromegabalanus (Notomegabalanus) kensleyi is rendered distinct from all

other species of the subgenus by its short, broad, rounded tergal spur at less than

its own width from the basi-scutal angle, especially if this character is considered

simultaneously with the very shallow spur furrow and subdued expression of the

spur ridge on the valve interior.

The comparison of A. (N.) kensleyi with the austral species of Notomegabala-

nus indicates that the deep, narrowly triangular lateral depressor pit distinguishes

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ANNALS OF THE SOUTH AFRICAN MUSEUM

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(£96 ‘UdSIIOAIS 2 SINYIJOH]) apjos1 (snuvjvqvsawousny) snuvjvqvsawousnpy

(O8LT ‘UITOWID) snoupuydd (snuvjoqvsawoysny) snunjvgvsawousny

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(QIQT ‘yorewey) suaosausiu (snuvjoqvsawoysny) snuvjvgvsawousny

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AUSTROMEGABALANUS FROM THE PLIOCENE OF SOUTH AFRICA 9

the scutum of the new species from other representatives of the subgenus. The

lack of a definite adductor ridge separates the new species from all other austral

Notomegebalanus species except A. (N.) algicola (Pilsbry, 1916) which, however,

has a very broad scutum. The absence of scutal radial sculpture is a feature in

common with A. (N.) algicola and A. (N.) campbelli (Filhol, 1885). The basal

end of the articular ridge of the new species is not pointed or dependant, as is the

tendency in other austral species of Notomegabalanus. The lack of a tergal beak

distinguishes the new species from the weakly beaked Notomegabalanus species

of New Zealand.

The shells of A. (N.) kensleyi are slightly larger than those of A. (N.) algicola

but smaller than the New Zealand representatives of the subgenus, the ribbed

A. (N.) campbelli being closest in dimensions. Whereas the latter and

A. (N.) decorus (Darwin, 1854) have transverse septa in the parietal pores, the

new species does not, a feature in common with A. (N.) algicola and evidently

A. (N.) miodecorus Buckeridge, 1983.

In the Northern Hemisphere, the extant Balanus (Megabalanus) squillae

Daniel & Ghosh, 1963, possesses austral megabalanine radii. This rather cryptic

and very small species from the coast of Madras is parasitic on a stomatopod,

Squilla sp. It is problematic in possessing a tergum that is non-beaked but has a

closed spur furrow. On the basis of the overall resemblance of the opercula to

those of A. (N.) algicola, this species has been tentatively placed in Notomega-

balanus by Henry & McLaughlin, 1986. The small broad scutum with an adductor

ridge further distinguishes it from the new species. Austromegabalanus

(N.) wilsoni (Zullo, 1969) from the Upper Pliocene of California also possesses an

unbeaked tergum with a closed spur furrow and similarly has been placed in

Notomegabalanus on the basis of its overall resemblance to A. (N.) algicola.

Additional discoveries of such species that do not strictly conform to the definition

of Notomegabalanus might render a redefinition of the subgenus or erection of

another subgenus advisable.

The scutum of A. (N.) wilsoni resembles that of A. (N.) kensleyi in respect of

the small, deep, lateral depressor pit and lack of an adductor ridge, but differs in

respect of the straight basal margin, radially aligned pits on the exterior of some

Specimens and overall larger size. Austromegabalanus (N.) lepidus Zullo, 1986,

from the Pliocene of the Western North Atlantic is a species of comparable size to

that of A. (N.) kensleyi but the scutum, although also lacking an adductor ridge, is

thin with a shallow lateral depressor pit and occasionally has exterior radial

Striae. In addition to the deep spur furrow, the tergum of A. (N.) lepidus differs

from the new species in the strongly concave scutal margin, presence of distinct

depressor crests, and radially striate exterior scutal portion. The opercula of

A. (?N.) obliquus (Ross, 1964) (Pliocene, Western North Atlantic) have not been

recovered but that species is larger than the new species and has externally ribbed

parietes with deeply inset radii. The opercular valves of A. (?N.) insperatus Zullo

& Guruswami-Naidu, 1982 (Late Miocene, California), are also unknown but the

shell is also larger than A. (N.) kensleyi and has prominent external ribs.

i0 ANNALS OF THE SOUTH AFRICAN MUSEUM

It is noteworthy that of all the representatives of Notomegabalanus, the

opercula of the new species are overall most similar to those of the extinct genus

Fosterella Buckeridge, 1983. This applies particularly to the deep, small, scutal

lateral depressor pit (this feature having been considered a defining character of

Fosterella) and the short tergal spur and shallow spur furrow. In general appearance

the opercula of Fosterella tubulatoides (Newman, 1979) from the Late Pleistocene

of the continental shelf off Tierra del Fuego are closest. However, several other

features remain to distinguish the opercula of species of Fosterella, such as the

presence of weak adductor ridges, pointed basal ends of articular ridges, and

strongly developed tergal depressor crests (the latter not present for F. hennigi

(Newman, 1979) but that species has very distinctive elongate opercula).

Furthermore, the shells of Fosterella are all strongly ribbed and, in addition to the

defining character of multitubate parietes, have transverse septa in the parietal

pores.

GEOLOGICAL SETTING

Austromegabalanus (Notomegabalanus) kensleyi is abundantly represented

in a formation informally called the 50-metre Package (Pether 1986). This

formation is comprised of gravels and fine sands laid down during regressive

shoreline progradation from a transgressive maximum near 50 m above present

sea-level. Depositional environments include the nearshore shelf (locally

preserved only), lower shoreface, upper shoreface, foreshore, and back-barrier

settings. The age of the 50-m Package is not yet unequivocally determined but is

constrained by the presence of a remanie vertebrate assemblage of Mio-Pliocene

aspect at the base (Q. B. Hendey pers. comm.) and the occurrence of Equus near

the top, the latter probably representing stratigraphic leakage from a closely

overlying erosion surface. Furthermore, the highest elevation marine deposits

(90-m Package, Fig. 1) inland of the 50-m Package have been correlated with the

Early Pliocene Varswater Formation at Langebaanweg (Hendey 1981). The 50-m

Package is thus considered to be Middle to Late Pliocene in age.

Austromegabalanus (Notomegabalanus) kensleyi has not been found in

barnacle coquina from the 30-m Package, the regressive wedge overlying the

transgressive truncation of the 50-m Package. Thus it appears to have become

extinct on the Namaqualand coast around the Plio-Pleistocene. The lower limits

of the new species are not yet determined, due to lack of material, but it is

considered quite likely to extend into the Early Pliocene 90-m Package. At least

one other species of barnacle is present in the 50-m Package. This is a

non-megabalanine but opercula permitting determination have not yet been

recovered.

In the barnacle coquina of the subsequent Early Pleistocene 30-m Package,

disassociated opercula indicate the presence of three barnacle species, whereas

parietal fragments indicate that Austromegabalanus, Megabalanus s.s. and

Balanus species occur. The opercular material is too abraded for the purposes of

unequivocal specific determination but one scutum type and one tergum type

AUSTROMEGABALANUS FROM THE PLIOCENE OF SOUTH AFRICA 11

(probably from the same species) appear sufficiently unique to suggest a new

species. Other fragmentary opercula suggest that the Austromegabalanus sp.

indicated by parietes may by Austromegabalanus s.s. It is hoped that continued

fieldwork will produce better-preserved and associated examples of these taxa.

PALAEOENVIRONMENT

The types of A. (N.) kensleyi were found in situ on a boulder resting on the

gneissic bedrock. The palaeoenvironment of the type area, reconstructed from

the overlying sediments, was the inner, shallow portion of a protected

embayment. The boulder was at shallow subtidal palaeodepth (<—5 m) when the

live barnacles were covered by fine sands—probably during or immediately

subsequent to a storm. Disarticulated barnacle fragments are a major component

of coarse, basal lower shoreface deposits in the deeper portion of the bay and,

within the fine sandy portion of the lower shoreface bay infill, the new species 1s

commonly found encrusting Striostrea margaritacea (Lamarck) valves. These

oysters were swept from the lower intertidal to shallow subtidal margin of the bay

during storms and deposited in deeper water. Lagoonal deposits within which

barnacle coquina lenses are present represent the final infilling of the embayment.

These lenses are interpreted as coarser lags accumulated along the very shallow

subtidal bases of tidal creeks within the lagoonal system, with barnacle fragments

likely to have been transported by flood tides from the proximal lagoon and

adjacent coast.

In exposed coastal settings, detached clusters of barnacles are found in shelly

lenses within the storm-deposited lower shoreface facies. Disarticulated frag-

ments are also an important component of the cross-stratified, coarse sandy upper

shoreface facies.

In all the above instances the dominant barnacle is referable to the new

species and its preferred environment was evidently the intertidal to shallow

subtidal of normal marine salinity. The presence of warm-water West African and

southern African east-coast molluscs in the 50-m Package indicates that the sea

temperature was warmer than the present interglacial, at least seasonally.

However, upwelling influences were present, as is attested by the occurrence of

authigenic phosphorite rinds within nearshore shelf and basal lower shoreface

facies. It is likely that changes in faunal structure associated with cooling,

sea-level fall, and altered shelf currents account for the extinction of

A. (N.) kensleyi.

CONCLUDING REMARKS

It is appropriate and not unexpected that the first extinct barnacle to be

described from the coastal plain deposits of South Africa is a species of

Notomegabalanus. Newman (1979) remarked that the austral region must have

been richer in austral megabalanines during the Pliocene than it is today. The

subsequent discovery of Fosterella chathamensis Buckeridge, 1983, from the

i, ANNALS OF THE SOUTH AFRICAN MUSEUM

Pliocene of the Chatham Islands, Austromegabalanus (A.) piscoensis Carriol et

al., 1987, from the Pliocene of Peru, and the present species bears out that

supposition that, with inclusion of the North American fossil species, may now be

amplified to include both hemispheres during the Neogene.

Another species of Austromegabalanus is represented in the Early Pleisto-

cene 30-m Package and may be Austromegabalanus s.s., but the abraded state of

the available opercula preclude unequivocal specific identification at present.

Significantly, material conclusively attributable to the extant South African

Austromegabalanus species has not been recovered from the Upper Pliocene and

Lower Pleistocene deposits. This suggests that A. (A.) cylindricus (Gmelin, 1780)

does not have a range extending from the Pliocene, in contrast to A. (A.) psitt-

acus (Molina, 1782) from South America. However, the former species is well

represented in Last Interglacial beach deposits at Milnerton near Cape Town

(Kensley 1985) and at other Late Pleistocene outcrops in the south-western Cape.

Austromegabalanus (N.) algicola is also present in Last Interglacial deposits at

Milnerton.

This paper represents the results of an initial reconnaissance of fossil

barnacles from the Hondeklip study area. The recognition of a new species of

barnacle from Pliocene coastal plain deposits, with possibilities of others in the

future and as yet undetermined lower limits for extant South African

Austromegabalanus species, holds the promise that barnacles will be of value in

the correlation of the Late Tertiary and Quaternary shallow marine sequences of

the southern African margin. Their calcitic composition favours preservation and,

together with their abundance in shallow marine deposits, barnacle assemblages

should emerge as useful biostratigraphic markers in the local coastal stratigraphy.

ACKNOWLEDGEMENTS

I am indebted to Mr F. Hoffman of Transhex Group Ltd for access to the

Hondeklip mine exposures and the mine manager and geologists for their

hospitality; to Dr Q. B. Hendey of the South African Museum and Dr J. Rogers

of the University of Cape Town, for their advice and support; to Marilyn Pether

for photographic assistance; to the referees for their constructive comments, and

to the CSIR and the South African Museum for financial assistance.

REFERENCES

BUCKERIDGE, J. S. 1983. Fossil barnacles (Cirrepedia: Thoracica) of New Zealand and

Australia. Palaeontological Bulletin. Geological Survey, New Zealand 50: 1-151.

CaRRIOL, R. P., DE Muizon, C. & SECRETAN, S. 1987. Les Crustacés (Cirrepedia et Decapoda)

du Neogéne de la cdte Péruvienne. Annales de paléontologie 73 (3): 137-164.

DanigEL, A. & Guosn, A. 1963. A new cirrepede of the genus Megabalanus from the stomatopod

(Squilla sp.) from Madras. Annals and Magazine of Natural History (13) 6 (68): 477-479.

HeENpEY, Q. B.1981. Geological succession at Langebaanweg, Cape Province, and global events

of the Late Tertiary. South African Journal of Science 77 (1): 33-38.

AUSTROMEGABALANUS FROM THE PLIOCENE OF SOUTH AFRICA 13

Henry, D. P. & McLauGuiin, P. A. 1986. The Recent species of Megabalanus (Cirrepedia:

Balanomorpha) with special emphasis on Balanus tintinnabulum (Linnaeus) sensu lato.

Zoologische verhandelingen 235: 1-19.

KENSLEY, B. 1985. The faunal deposits of a Late Pleistocene raised beach at Milnerton, Cape

Province, South Africa. Annals of the South African Museum 95 (2): 111-112.

KENSLEY, B. & PETHER, J. 1986. Late Tertiary and Early Quaternary fossil mollusca of the

Hondeklip area, Cape Province, South Africa. Annals of the South African Museum 97 (6):

141-225.

Newman, W. A. 1979. On the biogeography of balanomorph barnacles of the Southern Ocean

including new balanid taxa; a subfamily, two genera and three species. In: Proceedings of

the International Symposium on Marine Biogeography and Evolution in the Southern

Hemisphere, Auckland, New Zealand, July 1978. Information series. Department of

Scientific and Industrial Research, New Zealand 137 (1): 279-306.

Newman, W. A. & Foster, B. A. 1987. Southern Hemisphere endemism among the barnacles:

explained in part by extinction of northern members of amphitropical taxa? Bulletin of

Marine Science 41 (2): 361-377.

PETHER, J. 1986. Late Tertiary and Early Quaternary marine deposits of the Namaqualand coast,

Cape Province: new perspectives. South African Journal of Science 82 (9): 464-470.

Ross, A. R. 1964. Cirrepedia from the Yorktown Formation (Miocene) of Virginia. Journal of

Palaeontology 38 (3): 484-491.

ZuLLo, V. A. 1969. Thoracic Cirrepedia of the San Diego Formation, San Diego County,

California. Contribution in Science. Los Angeles County Museum 159: 1-25.

ZULLO, V. A. 1986. Pliocene barnacles (Cirrepedia, Thoracica) from South Carolina, with a key

to Pliocene balanoids of the Eastern United States. South Carolina Geology 29 (2): 1-18.

ZuLLoO, V. A. & Guruswami-Naibu, R. B. 1982. Late Miocene balanid Cirrepedia from the

basal Wilson Ranch beds (‘Merced’ Formation), Sonoma County, Northern California.

Proceedings of the California Academy of Sciences 42 (2): 525-535.

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.,

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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

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:

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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-

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Note standard form of writing South African Museum registration numbers and date.

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Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text

e.8. . 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 ‘sebedth 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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Specific name must not stand alone, but be preceded by the generic name or its abbreviation to initial

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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.

JOHN PETHER

A NEW AUSTROMEGABALANUS

(CIRREPEDIA, BALANIDAE)

_ FROM THE PLIOCENE OF NAMAQUALAND,

CAPE PROVINCE, SOUTH AFRICA

4 |

VOLUME 99 PART 2 FEBRUARY 1990 . ISSN 0303-2515

if the

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

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KouNn, 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

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 99 Band

February 1990 Februarie

Part 2 Deel

CRETACEOUS FAUNAS FROM ZULULAND

AND NATAL, SOUTH AFRICA.

A KOLOCERAS (CEPHALOPODA,

AMMONOIDEA) FROM THE MZINENE

FORMATION (ALBIAN)

HERBERT CHRISTIAN KLINGER

WILLIAM JAMES KENNEDY

Cape Town Kaapstad

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are issued in parts at irregular intervals as material

becomes available

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C465

CRETACEOUS FAUNAS FROM ZULULAND AND NATAL, SOUTH

AFRICA. A KOLOCERAS (CEPHALOPODA, AMMONOIDEA) FROM

THE MZINENE FORMATION (ALBIAN)

HERBERT CHRISTIAN KLINGER

Department of Invertebrate Palaeontology, South African Museum, Cape Town

WILLIAM JAMES KENNEDY

Geological Collections, University Museum, Oxford

(With 2 figures)

[MS accepted 22 September 1989]

ABSTRACT

The monospecific, micromorphic ammonite genus Koloceras (of the subfamily Aconecera-

tinae), thus far only known from the Upper Albian of Estancia La Vega in Santa Cruz Province,

Argentina, is recorded for the first time from the Upper Albian of Zululand.

CONTENTS

PAGE

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IR ENIEIRSTD COG ig Bee aes Ele oe cs CR Le Oe eee Rr yt SIE eo ais eae ZAM

INTRODUCTION

On three separate occasions, micromorph, platyconic ammonites have been

recorded from the Upper Albian of the Southern Hemisphere as Boris-

siakoceras—a genus generally regarded as being of Middle Cenomanian to Turo-

nian age, but which may occur as early as the Lower Cenomanian and persist up

to the Coniacian (see e.g. Kennedy & Klinger 1979: 115-116). These records are:

1. Borissiakoceras sp. Nullo et al., 1981 (pl. 2 (fig. 9a—c)) from Patagonia,

Argentina.

2. Falciferella breadeni Brunnschweiler, 1959 (p. 15, pl. 1 (figs 5-6)) and

Falciferella reymenti Brunnschweiler, 1959 (p. 16, pl. 1 (figs 7-8)), both referred

to Borissiakoceras by Cooper & Kennedy (1979: 297) from South Australia.

3. Borissiakoceras sp. nov. ? aff. reymenti Cooper & Kennedy (1979: 297,

fig. 31H-I) from Porto Amboim, Angola.

Ann. S. Afr. Mus. 99 (2), 1990: 15-21, 2 figs.

16 ANNALS OF THE SOUTH AFRICAN MUSEUM

Cooper & Kennedy (1979: 296-7) regarded the Angolan and Australian

material as bridging the phylogenetic gap in the family Binneyitidae between the

last appearance of the genus Falciferella Casey, 1954 (type species F. milbournei

Casey, 1954: 271, pl. 7 (figs 1-5), text-fig. 3), in the Middle Albian and the first

occurrence of Borissiakoceras in the Middle Cenomanian. Recent data suggest

that this view seems to be too simplistic.

The Australian material was recently (McNamara 1985) referred to a new

genus Naramoceras (type species Falciferella breadeni Brunnschweiler, 1959: 15,

pl. 1 (figs 5—6)) in the subfamily Aconeceratinae. The Patagonian material was

also referred to a new genus (Riccardi et al. 1987: 167), Koloceras (type species

Koloceras talenkanum Riccardi et al., 1987: 169, pl. 14 (figs 1-17b), text-figs 42—

46), also of the subfamily Aconeceratinae.

Both Koloceras and Naramoceras have distinctive, little-incised suture lines

with a trifid lateral lobe (L), which clearly separate them from Borissiakoceras.

Koloceras in turn differs from Naramoceras in having a trifid second lateral (U,)

lobe, rather than bifid as in the latter.

The erection of these two (apparently endemic) genera casts doubt on the

identity of the Angolan specimen. Is it indeed the only true Upper Albian

Borissiakoceras, or might it belong to Koloceras or Naramoceras? Unfortunately,

we have not seen the suture lines of the Angolan specimen to be able to comment

further on this matter.

In Patagonia and South Australia, Koloceras and Naramoceras respectively

occur in association with a diverse and distinct heteromorph ammonite fauna

belonging to the subfamily Labeceratinae (see e.g. Whitehouse 1926; McNamara

1978; Aguirre Urreta & Riccardi 1988). The subfamily Labeceratinae has a typi-

cally South Gondwanid distribution, and is locally very common in Zululand and

southern Mozambique (see Spath 1925; Klinger 1976; Klinger 1989). When

Naramoceras and Koloceras were first described, our immediate reaction was to

see if we had any representatives of either genus in our collections from

Labeceras-bearing localities. We did not find any.

Recently, in the course of routine curation of the Van Hoepen collection,

housed in the South African Museum, a small ammonite was brought to the

attention of one of us (H. C. Klinger), who identified it as belonging to the genus

Koloceras; later a second specimen was found, which is tentatively referred to that

genus.

We here describe the first record of the genus Koloceras from South

Africa. We cannot be certain whether this is only a fortuitous discovery of a

post-mortem drifted occurrence of this apparently endemic, monospecific

Patagonian genus, or whether it indeed represents part of a population as yet not

localized. Its discovery should encourage the search for further material of

both Koloceras and Naramoceras in the Southern Hemisphere, to clarify

the interrelationship between these two genera and Borissiakoceras of the

subfamily Binneyitinae.

CRETACEOUS FAUNAS FROM SOUTH AFRICA NY

SYSTEMATIC DESCRIPTION

Superfamily HAPLOCERATACEAE Zittel, 1884

Family Oppelidae Bonarelli, 1894

Subfamily Aconeceratinae Spath, 1923

Genus Koloceras Riccardi, Aguirre Urreta & Medina, 1987

Koloceras talenkanum Riccardi, Aguirre Urreta & Medina, 1987

Figs 1-2

Borissiakoceras sp. Nullo et al., 1981: 200, pl. 2 (fig. 9a—c).

?Falciferella sp. Medina & Rinaldi, 1986: 16, pl. 2 (figs 3-5), pl. 4 (figs 6-9).

Koloceras talenkanum Riccardi et al., 1987: 169, pl. 14 (figs 1-17b), text-figs 42—46.

Type

Holotype is the specimen (MLP 19981) figured by Riccardi et al. (1987, pl. 14

(fig. 15a—b)), an adult macroconch from the upper part of the Rio Mayer For-

mation, Estancia La Vega, Santa Cruz Province, Argentina, HOMIES in the col-

lections of the Museo de Ciencias Naturales, La Plata.

Material

NMB D380 (Van Hoepen Collection) from locality 51 and NMB D872 (Van

Hoepen Collection) from locality 55, both from the Mzinene Formation,

Albian IV—V, Zululand. (For precise locality data see Kennedy & Klinger 1975.)

Dimensions

Dimensions of specimens are in millimetres; abbreviations are as follows:

D = diameter, Wb = whorl breadth, Wh = whorl height, U = umbilical diameter.

Figures in parentheses are dimensions as a percentage of total diameter.

Specimen i) Wb Wh Wb/Wh U

D380 8,6 lis (209) 455) (6,9) 0,4 Los (192)

D872 139 Les (3,1) 3,0 (4S,1) 0,51 135 (0,9)

Description

D380 (Figs 1E—-F, 2A) is a microconch; the phragmocone is preserved in

clear calcite and the body chamber is preserved in grey glauconitic silt and occu-

pies half of the outer whorl. The umbilicus is partially occluded with matrix but

it seems to have steep umbilical walls. The flanks are flat and parallel, and the

venter rounded.

The small size, platycone shell shape, lack of ornament and simplified sutures

(Fig. 2A) with a trifid lateral (L) lobe clearly identify it with Patagonian Koloceras

talenkanum.

D872 (Fig. 1G) is part of a phragmocone preserved in sparry calcite. The

sutures are not preserved well enough to permit a definite identification. The

small size and shell shape suggest that it might be a macroconch of Koloceras

talenkanum.

18 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 1. Koloceras talenkanum Riccardi et al., 1987. A-C. SAM-—PC8394. D. SAM-—PC8393.

E—-F. NMB D380. G. NMB D872. H-I. SAM-—PC8392. A-D, H-I. From Estancia La Vega,

Santa Cruz Province, Argentina, upper part of the Rid Mayer Formation, Late Albian. E-F.

From locality 51, Zululand, Mzinene Formation, Albian IV-V. G. From locality 55, Zululand,

Mzinene Formation, Albian V. All x 4.

CRETACEOUS FAUNAS FROM SOUTH AFRICA 19

Fig. 2. External suture lines of Koloceras talenkanum Riccardi et al., 1987. A. NMB D380.

B. SAM—PC8392. Scale bar for size.

Discussion

These two specimens do not add anything to the extensive descriptions of

Koloceras talenkanum provided by Riccardi et al. (1987), but they do present a

palaeobiogeographic enigma.

Thus far, Koloceras talenkanum, or the genus Koloceras for that matter, were

only known from a single locality, 0,7 km to the south-east of the Estancia La

Vega, approximately 34 km north-west of the village of Tres Lagos, where about

30-35 metres of sediment referable to the upper part of the Rid Mayer Formation

are exposed. According to Riccardi et al. (1987: 118), Koloceras talenkanum

seems to occur throughout the extent of the exposure. The associated ammonite

fauna consists of a strange mixture of apparent endemic, South Gondwanid, and

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

cosmopolitan species respectively, which have been referred to the Puzosia

vegaensis Assemblage Zone (Riccardi et al. 1987: 121), and includes Puzosia

vegaensis Leanza, Hypophylloceras lestai Leanza, Partschiceras sp., Anagaudry-

ceras pulchrum (Crick), Eomarshallites espinosum Medina & Rinaldi,

E. hibridum Medina & Rinaldi (= ‘Parasilesites’ desmoceratoides (Stolley)),

? Umsinenoceras, Mortoniceras (Neokentroceras) tarense H. Leanza, Labeceras

singulare (A. Leanza), L. crassetuberculatum magnum Aguirre Urreta & Ric-

cardi, and Myloceras (Calliscaphites) andinus A. Leanza, (cf. A. Leanza 1970,

Nullo et al. 1981; H. Leanza 1986; Medina & Rinaldi 1986; Riccardi et al. 1987;

Aguirre Urreta & Riccardi 1988). The bivalve genus Maccoyella Etheridge

(1892)—thus far only known from Patagonia, Australia and New Zealand—is

also common (see Waterhouse & Riccardi 1970).

Detailed synonymies still have to be worked out for some of the above spe-

cies, but cosmopolitan forms include Phylloceras (H.), Partschiceras and Ana-

gaudryceras pulchrum; the labeceratinae are typical South Gondwanid forms (see

Klinger 1989 for a review); and endemic forms are Eomarshallites and Kolo-

ceras (the latter with the exception of the Zululand specimen(s?)). Representa-

tives of the subfamily Mortoniceratinae are conspicuously rare. Up to now only

a fragment of Neokentroceras has been recorded from Estancia La Vega

(H. Leanza 1985).

Judging by the similarity of some of the labeceratine faunas (see Aguirre

Urreta & Riccardi 1988; Klinger 1989), an open marine connection must have

existed between part of the Patagonian Austral basin and the Zululand basin. We

can think of no single factor to explain the selective barrier between the two

basins, permitting free interchange of cosmopolitan and part South Gondwanid

faunas but yet restricting interchange of Eomarshallites and Koloceras on the one

side and Mortoniceratinae on the other. On a less negative note, we have to bear

in mind that Estancia La Vega is thus far the only Upper Albian locality known

in Patagonia. Future exploration may well diminish the apparent endemic char-

acter of the Upper Albian ammonoid fauna of Patagonia.

ACKNOWLEDGEMENTS

We thank C. M. Joubert for bringing these specimens to our notice. Both

were collected in 1922-23 by the late Dr E. C. N. van Hoepen in a section of the

Mzinene River that is now flooded due to extensive damming downstream. We

pay tribute to Van Hoepen’s meticulous collecting activities—the fruits of which

lay undiscovered for more than sixty years. Drs M. B. Aguirre Urreta, F. Med-

ina, G. Blasco de Nullo (Buenos Aires) and A. C. Riccardi (La Plata) allowed

access to their collections and provided stratigraphic and other data to H.C.K.

during a visit to Argentina in 1986. Dr G. Blasco de Nullo provided the figured

Argentinian specimens. Their generous assistance is gratefully acknowledged.

Sally Dove prepared the photographic plates and Jacqueline Blaeske did the

CRETACEOUS FAUNAS FROM SOUTH AFRICA TA

drafting. Financial assistance from the Foundation for Research Development to

Klinger and from the NERC to Kennedy is gratefully acknowledged.

REFERENCES

AGUIRRE URreETA, M. B. & Riccarpi, A. C. 1988. Albian heteromorph ammonoids from

southern Patagonia, Argentina. Journal of Paleontology 62 (4): 598-614.

BRUNNSCHWEILER, R. O. 1959. New Aconeceratinae (Ammonoidea) from the Albian and

Aptian of Australia. Bulletin. Bureau of Mineral Resources, Geology and Geophysics,

Australia 54: 1-19.

Casey, R. 1954. Falciferella, a new genus of Gault ammonites, with a review of the family

Aconeceratidae in the British Cretaceous. Proceedings of the Geologists’ Association 65

(3): 262-277.

Cooper, M. R. & KENNEDY, W. J. 1979. Uppermost Albian (Stoliczkaia dispar Zone) ammo-

nites from the Angolan littoral. Annals of the South African Museum 77 (10): 175-308.

KENNEDY, W. J. & KLINGER, H. C. 1975. Cretaceous faunas from Zululand and Natal, South

Africa. Introduction, stratigraphy. Bulletin of the British Museum (Natural History)

(Geology) 25 (4): 263-315, 1 pl.

KENNEDY, W. J. & KLINGER, H. C. 1979. Cretaceous faunas from Zululand and Natal, South

Africa. The ammonite superfamily Haplocerataceae Zittel, 1884. Annals of the South

African Museum 77 (6): 85-121.

Kuincer, H. C. 1976. Cretaceous heteromorph ammonites from Zululand. Memoirs. Geologi-

cal Survey South Africa 69: 1-142.

Kuincer, H. C. 1989. The ammonite subfamily Labeceratinae Spath, 1925: systematics, phy-

logeny, dimorphism and distribution (with a description of a new species). Annals of the

South African Museum 98 (7): 189-219.

LEANZA, A. F. 1970. Ammonites nuevos 0 pocos conocidos del Aptiano, Albiano y Ceno-

maniano de los Andes australes con notas acerca de su posicion estratigrafica. Revista de

la Asociacion geologica argentina 25 (2): 197-261.

LEANZA, H. 1986. Un nuevo mortoniceratido (Cephalopoda-Ammonoidea) del Albiano

superior de Santa Cruz, Argentina. Ameghiniana 22 (3-4): 249-254.

McNamara, K. J. 1978. Myloceras (Ammonoidea) from the Albian of central Queensland.

Alcheringa 2 (3): 231-242.

McNamara, K. J. 1985. A new micromorph ammonite genus from the Albian of South Aus-

tralia. Special Publication. South Australia Department of Mines and Energy 5: 263-268,

Ol, Ie

Mepina, F. & Rinatpi, C. A. 1986. Ammonoideos del Albiano superior en la Formacion Rid

Mayer de la Estancia La Vega, Provincia de Santa Cruz. Contribucion. Direccion Nacional

del Antartico, Instituto Antarctica Argentina 324: 1-34, pls 1-S.

NuLLo, F., PRosErpIO, C. A. & BLASCO DE NULLO, G. 1981. El Cretacico de la Cuenca Austral.

In: VOLKHEIMER, W. & Mussacuio, E. eds. Cuenas sedimentarias del Jurdsico y

Cretacico 1: 181-220. Buenos Aires: Comité Sudamericano del Jurasico y Cretacico.

RIccARDI, A. C., AGUIRRE URRETA, M. B. & MepINA, F. A. 1987. Aconeceratidae

(Ammonitina) from the MHauterivian—Albian of southern Patagonia. Palaeonto-

graphica (A) 196 (4-6): 105-185.

SPATH, L. F. 1925. On Upper Albian Ammonoidea from Portuguese East Africa, with an

appendix on Upper Cretaceous ammonites from Maputoland. Annals of the Transvaal

Museum 11 (3): 179-200.

WATERHOUSE, J. B. & Riccarpi, A. C. 1970. The Lower Cretaceous bivalve Maccoyella in

Patagonia and its paleogeographic significance for continental drift. Ameghiniana 7 (3):

281-296.

WHITEHOUSE, F. W. 1926. The Cretaceous Ammonoidea of eastern Australia. Memoirs of the

Queensland Museum 8: 195-242, pls 34-41.

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

larly Articles 22 and 51).

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Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

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counter to Recommendation 23 of the Code, to meet the requirements of Biological Abstracts.

HERBERT CHRISTIAN KLINGER

WILLIAM JAMES KENNEDY

CRETACEOUS FAUNAS FROM ZULULAND

AND NATAL, SOUTH AFRICA.

A KOLOCERAS (CEPHALOPODA,

AMMONOIDEA) FROM THE MZINENE

FORMATION (ALBIAN)

7 VOLUME 99 PART 3 FEBRUARY 1990 | ISSN 0303-2515

-ANNALS

OF THE SOUTH AFRICAN

~ MUSEUM |

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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): 1-51.

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 99 Band

February 1990 Februarie

ant soe Dee!

A NEW GENUS OF RUTITRIGONIINAE

(BIVALVIA, TRIGONIACEA)

FROM THE LOWER CRETACEOUS

(APTIAN) OF ZULULAND

MICHAEL R. COOPER

Cape Town Kaapstad

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A NEW GENUS OF RUTITRIGONIINAE

(BIVALVIA, TRIGONIACEA)

FROM THE LOWER CRETACEOUS (APTIAN) OF ZULULAND

MICHAEL R. COOPER

Department of Geology, University of Durban-Westville, Natal

(With 1 figure)

[MS accepted 12 September 1989]

ABSTRACT

Trigonia (Rutitrigonia) pongolensis Rennie lacks the characters of Rutitrigonia, and is made

the type species of the new genus Zulutrigonia. Other species to be referred here include

T. krenkeli Lange and T. kigombana Aitken from the Neocomian—Aptian of Tanzania.

CONTENTS

PAGE

MN GHOCU Ci Omer we ateneg eater aie para a ent cna canynd ani cic. a ore mania eat Niel tease Op

SyStemavicdeSchipUOmMerme ie Atk Tae sean Crates at aiie jocles has eu: 24

INCKnOwledeementSraser tne tae: Globe Ie aati me oee se oss 28

INCH INCC Sarma ine ey Cen ines mie ener se niente aiean ees Si ue Siete a Miele 8 28

INTRODUCTION

In a benchmark paper representing a marked departure from the conser-

vative taxonomy of that period, Van Hoepen (1929) introduced a number

of subfamilies and genera for trigoniid bivalves from the Cretaceous of Zululand.

Among these was the genus Rutitrigonia, introduced for R. peregrina van Hoepen

from the upper Middle or low Upper Albian. This taxon formed the basis for the

monogeneric subfamily Rutitrigoniinae van Hoepen, corresponding broadly with

the Excentrica group of Lycett (1879).

Van Hoepen’s (1929) bold taxonomy failed to make immediate impact, and

Rutitrigonia was reduced to a subgenus of Trigonia by Rennie (1936), whereas

Cox (1952) and Aitken (1961) treated it as a subgenus of Megatrigonia. However,

Crickmay (1932), Kobayashi (1957), Saveliev (1958), Nakano (1961, 1963, 1965,

1977), Cox (1969), Poulton (1977, 1979) and Reyes & Pérez (1978) have all used

Rutitrigonia as a valid genus, and it is now widely accepted.

In addition to the type species, the following species have also been assigned

to Rutitrigonia: R. affinis (Sowerby), R. agrioensis (Weaver), R. amagensis (Koba-

yashi), R. beyrichi (Krumbeck), R. beyschlagi (Muller), R. bornhardti (Miller),

Ann. S. Afr. Mus. 99 (3), 1990: 23-29, 1 fig.

24 ANNALS OF THE SOUTH AFRICAN MUSEUM

R. coquandiana (d’Orbigny), R. dietrichi (Lange), R. dunscombensis (Lycett),

R. excentrica (Parkinson), R. jacksonensis (Packard), R. janenschi (Lange),

R. kigombana (Aitken), R. laeviuscula (Lycett), R. lerchi (Hill), R. longa

(Agassiz), R. mesembris (Tenison-Woods), R. niongalensis (Lange), R. nossae

Aitken, R.nyangensis Aitken, R. pongolensis (Rennie), R. sanchuensis

(Nakano), R. schwarzi (Miller), R. semiculta (Forbes), R. skorkovensis Pugaczew-

ska, R. syrdariensis (Archangelsk’ii), R. weaveri (Stoyanow), and R. yeharai

Kobayashi.

In introducing the new species Trigonia (Rutitrigonia) pongolensis, Rennie

(1936: 358) noted that *. . . It is not without some hesitation that I associate with

Rutitrigonia the species T. pongolensis sp. nov. and T. krenkeli. The concentric

ribbing of the neanic stage is of such a character in both these species as to exclude

them from /ndotrigonia in spite of adult similarities, and Rutitrigonia is the only

term available for species of Trigonia having features at all like those of the last

two species.’

Nakano (1963) excluded Trigonia (Rutitrigonia) pongolensis Rennie from his

concept of Rutitrigonia because of its coarse oblique flank costae, referring it

instead to Megatrigonia. Subsequently, he identified it (Nakano 1965) as

Megatrigonia? pongolensis (Rennie).

Given the large number of Rutitrigonia species now known, it is clear that

Trigonia pongolensis and its allies can no longer be accommodated satisfactorily

in this genus. It is proposed, therefore, to introduce a new genus for these forms.

SYSTEMATIC DESCRIPTION

Superfamily TRIGONIACEAE Lamarck, 1819

Family Trigoniidae Lamarck, 1819

Subfamily Rutitrigoniinae van Hoepen, 1929

Diagnosis

Shell small to moderately large, subovate to pyriform, subtrigonal and

subtrapezoidal. Typically with inner and marginal carina lacking, resulting in poor

discrimination between the smooth escutcheon and area. The latter is ornamented

only in the nepionic stages and may be weakly bipartite. In some taxa there is a

more or less prominent umbonal ridge. Nepionic ornament comprises concentric

costae that pass uninterrupted on to the area where they weaken across the

longitudinal furrow. Later flank ornament comprises costae that are subconcen-

tric anteriorly, but posteriorly may be subconcentric, strongly oblique, or effaced.

Discussion

The origins of Rutitrigonia are cryptic. It first appears in the Bathonian of

East Africa (Venzo 1942), and its ancestry has been linked to the myophorelline

Frenguelliella (Kobayashi 1957; Nakano 1963). However, it seems to be Gon-

A NEW GENUS OF RUTITRIGONIINAE Ds,

dwanic in origin and there is a noteworthy similarity with primitive Anditrigonia

species such as the Lower Callovian A. keideli (Weaver) (cf. Leanza & Garate,

1987, pl. 3 (fig. 1)). Like Van Hoepen (1929) and Cox (1952), the writer considers

the Megatrigoniinae and Rutitrigoniinae to be closely allied; they are here

regarded as sister taxa.

Genus Zulutrigonia gen. nov.

Type species. Trigonia (Rutitrigonia) pongolensis Rennie, 1936; by original

designation herein.

Diagnosis

Shell medium sized, subtrapezoidal, with inconspicuous subterminal

umbones and opisthogyrous beaks. Anterior margin weakly convex, posterodorsal

margin almost straight and respiratory margin obliquely truncate. Escutcheon

narrow, indistinct, not sunken. Broad subtrigonal area essentially flat, meeting

the posterodorsal commissure in an acute angle, so that the shell is fastigiate

(roof-shaped) posteriorly. Ornament comprises coarse, robust, relatively sparse

costae that are subconcentric anteriorly and curve strongly upwards posteriorly,

cutting obliquely across the growth striae. Age: Lower Cretaceous (Neocomian—

Aptian).

Discussion

Zulutrigonia gen. nov. is distinguished from Rutitrigonia by its subtrapezoidal

outline, with subterminal umbones, almost straight posterodorsal margin and

obliquely truncate siphonal margin. In addition, the shell is fastigiate postero-

dorsally, due to the broadly flattened areas meeting the commissure at an acute

angle, and the coarse, robust flank costae curve strongly upwards posteriorly.

There are noteworthy similarities between Zulutrigonia gen. nov. and Bucho-

trigonia (Syrotrigonia) (Cox, 1952). Both are characterized by moderate inflation,

subterminal umbones, a straight posterodorsal margin and obliquely truncate res-

piratory margin, an inconspicuous to near-obsolete escutcheon, and broad flat-

tened areas that produce a strongly fastigiate profile to the shell posterodorsally.

The concentric ornament of the nepionic and middle growth stages is identical.

Syrotrigonia may be distinguished, however, by its more ovate outline, the posses-

sion of a marginal carina that becomes a tuberculate umbonal ridge in maturity,

the presence of a broad shallow antecarinal sulcus in maturity across which ribbing

weakens or is effaced, and the generally V-shaped costae of the adult. Significantly,

however, the V-shaped costae are far from uniformly developed and in some indi-

viduals examined they simply bend strongly upwards as in Zulutrigonia gen. nov.

Similarly, the strong marginal carina that was emphasized in the original diagnosis

is not a true carina (the latter is restricted to the nepionic stages) but an umbonal

ridge whose prominence is enhanced by the bounding antecarinal sulcus. Initially

the tubercles on this ridge are no more than rib endings that are emphasized by

the weakening of ornament across the antecarinal sulcus. Whether the similarities

26 ANNALS OF THE SOUTH AFRICAN MUSEUM

are due to phylogeny or are the result of convergence is, at present, unclear. Syro-

trigonia is here elevated to full generic status because Buchotrigonia has radial flank

costae in the early to middle growth stages, an internally crenulated postero-

ventral margin and transverse costellae to the escutcheon (cf. Pérez & Reyes 1980),

characters that ally it with the Apiotrigoniinae and Pterotrigoniinae.

Zulutrigonia pongolensis (Rennie, 1936)

Fig. LA-E

Trigonia (Rutitrigonia) pongolensis Rennie, 1936: 359, pl. 41 (figs 5-6), pl. 42 (figs 5-7).

Megatrigonia (Rutitrigonia) pongolensis (Rennie) Cox, 1952: 59.

Megatrigonia pongolensis (Rennie) Nakano, 1963: 527.

Megatrigonia? pongolensis (Rennie) Nakano, 1965: 17.

Type material

The holotype, SAM—PCZ8528, and paratype, SAM—PCZ8529, both in the

South African Museum, Cape Town.

Type locality

The material was collected by Drs S. H. Haughton and A. W. Rogers along

the Mfongosi stream, a tributary of the Pongola, at their locality Z2. It is from the

Mlambongwenya Formation (Cooper & McCarthy 1988), which here is of late

Aptian age and corresponds broadly with Locality 164 of Kennedy & Klinger

OTS).

Description

Shell medium sized (maximum length about 60 mm), longer than high

(H/L = 0,73-0,80), subtrapezoidal, with low subterminal umbones and incurved

opisthogyrous beaks. The valves are moderately inflated (W/L = 0,20—0,22) and

maximum inflation is just behind the umbones. The broad, weakly curved

anterior margin is subvertical and curves evenly into the long, almost straight,

ventral margin. The broad, obliquely truncate respiratory margin meets the

straight posterodorsal margin in a very obtuse angle. The dorsal and ventral

margins are subparallel.

The smooth, narrow, flat, lanceolate escutcheon is poorly discriminated and

can be distinguished from the area only in the nepionic stages when it is

ornamented. Marginal and inner carinae are lacking, but an umbonal ridge marks

the position of the marginal carina. The broad subtrigonal area is weakly concave,

almost flat, and with an indistinct longitudinal groove. It forms an acute angle

with the opposing area at the posterodorsal commissure so that the posterodorsal

part of the shell is fastigiate (roof-shaped).

The ornament of the nepionic stages comprises coarse concentric costae,

broader than the interspaces, which flex slightly at the umbonal ridge before

crossing onto the area. The ribs thin noticeably where they cross the longitudinal

groove. Later flank costae are coarse, robust, relatively distant and slightly

A NEW GENUS OF RUTITRIGONIINAE Di)

Fig. 1. A-E. Zulutrigonia pongolensis (Rennie). A-—C. Anterior, lateral and dorsal views of the

paratype, SAM-—PCZ8529. D-E. Lateral and dorsal views of the holotype, SAM—PCZ8528.

F. Zulutrigonia krenkeli (Lange). A plastercast of the holotype, in the South African Museum.

All x 1.

28 ANNALS OF THE SOUTH AFRICAN MUSEUM

narrower than the interspaces. They are subconcentric anteriorly but curve

strongly upwards to the posterior, cutting the growth striae obliquely and

approaching the area almost at right angles. In the middle growth stages the ribs

terminate at the umbonal ridge but in maturity they weaken and become effaced

before reaching the area. The flank costae become obsolete on the anterolateral

shoulders, leaving the narrow anterior face crossed only by growth striae.

Discussion

Other species to be assigned to Zulutrigonia include the Tanzanian Trigonia

krenkeli Lange (1914: 230, pl. 20 (fig. 2)) and T. kigombana Aitken (1961: 95,

pl. 13 (fig. 3)), both from the Rutitrigonia schwarzi Beds of the Tendaguru

succession and thus of Hauterivian age.

Zulutrigonia krenkeli (Lange) (1914: 230, pl. 20 (fig. 2)) (Fig. 1F) differs

from Z. pongolensis in having dorsal and ventral margins that converge

posteriorly and with a lower, less-fastigiate posterodorsal profile.

Zulutrigonia kigombana (Aitken) (1961: 95, pl. 13 (fig. 3)) differs from

Z. pongolensis in being strongly produced posteriorly, with a gently concave pos-

terodorsal margin, and with the posterior one-third of the flanks unornamented.

Trigonia inca Fritzsche (1921: 49, pl. 3 (figs 1-3)) from the Barremian of

northern Peru shows some similarity to Zulutrigonia. However, its flank costae

form distinct chevrons in maturity and hence it may be a Syrotrigonia

(Etayo-Serna 1985; Pérez & Reyes 1986).

Occurrence

Zulutrigonia pongolensis (Rennie) is currently known only from the late

Aptian of northern Zululand.

ACKNOWLEDGEMENTS

I thank Drs M. Tashiro (Kochi) and E. Pérez d’Angelo (Santiago) for their

constructive criticism of the manuscript. Betsie Greyling assisted with the

photography and the University of Durban-Westville is thanked for a grant in aid

of publication.

REFERENCES

AITKEN, W. G. 1961. Geology and palaeontology of the Jurassic and Cretaceous of southern

Tanganyika. Bulletin of the Geological Survey of Tanganyika 31: 1-144.

Cooper, M. R. & McCartny, M. J. 1988. The stratigraphy of the Uloa Formation. Extended

Abstract, Geocongress ’88, Durban: 121-124. Durban: Geological Society of South Africa.

Cox, L. R. 1952. Notes on the Trigoniidae, with outlines of a classification of the family.

Proceedings of the Malacological Society of London 29 (2-3): 45-70.

Cox, L. R. 1969. Trigoniacea Lamarck, 1859. In: Moore, R. C. ed. Treatise on invertebrate

paleontology. Part N, Mollusca 6 (1): N471—-N489. Boulder: Geological Society of

America; Lawrence: University of Kansas.

Crickmay, C. H. 1932. Contributions toward a monograph of the Trigoniidae. American

Journal of Science 24 (5): 443-464.

A NEW GENUS OF RUTITRIGONIINAE 29

ETAYO-SERNA, F. 1985. Paleontologia estratigrafica del sistema Cretacico en la Sierra Nevada

del Cocuy. Publicaciones Especiales Ingeominas, Colombia, No. 16, Proyecto Cretdacico,

Capitulo 24: 1-47.

FritzscHe, C. H. 1921. Neue Kreidefaunen aus Siidamerika (Chile, Bolivia, Peru, Colombia).

In: STEINMANN, G. ed. Beitrdge zur Geologie und Paldontologie von Stidamerika. Neues

Jahrbuch fiir Mineralogie, Geologie und Paldontologie 50: 1-56, 313-334.

KENNEDY, W. J. & KLINGER, H. C. 1975. Cretaceous faunas from Zululand and Natal, South

Africa. Introduction, stratigraphy. Bulletin of the British Museum of Natural History

(Geology) 25 (4): 265-315.

KopayasHl, T. 1957. Nipponitrigonia and Rutitrigonia in Japan. Transactions and Proceedings

of the Palaeontological Society of Japan (n.s.) 26: 51-61.

Lance, E. 1914. Die Brachiopoden, Lamellibranchiaten und Anneliden der Trigonia

schwarzi-Schicht, nebst verleichender Ubersucht der Trigonien der gesamten Tendagur-

uschichten. Archiv ftir Biontologie 3 (4): 193-289.

LEANZA, H. A. & Garate, J. I. 1987. Faunas de Trigonias (Bivalvia) del Jurasico y Cretacico

Inferior de la Provincia del Nequén, Argentina, conservadas en el Museo Juan Olsacher de

Zapala. In: VOLKHEIMER, W. ed. Bioestratigrafia de los Sistemas Regionales del Jurasico y

Cretacico de América del Sur 1: 201—255. Mendoza.

Lycetr, J. 1872-79. A monograph of the British fossil trigoniae. Monograph of the

Palaeontological Society: 1-52 (1872); 53-92 (1874); 93-148 (1875); 149-204 (1877);

205-245 (1879).

Nakano, M. 1961. On the Trigoniinae. Journal of Science of the Hiroshima University

(C, Geology and Mineralogy) 4 (1): 71-94.

NAKANO, M. 1963. On the Rutitrigoniinae. Geological Report of the Hiroshima University 12:

513-529.

NAKANO, M. 1965. On the Megatrigoniinae. Journal of Science of the Hiroshima University

(C, Geology and Mineralogy) 5 (1): 13-20.

Nakano, M. 1977. A new genus Levantotrigonia nov. gen. Journal of Science of the Hiroshima

University (C, Geology and Mineralogy) 7 (4): 199-202.

PEREZ, E. & Reyes, R. B. 1980. Buchotrigonia (Buchotrigonia) topocalmensis sp. nov.

(Trigoniidae; Bivalvia) del Cretacico superior de Chile. Revista Geologica de Chile 9:

37-55.

PEREZ, E. & Reyes, R. B. 1986. Presencia de Buchotrigonia (Syrotrigonia) Cox, 1952 (Bivalvia;

Trigoniidae) en Sudamérica y descripcion de dos especies nuevas. Revista Geologica de

Chile 28-29: 77-93.

PouLtton, T. P. 1977. Early Cretaceous trigoniid bivalves of Manning Provincial Park,

southwestern British Columbia. Geological Survey Paper. Mines and Geology Branch,

Canada 76-9: 1-25.

Poutton, T. P. 1979. Jurassic trigoniid bivalves from Canada and the western United States of

America. Bulletin. Geological Survey of Canada 282: 1-82.

RENNIE, J. V. L. 1936. Lower Cretaceous Lamellibranchia from northern Zululand. Annals of

the South African Museum 31 (3): 277-391.

Reyes, R. B. & PEREZ, E. 1978. Las trigonias del titoniano y cretacico inferior de la cuenca

andina de Chile y su valor cronoestratigrafico. Boletin de Instituto de Investigaciones

Geoldgicas de Chile 32: 1-105.

SAVELIEV, A. A. 1958. The Lower Cretaceous trigontids of Mangyschlak and western

Turkmenia. Trudy Vsesoyuznogo neftyanogo nauchno-issledovatel’skogo geologorazved-

ochnogo instituta (VNIGRI) 125: 1-516. (In Russian.)

VAN Hoepen, E. C. N. 1929. Die krytfauna van Soeloeland. 1. Trigontidae. Paleontologiese

Navorsing van die Nasionale Museum van Bloemfontein 1 (1): 1-38.

VeENZO, S. 1942. Trigonia (Laevitrigonia) stefaninii n. sp. del Batoniano dell’Oltregiuba.

Revisione del sottogenere e sua distribuzione. Atti della Societe italiana di scienze naturali,

e del Musei civico di storia naturale 81 (3-4): 210-229.

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

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Holotype

SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach, Port Eliza-

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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.

MICHAEL R. COOPER

A NEW GENUS OF RUTITRIGONIINAE

(BIVALVIA, TRIGONIACEA)

FROM THE LOWER CRETACEOUS

| (APTIAN) OF ZULULAND

'OF THE SOUTH AFRICAN

OR EE Mose

CAPE TOWN

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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, 1969b) 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)

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

FiscuEr, 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

1903-1905 4 (15). Denkschriften der medizinisch-naturwissenschaftlichen Gesellschaft zu Jena 16: 269-270.

(continued inside back cover)

‘

itn)

‘ :

fe ?

Fig. 1. Pennatulaceans of southern Africa. A-C. Actinoptilum molle Kikenthal, 1910).

D-F. Virgularia schultzei Kikenthal, 1910. G. Echinoptilum echinatum (Kikenthal, 1910).

H-I. Echinoptilum macintoshii Hubrecht, 1885.

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 99 Band

May 1990 Mei

Part 4 Deel

THE PENNATULACEA OF SOUTHERN AFRICA

(COELENTERATA, ANTHOZOA)

GARY C. WILLIAMS

Cape Town Kaapstad

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C460

THE PENNATULACEA OF SOUTHERN AFRICA

(COELENTERATA, ANTHOZOA)

Gary C. WILLIAMS*

Department of Marine Biology, South African Museum, Cape Town, South Africa

and Department of Zoology, University of Cape Town, Rondebosch, South Africa

(With 41 figures)

[MS accepted 29 April 1988]

ABSTRACT

Nineteen species of pennatulacean octocorals from sublittoral and bathyal regions of

southern African coastal waters are described and illustrated. Six additional species that are

recorded in the literature are mentioned, although material was not available for examination.

Two of these, Cavernulina cylindrica and Virgularia juncea, are considered dubious records and

are not included in the faunal list provided for the region. Presumably young or partial colonies

or problematic taxa representing seven genera (Cavernulina, Kophobelemnon, Scleroptilum,

Calibelemnon, Virgularia, Halipteris, and Pteroeides) are not identified to species but are

mentioned in the text and included in the list of species. A total of 23 determined and at least

seven undetermined species representing 20 genera in 12 families are recorded. Three species

that have been recorded from Angola, which is north of the geographic limit of this paper, are

also mentioned: Virgularia tuberculata, Pteroeides griseum, and Crassophyllum cristatum; these

may possibly be encountered further south.

Fourteen new geographic records are established, including the first southern African

records for the following ten genera: Cavernulina, Echinoptilum, Kophobelemnon, Distichop-

tilum, Scleroptilum, Calibelemnon, Chunella, Amphiacme, Scytaliopsis, and Halipteris.

Scanning electron micrographs of sclerite form or axial morphology are presented for many

species. A key is provided to all species treated in the present work. A brief discussion of

faunistic considerations is also included.

CONTENTS

PAGE

MTGhOGUCtIOM peer teeta rm ye ein ug is retin arn OaMtve aia Seg 32

Watentalsrandumethods qs sn cae eines ottawa ca aes o.oo clea es 55

SV SUCIMAUCrACCOUMUr ier eerie raesst sie er alam sie ene me Laicia Sue lane urun leit 55

Key to the families of Pennatulacea from southern Africa.......... 315)

Key to the species of Pennatulacea from southern Africa .......... 36

List of Pennatulacea recorded from southern Africa............... 39

DD CSCHIP CONS ye See etn te ioe ohio Fie e soa iennnte alee 40

Any ene Clllidae rem metre meena rye Macias ok mys. shee a oi 40

Banulyse chino ppilidacwwe. 4 omecce ne ciate ns ane ere: sheen es 50

BanulyaCophobelemmidaeyes x20. osinee se ce te ees se dante G 64

Banuilye Amino ptilidacwmiere sare sya si-) Sellers cece oi olsun rosie weary 68

RannilyaumiculiniGdaen ie ee cas ete so hae ols sd ain os so alate ale lores 69

Bamilyperocopulidace nr tei. ese | ee ear oe rea ei mal

* Present address: Department of Invertebrate Zoology, California Academy of Sciences,

Golden Gate Park, San Francisco, California 94118, U.S.A.

Sil

Ann. S. Afr. Mus. 99 (4), 1990: 31-119, 41 figs.

32 ANNALS OF THE SOUTH AFRICAN MUSEUM

PAGE

Family Scleroptilidaci...5 3225 (ae ace re ieee ne i

Family Chunellidae. 0%... 40055 eer ek nk ee ene 76

Family Umbellulidae <...0.25 29) 58s eos ee eels eee eee 81

Family Virgulamidae. .0.5..a.36 5 eae ote oot soe eee tee oe 86

Family Pennatulidaes.%..5 22) 44s ce tee er een 99

Family Pteroeididac’s 0.2). a aes a serees oehe eeee 106

DISCUSSIONS ie 8 ot occ eee tas a eae ce cs oy ee ee 112

Summary and conclusions sxe. 8. ae aaerecroe encie roe ae eee Rear ee 113

Acknowledements...4 2. s5.ncgss sake so sea Ooenak os eee ee 11S

FRELETENCES > ac: oh Gif whee Ps Gece ea een cs NOIRE TRS OS te EE ne eet CC 114

INTRODUCTION

At present, octocoral classification is frequently an equivocal and disputa-

tious field (Weinberg 1976: 63). It is clear that our knowledge concerning this

group is far from adequate; a great deal of research needs to be accomplished in

order to attain the goal of a world-wide synthesis of the biology and classification

of octocorals (Bayer 1981a: 11). A substantial amount of confusion has arisen in

the literature because of a general lack of knowledge concerning variability; some

octocoral species can be exceedingly variable and many authors have described

species or even genera based on intraspecific variants. Extensive collecting in

many different geographical localities and detailed comparison of material is

needed to assess the degree of variation in many taxa due to genetic, geo-

graphical, or ecological differences.

The goal of the present study of the systematics and faunistics is to contribute

toward a world-wide synthesis of the classification and distribution of octocorals by

concentrating on a localized fauna that has been previously only minimally to

moderately known. This work is the first of four proposed parts describing the

octocorals of southern Africa. Subsequent papers will deal with additional species

of pennatulaceans from the Cape, Transkei and Zululand; stoloniferous octocorals

and soft corals (families Clavulariidae, Tubiporidae, Coelogorgiidae, Alcy-

oniidae, and Nephtheidae); and the shallow-water gorgonians (families Antho-

thelidae, Melithaeidae, Keroeididae, Acanthogorgiidae, Plexauridae, Gorgonii-

dae, Ellisellidae, Chrysogorgiidae, Primnoidae, and Isididae). The four papers

together will attempt to synthesize our knowledge of southern African octocorals.

The studies exclude the deep-water gorgonians, primarily of the families

Ellisellidae, Primnoidae, and Chrysogorgiidae, groups that are less well known

regionally or that require large-scale revisions before accurate identifications can be

made. Many octocoral species that have been collected still need to be identified

and new species are continually being discovered.

The present work is intended not only for specialists in the field of octocoral

systematics but also as a guide for other biologists, e.g. ecologists, fisheries

biologists and biogeographers, who share the common need to identify material

from benthic surveys and other studies. Southern African octocorals are of

interest to non-specialists, since they comprise such a major part of the

SOUTHERN AFRICA PENNATULACEA 33

shallow-water benthic fauna. Therefore, sections have been added to the texts of

these papers that would not normally be included in a work intended solely for

specialists. These include brief diagnoses of the families and genera, sections on

general characteristics of the octocoral groups, and illustrations of the entire

colonies of all species, as well as figures depicting diagnostic features, sclerites,

and other features.

The monograph by Kikenthal (1915) remains the most comprehensive work

on pennatulaceans of the world and is certainly the most thorough account up to

that time. Modern revisions of many of the genera are obviously needed. Other

major reviews on sea pens are mainly monographs produced from the large

Oceanographic expeditions of the past and include K6lliker (1872a, 1872b),

Kolliker (1880) (Challenger), Danielssen & Koren (1884) (Norwegian North

Atlantic), Kukenthal & Broch (1911) (Valdivia), Hickson (1916) (Siboga), and

Broch (1958b) (Discovery). Thomson (1915, 1917) described the pennatulacean

fauna of South Africa known at that time and Bayer (1956) reviewed the known

world genera. Tixier-Durivault (1954, 1960, 196la, 19666, 1972) and Tixier-

Durivault & D’Hondt (1974) described material from South Africa, Mozambique,

Angola, and Madagascar (including neighbouring islands). General accounts of

the biology of sea pens are by Hickson (1909) and Hyman (1940). More recently,

the distribution and taxonomy of pennatulaceans has been reported on by

Pasternak (1960, 1961a, 1961b, 1962, 1964, 1970, 1975a, 1975b, 1980), Grasshoff

(1972, 1981, 1982a, 1982b), and D’Hondt (1984a, 19845).

Bayer (1981a: 7) classified various geographic regions of the world into four

general categories of taxonomic knowledge pertaining to octocorals: essentially

complete; moderately well known; poorly known; and minimally known. He

considered South Africa, in the region of the Cape of Good Hope, to be

moderately well known and described this category as follows: ‘where there is

extensive literature, but many more species remain to be described and taxonomic

problems to be solved, and the major patterns of distribution must yet be worked

out. Much descriptive work remains to be done before ecological and experimental

studies can proceed at an effective level’. The rest of southern Africa is less well

known and certain regions fall into Bayer’s latter categories—being poorly or even

minimally known. Two such regions include the Atlantic coast between 20°S and

33°S, and the northern Natal—southern Mozambique region of the Indian Ocean

between 20°S and 29°S.

Southern Africa is here defined as the African coastal region that lies south

of 20°S—an arbitrary limit used by Millard (1978) in a zoogeographical analysis

of southern African hydroids, which are another diverse and widespread benthic

group. This region extends from the Skeleton Coast of Namibia in the Atlantic,

round the coast to the vicinity of Beira, Mozambique, in the Indian Ocean.

Material examined in the present work was obtained throughout this range from

the continental shelf, slope and rise, and from the littoral to depths of 3 000 m.

The island of Madagascar is not included here, since its affinities are strictly

Indo-Pacific and not directly associated with the southern African fauna.

34 ANNALS OF THE SOUTH AFRICAN MUSEUM

Pennatulaceans from geographical regions marginal or in proximity to the area of

the present study have been described by Broch (19585) and Tixier-Durivault

(196la, 1963) for Angola, and Tixier-Durivualt (19666, 1972) and Tixier-

Durivault & D’Hondt (1974) for Madagascar.

Of the 14 presently recognized pennatulacean families of the world, 12 (or

86 %) are represented in southern Africa, and of the 34 world genera, at least 20

(or 59 %) are known from this region. The only families not represented in the

regional fauna are two monogeneric ones: the Renillidae—genus Renilla

Lamarck, 1816, with four or five species restricted to South America and the

southern part of North America; and the Stachyptilidae—genus Stachyptilum

Kolliker, 1880, with three species presumably restricted to warmer parts of the

Pacific Ocean.

The classification system used in this study at the subordinal and familial

levels conforms to Kikenthal (1915) and Bayer (1956). Terminology conforms to

the trilingual glossary of Bayer et al. (1983). All illustrations, photographs, and

scanning electron micrographs are by the author. The abbreviation ‘Lit.!’ in a

synonymy indicates where a complete or more detailed synonymy can be found.

The notation non in the synonymies indicates a misidentification by the author

cited; partim indicates ‘in part’.

Characteristics of the Pennatulacea

The pennatulaceans, or sea pens, are considered a very distinct and greatly

specialized group of octocorallian cnidarians (Hickson 1909: 358; 19306: 229; Bayer

1956: 181). Unlike other octocorals, pennatulacean colonies are formed by a very

large polyp, the oozooid (also known as the axial, initial, principal, or primary

polyp), that by lateral budding of its body walls gives rise to all the secondary zooids

of the colony. The oozooid forms a fleshy, unbranched peduncle to anchor the

colony by peristaltic contractions in soft substrata such as mud, sand, or

abyssal-ooze deposits. The polyps of an individual colony (excluding the oozooid)

are always dimorphic, more rarely trimorphic; the distal region of the oozooid

forms a rachis bearing several to many daughter polyps, which take the form of

autozooids, siphonozooids, and sometimes mesozooids. Disregarding the oozooid.

the autozooids (Fig. 11E) are the largest polyps in polymorphic colonies and possess

eight well-developed tentacles and mesenteries; these are the feeding polyps of the

colony. Siphonozooids (Fig. 11C) are much smaller than autozooids and have

strongly developed siphonoglyphs that produce currents for internal water

circulation; the tentacles are reduced or absent. Mesozooids (Fig. 32B, D) are

polyps that are intermediate in structure between autozooids and siphonozooids.

In colonies with bilateral symmetry and well-developed polyp leaves (such as

Virgularia, Pennatula, and Pteroeides), the dorsal side refers to the side of the

rachis adjacent to the oldest (usually largest) autozooids, where the rachis appears

mostly bare and uncovered by the polyp leaves; hence, the ventral side refers to

the side of the rachis opposite to the dorsal side, where the polyp leaves cover

most or all of the rachis surface.

SOUTHERN AFRICA PENNATULACEA 35

Characteristics that are important in the classification of pennatulaceans

include the form, number, and arrangement of autozooids; distribution of

siphonozooids; the colonial symmetry, growth form, and axial development; and

form and distribution of sclerites. Pennatulacean sclerites differ from those of

other octocorals by commonly being smooth and unsculptured, sometimes

grooved or three-flanged, with tubercles usually absent but, if present, then

simply knobby or rounded, not thorny or elaborate.

MATERIALS AND METHODS

Recently procured material was collected by means of SCUBA, dredge, or

beam trawl. Colonies were relaxed in a solution of magnesium chloride mixed

with sea-water and preserved in 70 per cent ethanol. Small and fragile specimens

were first fixed in 10 per cent formalin before transfer to ethanol. Sclerites and

axial fragments were isolated by dissolving tissue in concentrated sodium

hypochlorite, with repeated rinsing in water and finally in ethanol. Minute

sclerites were pipetted from alcohol suspension and dropped on to stubs that had

circular pieces of exposed and developed black-and-white photographic paper

glued to them. After evaporation of the alcohol, the sclerites adhered well to the

paper surface. Large sclerites and fragments of axial material were applied

directly to the stub surface with SEM glue. Specimens were coated with

gold/palladium (Au/Pd). Cambridge S200 and S180 scanning electron micro-

scopes were used to examine sclerites and axial fragments.

All material examined is housed in the marine invertebrate collection of the

South African Museum (SAM), Cape Town.

SYSTEMATIC ACCOUNT

KEY TO THE FAMILIES OF PENNATULACEA FROM SOUTHERN AFRICA

eee NayaceitpOlyps tree, Not fused to) amy degree... 2.2... .... 2520025. I

1B. Proximal portions of adjacent polyps fused to some degree, forming polyp

leavestonmmndiscd mideess on ,omed only atthe bases)... /.0.........-.-- 9

2A. Polyps crowded and distributed on all sides of rachis, or rachis with a single

longitudinal furrow or V-shaped region without polyps .............. 3

2B. Polyps confined to a terminal cluster, arranged biserially along rachis, or in

groups of two to four arranged sparsely along rachis ................ 4

eee OLY I SAWALMOUC CAV CES « ayn nn. So yee sues dae be Veretillidae

3B. Polyps with non-retractile bifurcated calyces ............ Echinoptilidae

4A. Polyps arranged in a terminal cluster at the end of a long slender stalk

PE ee a eee. eG SAME in os eb aes Umbellulidae

TPE Lolypssamansed along the sides of the tachis. .7..4..:...5...0..0..- 5

SA. Coloniés clavate, distal portion of rachis distinctly wider than rest of colony

56-G.8 cig, S.9n6 Gra TRANG ne Once Oa ne ne Kophobelemnidae

5B. Colonies not clavate, but are long, thin and whip-like or staff-like..... 6

6A.

6B.

TA.

is}.

8A.

SB.

9A.

OB.

10A.

10B.

11A.

11B.

IVAN.

1B.

2A.

2B.

3A.

318)

4A.

4B.

SA.

ANNALS OF THE SOUTH AFRICAN MUSEUM

Polyps numerous (> 40) and contiguous, arranged in two longitudinal series

along rachis. Sclerites dense. Polyps retractile into calyces............ 8

Polyps few (<40), arranged sparsely in clusters of 2—4 at intervals along

rachis. Calyces and sclerites not apparents..554 554-5 450 iy

Polyp pairs 10-20. Intervals between polyp pairs <10 mm .. Scleroptilidae

Polyp clusters <5. Intervals between clusters of 1-4 polyps is >20 mm

pbug oie Reet ow 4 le tee ee ee See eee Chunellidae

Calyxawithvercht terminaliteeth( eee ne eee Funiculinidae

Calyx with 0—6.tenminalitcethy anaes ene ee eee Protoptilidae

Polyps non-retractile, calyces absent. Adjacent polyps united at their bases

only, not forming polyp leaves or raised ridges........... Anthoptilidae

Polyps retractile into calyces or into margins of fleshy polyp leaves or raised

TUDES istic is whe peal see nade oe ee 10

Polyps have tubular calyces with eight terminal teeth ...... Pennatulidae

Polyps have calyces with two terminal teeth, or calyces absent....... i

Polyp leaves stiffened with large white needle-like sclerites up to 10 mm

that formiprojectineysuppentingenays are eee Pteroeididae

Polyp leaves soft without sclerites, or polyps arranged in raised ridges with

calyces having) tworterminalatee thie acer Virgulariidae

KEY TO THE SPECIES OF PENNATULACEA FROM SOUTHERN AFRICA

Adjacent polyps free, not united to any degree. Individual polyps arise

directly from: the surface of the rachis) (e3e4 Fig tA) 2 ane Z

Proximal portions of adjacent polyps fused to some degree, often forming

flattened expansions (polyp leaves, e.g. Fig. 32A, D), raised ridges or swellings

that emanate laterally from the rachis (e.g. Fig. 30B, D), or joined only at their

bases with major portions of polyps free (e.g. Fig. 17B).............. 20

Polyps crowded and distributed evenly on all sides of rachis, or rachis with

a single longitudinal furrow or elongate V-shaped region devoid of polyps.

Colony radially symmetrical or somewhat bilaterally symmetrical, capitate,

clavate, or, Gigitiform. «44. se. cent ores ee ee ee 3

Polyps restricted to a limited portion of rachis, arranged biserially along

rachis, or concentrated in a terminal cluster, or in groups of two to four

distributed! sparsely, along machisi= 442 4542 oe oe ne ee eee 11

Polyps. without:calyces: c.f ye an See ease Sa oe +

Polyps with non=retractile biturcated (calyices 4254.) .5- 5000 ooo 9

Sclerites are smooth ovals, rods that are bilobed or branched at the ends, or

spindles up to 0,4 mm in length. Colonies clavate or capitate ......... 5

Sclerites are small flat plates (<0,1 mm in length) medially constricted with

rounded ends or with terminal denticles. Colonies cylindrical to elongate

ClAV ALE: 2854) Scie Re RN Ee BAS IE 7

Rachis sclerites are primarily rods with bilobed or branched ends

Te er Pn Mes ies sa ae OB Ordos So ad 5:2 Cavernulina sp.

SB.

6A.

6B.

WES

73%

8A.

SB.

9A.

OB.

10A.

10B.

11A.

11B.

12A.

12B.

13A.

13B.

14A.

14B.

ISA.

1SB.

16A.

SOUTHERN AFRICA PENNATULACEA 37

Rachis sclerites are ovals or spindles that are usually not bilobed or

ip TeeATNC NS Gl ie es ee at Pe ce eo es tahoe ad al A Naa danse Pu Or ered UAC ke ds 6

Internal axis absent. Sclerites are mostly spindles... Cavernularia elegans

Internal axis present. Sclerites are mostly ovals....... Cavernularia dayi

Colonies elongate clavate. Sclerites are small biscuit-shaped plates, many

Wwithecerminal GEmticleS 20g jccn 5c cians Stas yo Lituaria valenciennesi

Colonies cylindrical or elongate clavate. Sclerites are flat plates with medial

Consinictions andi smooth, roundediends...2..4.+.2..-5.555055-.-..- 8

Colonies plump and cylindrical with stalk shorter than rachis. Siphono-

zooids aligned in longitudinal rows. Colour yellowish to orange-yellow

56 0.0.8.0 9 B16 80.0 Ne een tet Oe Se nan ar a ge ace Veretillum cynomorium

Colonies long and narrow, clavate, with stalk equal to or longer than rachis.

Siphonozooids not aligned in longitudinal rows. Colour cream to greyish-

WHIGIRS c's oA eee gin tg ae they Sen rer ee ena ane ae eee Veretillum leloupi

Colonies radially symmetrical; polyps evenly distributed and covering all

sides of rachis. Calyces often not conspicuous with unaided vision. Colonies

usually straight or sometimes curved............... Actinoptilum molle

Colonies bilaterally symmetrical; rachis curved, concave side with longitu-

dinal furrow or V-shaped region near base of rachis free of polyps. Calyces

Conspicuous) givins colony a spiny appearance ..................... 10

Calyx length approximately three times the width. Calyx teeth long and

thin, lanceolate. Peduncle sclerites are small ovals 0,04-0,15 mm in

length. Colour of rachis often brick-red or deep reddish-purple

ee ee cree on vats ech ale nnd Oa igaeees Echinoptilum echinatum

Calyx length approximately twice the width. Calyx teeth conical or deltoid.

Peduncle sclerites are rods or spindles 0,07—0,47 mm in length. Colour of

rachis) Orange to reddish-brown. |... 24:5.-..- Echinoptilum macintoshii

Polyps arranged in a terminal cluster at the distal end of a long naked

DECINACKS 9:9 6 a:& ovBsh G4 CRO AOae een ee ee ear ESP ar oe 12

Roly ps anrangedsalone the sides of the rachis....0. 2. 4-0. 24-52.5.55 4. 13

Polyps and distal portion of stalk with sclerites ..... Umbellula thomsoni

Colony lacking sclenites altogether... 2. 2.5.-..5... Umbellula lindahli

Colonies clavate, distal portion of rachis wider than rest of colony; terminal

end obtuse. Polyps large (>10 mm), non-retractile, without calyces .. 14

Colonies not clavate, but are long, thin and uniform. Distal portion of rachis

NOtwiderthaminest Of COLONY i604 ae Hs el eek PR we 5

Rolyps Zon smear tenminalend Of rachis... 5. ...-- Kophobelemnon sp.

Polyps numerous and arranged biserially along rachis ................

a 6'o oo Bhacs bad Cha Sich hee a Ree Kophobelemnon stelliferum

Polyps numerous (usually >40) and contiguous, arranged along rachis in

two longitudinal series. Polyps retractile into distinct calyces......... 16

Polyps few (<40), arranged sparsely in clusters or whorls of 2—4 at intervals

AM OMemiaciisMiG@alyCesADSEME Me ic don wad Aliete yds ce alle elena dee ee LSS 18

Calyxaqwithie2=—omenminal tect a... . oc. y<sete oe be Seana ee 17

18B.

19A.

19B.

20A.

20B.

DANS

21B.

22A.

Dig) Bye

ip) 8).

24A.

24B.

ZA.

ZB:

ANNALS OF THE SOUTH AFRICAN MUSEUM

. Calyxowithout temminallitecthe sane ers nee eee eee Scleroptilum sp.

@alyxowithtereht tenninalitcethGanre ser see Funiculina quadrangularis

, Calyx usually with)2—6 terminaliteeth eee Distichoptilum gracile

. Polyps number 20-40, generally arranged in 10-20 pairs with 5-7 mm

intervals of bare rachis between each pair. Colonies very delicate, usually

<120-mniin leneth et eet a eee Calibelemnon sp.

Polyps fewer than 15. Generally <7 polyp clusters per colony with intervals

between clusters >20 mm. Mature colonies >120 mm in length...... 19

Polyps of rachis arranged in pairs. Distal end of colony with a single,

well-developed, somewhat asymmetrical terminal polyp................

ee a le Nees ee hee eG ic od Amphiacme abyssorum

Polyps of rachis in clusters of one to four, but usually three. Distal end of

colony with terminal polyp highly reduced or absent .. Chunella gracillima

Polyps non-retractile, without calyces. Sclerites not evident. Adjacent

polyps united at their bases only, major portion of polyps are free ......

ry oe ee eR LN re ORM, Sh Anthoptilum grandiflorum

Polyps retractile into conspicuous calyces or into margins of fleshy polyp

leaves. Fused proximal portions of adjacent polyps form prominent polyp

leaves, ridges, or basal swellings which emanate laterally from rachis in two

longitudinal series... os. ae eos ee ee 21

Polyps arranged in short oblique rows of four to six polyps in two

longitudinal series. Adjacent polyps united at their bases only, often

forming a common raised ridge or basal swelling. Polyps possess calyces that

each havestworterninalatce thine seer eee Halipterus africana

Adjacent polyps united most of their length to form distinct and flattened

lateral expansions (polyp leaves) that emanate from the rachis....... DD

Colonies lack visible sclerites (sclerites too small to be seen with unaided

eye). Peduncles usually thin and flexible or limp and worm-like...... 73

Colonies with numerous and conspicuous sclerites (sclerites easily seen with

unaided eye), Reduncles thick andsnieid 3. 26

Polyps usually 5-12 per leaf. Polyp leaves sparsely arranged, with

conspicuous intervals of bare rachis between leaves ................ 24

Polyps >12 per leaf. Polyp leaves densely situated, often overlapping one

another with little or no interval of bare rachis between leaves....... 25

Polyps never more than five per leaf. Peduncle somewhat stiff. Adjacent

polyps of a particular leaf gradually become larger from the innermost to

the outermost polyp. Gradual transition between peduncle and rachis

EMT ee ee Bat shee 5 ett Sth sch is een 0) tals Scytaliopsis djiboutiensis

Polyps 5-12 per leaf. Peduncle long and thin, vermiform. All polyps of a

given leaf approximately same size. Peduncle and rachis abruptly delimited

ee ST oS icin oa ny nS OG 8.4 8 Virgularia mirabilis

Polyps 50—200 pemleaii eee noe Virgularia gustaviana

Polyps 15=35:perdéatee sean ee ee eee ee Virgularia schultzei

SOUTHERN AFRICA PENNATULACEA 39

26A. Polyps possess tubular calyces with eight terminal teeth and dense sclerites

of three-flanged spindles. Sclerites usually orange in colour.............

ra a8 abet inciy Ue eee Maan as eA ake, nome ine ae Pennatula inflata

26B. Polyps retract directly into outer margin of polyp leaf. Sclerites of polyp

leaves are white or transparent spine-like needles up to 10 mm in length that

align to form projecting supporting rays. Sclerites not three-flanged. Colour

of colonies pale yellowish-grey or brownish-grey..... Pteroeides isosceles

LIST OF THE PENNATULACEA RECORDED FROM SOUTHERN AFRICA

Order PENNATULACEA Verrill, 1865

Suborder SESSILIFLORAE Kiikenthal, 1915

Family Veretillidae

* Lituaria valenciennesi D’Hondt, 1984

Cavernulina sp.

* Veretillum cynomorum (Pallas, 1766)

* Veretillum leloupi Tixier-Durivault, 1960

Cavernularia dayi Tixier-Durivault, 1954

Cavernularia elegans (Herklots, 1858)

Family Echinoptilidae

Echinoptilum macintoshii Hubrecht, 1885

Echinoptilum echinatum (Kikenthal, 1910)

Actinoptilum molle (Kukenthal, 1910)

Family Kophobelemnidae

Kophobelemnon stelliferum (Miller, 1776)

Kophobelemnon sp.

Family Anthoptilidae

Anthoptilum grandiflorum (Verrill, 1879)

Family Funiculinidae

* Funiculina quadrangularis (Pallas, 1766)

Family Protoptilidae

Distichoptilum gracile Verrill, 1882

Family Scleroptilidae

Scleroptilum sp.

Calibelemnon sp.

* Literature records only; material not available for examination.

40 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Chunellidae

Chunella gracillima Kukenthal, 1902

Amphiacme abyssorum (Kikenthal, 1902)

Family Umbellulidae

Umbellula thomsoni Kolliker, 1874

Umbellula lindahli Kolliker, 1874

Suborder SUBSELLIFLORAE Kiikenthal, 1915

Family Virgulariidae

Subfamily Virgulariinae

Virgularia schultzei Kikenthal, 1910

Virgularia mirabilis (Muller, 1776)

Virgularia gustaviana (Herklots, 1863)

Virgularia sp.

Scytaliopsis djiboutiensis Gravier, 1906

Subfamily Balticininae

Halipteris africana (Studer, 1879)

Halipteris spp.

Family Pennatulidae

Pennatula inflata Kikenthal, 1910

Family Pteroeididae

Pteroeides isosceles Thomson, 1915

Pteroeides spp.

DESCRIPTIONS

Family Veretillidae Herklots, 1858

Colonies radially symmetrical without a trace of external bilateral sym-

metry. Colony cylindrical, clavate (club-shaped), or capitate (forming a head).

Autozooids without calyces, completely retractile. Siphonozooids numerous

between the autozooids. Axis developed to varying degrees: conspicuous, minute,

or absent. Sclerites are plates, platelets, ovals, rods or spindles; none are

three-flanged.

Distribution widespread in shallow tropical and subtropical seas, mostly from

the Atlantic and Indo-Pacific, shallow sublittoral to approximately 190 m in

depth. Five genera, four of which are included below, in addition to Pollicella

Gray, 1870.

SOUTHERN AFRICA PENNATULACEA 41

Genus Cavernularia Valenciennes in Milne Edwards & Haime, 1850

Veretillum: Philippi, 1835: 277. Gray, 1870: 28.

Cavernularia Valenciennes in Milne Edwards & Haime, 1850: 84. Kukenthal & Broch, 1911: 180

(Lit.!). Kukenthal, 1915: 12.

Sarcobelemnon Herklots, 1858: 25.

Stylobelemnon Kolliker, 1872a: 336. Balss, 1910: 79.

?Fusticularia Simpson, 1905: 561. Balss, 1910: 80.

?Parabelemnon J. A. Thomson & Simpson, 1909: 307.

Diagnosis

Colonies cylindrical, clavate or capitate. Sclerites are smooth ovals, rods or

spindles; many are irregular in shape. Siphonozooids numerous and minute

(<0,3 mm in diameter). Autozooids with or without sclerites. Axis variable; large

to very small or absent. Colour usually white to cream, yellowish or greyish-white.

A genus of approximately 14 species. Indo-Pacific, west coast of Africa,

Mediterranean Sea, and Bay of Biscay.

Type species. Cavernularia obesa Valenciennnes in Milne Edwards & Haime,

1850; Indian Ocean.

Cavernularia dayi Tixier-Durivault, 1954

FIGS Z, Bo dks

Cavernularia Dayi Tixier-Durivault, 1954: 626, fig. 1.

Material

SAM-H3594 and SAM-—H3595, off East London (33°02'S 27°56’E),

25-30 m, 16 July 1984, 6 specimens, dredge; G. C. Williams, R.V. Meiring

Naude. SAM-—H3817 and H3818, off Umlaas River, Natal (29°58'S 31°01’'E),

50 m, 10 July 1985, 2 colonies, dredge; G. C. Williams, R.V. Meiring Naude.

Description

The colonies examined include three complete colonies, 23 mm, 30 mm and

70 mm in length, and five larger colonies, which have the proximal portions of the

peduncles missing. These partial colonies are between 30 mm and 62 mm in

length. Colonies club-shaped with tapering peduncle, length of which is 0,9-1,8

times the rachis length. Rachis conspicuously wider than peduncle. Autozooids

concentrated on distal half of rachis; capable of complete retraction into rachis.

Retracted autozooids less than 1,5 mm in diameter. Siphonozooids numerous,

minute, c. 0,16 mm in diameter. Siphonozooids may extend down proximal

portion of rachis, but are not conspicuous. Axis is a short rod (c. 8 mm long and

0,8 mm wide) in the centre of the proximal portion of the rachis. It is roughly

circular in cross-section. The surface of the axis may be partially impregnated with

oval sclerites. Axis length is less than 15 per cent of total colony length.

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 2. Cavernularia dayi. A. Entire colony. Scale = 5mm. B. Single autozooid showing

placement of sclerites; polyp length 2,5 mm. C. Entire colony. Scale = 6mm. D. Entire

colony. Scale = 20 mm. E. Longitudinal section of a colony showing position of axis. Scale =

20 mm. F. Axis, length 8,0 mm. G. Cross section of axis, diameter 0,8 mm. H. Sclerites from

surface of rachis. I. Sclerites from interior of rachis. J. Sclerites from surface of peduncle.

K. Sclerites from interior of peduncle. Scale: H-K = 0,15 mm.

Sclerites are mostly small ovals, or elongate rods, usually 0,12 mm or less in

length. They are present both in the rachis exterior and interior, as well as the

peduncle exterior and interior. Some ovals may have a cross-pattern originating

in the centre. Some sclerites of the rachis may be constricted in the middle with

rounded or truncate ends. Sclerites are densely distributed, and are more or less

consistent in shape throughout the colony. They vary from 0,02—0,13 mm in

length. Autozooids may have a few scattered sclerites at the bases, extending up

to less than one-third total length of polyp. The tentacles have many rod-like

sclerites, 0,06 mm in length.

SOUTHERN AFRICA PENNATULACEA 43

Fig. 3. Cavernularia dayi. A. Entire colony, length 30 mm. B. Partial colony, length 60 mm;

peduncle absent. C. Entire colony, length 23 mm. D-I. Scanning electron micrographs of

sclerites. D-—F. From surface of peduncle. D. 0,04 mm. E. 0,03 mm. F. 0,03 mm.

G-I. From surface of rachis. G. 0,04 mm. H. 0,03 mm. I. 0,07 mm.

44 ANNALS OF THE SOUTH AFRICAN MUSEUM

Colour greyish-white in life. Neck zones of autozooids may have eight

longitudinal subcutaneous regions of deep purple pigmentation, which is

conserved in alcohol.

Distribution (Fig. 38)

East London to the northern Natal coast, 25-50 m depth. Tixier-Durivault

(1954: 626) described the type locality from off Lake St. Lucia, Natal at 27 m

depth. The present study extends the range from the type locality south-west to

the eastern Cape Province, and north to Hully Point, Zululand. This species is

possibly a southern African endemic but many more records are necessary to

accurately establish its geographical distribution.

Remarks

Cavernularia dayi is distinguished from other southern African species of

Cavernularia by its clavate shape, small axis, and short, robust, oval or rod-like

sclerites of the rachis and peduncle.

Cavernularia elegans (Herklots, 1858)

Figs 4, 40

Sarcobelemnon elegans Herklots, 1858: 25, pl. 7 (fig. 3).

Veretillum elegans Richiardi, 1869: 127, pl. 13 (fig. 112).

Cavernularia elegans Kolliker 1872a: 165, pl. 23 (figs 207-208). Kikenthal & Broch, 1911: 185,

text-figs 13-15, pl. 18 (fig. 29). Kiikenthal, 1915: 16, fig. 20. Broch, 1958: 249.

Cavernularia elegans (non Herklots, 1858) Hickson, 1900: 89. J. A. Thomson & Simpson, 1909:

303. Balss, 1910: 86, figs 26-28.

Material

SAM-—H3871, 67 km west of Cape Fria, Namibia (18°25'S 11°15’E), 274 m,

May-June 1982, 9 colonies, trawl; collector unidentified.

Description

Colonies examined are 29-58 mm in length. Axis absent. Colonies clavate,

with peduncle as long as the rachis or somewhat longer. Autozooids usually

c. 1,3 mm in diameter. Siphonozooids conspicuous and numerous, in more or

less longitudinal rows between autozooids, 0,1 mm in diameter. Peduncle slightly

swollen near junction of rachis, with longitudinal striations on the external

surface. Sclerites from base of autozooids as well as the surface of the rachis and

peduncle are smooth elongate rods and spindles, 0,30—0,42 mm long. These are

not three-flanged and may have bluntly rounded or somewhat pointed ends. In

the rachis interior, similar sclerites are common; 0,2—0,4 mm long. The interior of

the peduncle contains minute ovals, 0,01 mm in length.

Colour yellowish-white with some brownish pigmentation in the polyps

below the tentacles.

SOUTHERN AFRICA PENNATULACEA

B. Sclerites from surface

Sclerites from surface of peduncle.

terior of peduncle. Scale = 0,01 mm

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46 ANNALS OF THE SOUTH AFRICAN MUSEUM

Distribution (Fig. 40)

Along the west African coast from Morocco to northern Namibia. The

present study extends the range south from near Luanda, Angola, to the Skeleton

Coast of Namibia, and represents a new record for southern Africa. The type

locality is an unspecified station off the west African coast.

Remarks

One colony of Cavernularia elegans (Herklots, 1858) was recorded by Broch

(1958: 249) from a depth of 65m off the coast of Luanda, Angola. Tixier-

Durivault (1963: 72) recorded one specimen as C. pusilla (Philippi, 1835)

from Lobito Bay, Angola, at 35m depth. These two species, together with

C. madeirensis Studer, 1878, were considered synonymous by Broch (1958: 250),

yet he retained C. elegans (instead of C. pusilla) as the correct binomial. I here

consider C. elegans and C. pusilla as separate. Kikenthal (1915: 13) maintained

that C. pusilla has irregular rod-shaped sclerites in the peduncle surface, whereas

C. elegans has spindles. Balss (1910: 86) repeated Hickson’s (1900: 89) misidenti-

fication of Actinoptilum molle as C. elegans, from 45 m depth off the Cape of Good

Hope. Records of C. elegans from Japan (Balss 1910) and the Indian Ocean (J. A.

Thomson & Simpson 1909) were considered dubious by Kikenthal (1915: 16).

Cavernularia elegans is characterized by Ktikenthal (1915: 13, 16) as having dense

spindles at the surface of the peduncle, whereas these sclerites are absent in the

peduncle interior; axis absent; sclerites from the surfaces of the peduncle and rachis

are predominantly elongate rods and spindles. The rachis interior has similar

sclerites but the peduncle interior has minute ovals. Cavernularia obesa, from the

Indo-West Pacific, also lacks an axis, but possesses sclerites in the interior of the

peduncle that are similar to those from the surface (according to Ktikenthal, 1915:

13; also see D’Hondt 19845: 634). Cavernularia elegans is distinguished from other

southern African species of the genus by its clavate growth form, absence of an axis,

and sclerites of elongate rods and spindles.

Remarks on the genus Cavernularia

Day et al. (1970: 17) listed Cavernularia luetkeni Kolliker, 1872, from

80-102 m depth in False Bay, near Cape Town. Since the specimen upon which

this identification was made cannot be traced and since no description or figure is

provided, it must be considered a dubious record. A 19-mm long specimen

collected at Algoa Bay, near Port Elizabeth (SAM-—H3236), was originally

identified by Tixier-Durivault as C. luetkeni. I have examined this specimen and

found the polyps to contain small bifurcated calyces. I therefore identify it as a

young colony of Actinoptilum molle (Kiikenthal, 1910). Tixier-Durivault (1960:

366) listed C. luetkeni from Inhaca Island, southern Mozambique. Material of

C. luetkeni is not available at present for examination. Kiikenthal (1915: 14)

distinguished the species by the possession of a small axis in the base of the rachis

and top part of the peduncle; stalk sclerites <0,19 mm long; and rachis sclerites

<0,31 mm long.

SOUTHERN AFRICA PENNATULACEA 47

Approximately 18 species of the genus have been named. Of these, four were

considered dubious species or synonyms by Kikenthal (1915). Most of the

recognized species are from the eastern Atlantic or Indo-West Pacific, and one is

from the Pacific coast of Central America and the Galapagos Archipelago. The

genus is in need of revision, since clear distinctions between some species are not

evident from the literature. A comparison of type material is necessary to

establish the validity of certain members of the genus.

Other species of the family Veretillidae

Four additional veretillid species have been recorded from southern Africa,

but for three of these material is not presently available to the author for

examination, and in one species only partial material is available.

Genus Veretillum Cuvier, 1798

Pennatula: Pallas, 1766: 177 (partim).

Alcyonium Linnaeus, 1767: 342 (partim).

Veretillum Cuvier, 1798: 675. Herklots, 1858: 26. Gray, 1870: 33. Kolliker, 1872a: 324. Balss,

1910: 79. Kiikenthal & Broch, 1911: 175. Kiikenthal, 1915: 11.

Diagnosis

Colonies cylindrical and plump or elongate and clavate. Size of axis variable, or

altogether absent. Autozooids are distributed over surface of rachis; siphono-

zooids are contained in longitudinal rows between the autozooids. Sclerites are small

plates (<0,1 mm), unbranched, often bone-shaped, biscuit-shaped, or at least

constricted in the middle to some degree.

Three species from southern Europe, west coast of Africa, southern

Mozambique and the Malay Archipelago.

Type species. Pennatula cynomorium Pallas, 1766; Europe.

Veretillum cynomorium (Pallas, 1766)

Figs 38, 40

Pennatula cynomorium Pallas, 1766: 177.

Alcyonium epipetrum Linnaeus, 1767: 1294.

Veretillum cynomorium Kikenthal & Broch, 1911: 176, pl. 13 (fig. 1), pl. 18 (figs 32-36) (Lit.!).

Kukenthal, 1915: 12, figs 12, 13. Broch, 1958: 249. Tixier-Durivault, 1960: 362.

Material

Material was not available for examination.

Remarks

According to Kiikenthal (1915: 11-12) colonies are cylindrical and plump,

sausage-shaped, with rachis longer than peduncle. Siphonozooids in longitudinal

rows. Sclerites are thin flat plates (<0,06 mm), usually medially constricted with

ends rounded.

48 ANNALS OF THE SOUTH AFRICAN MUSEUM

Broch (1958: 249-250) recorded this species from the Atlantic and Medi-

terranean coasts of southern Europe, and along the west African coast as far south

as Walvis Bay, Namibia; the latter record follows Ktikenthal & Broch (1911: 177).

Molander (1929: 15) extended the range south to the mouth of the Orange River

(border of Namibia and South Africa), 13 m depth. Tixier-Durivault (1960: 362)

recorded a colony from Inhaca Island, southern Mozambique. The known depth

range is 13-91 m.

Veretillum leloupi Tixier-Durivault, 1960

Fig. 38

Veretillum leloupi Tixier-Durivault, 1960: 363, figs 1-2.

Material

Material was not available for examination.

Remarks

According to Tixier-Durivault (1960: 363-365), colonies are long and nar-

row, clavate with stalk length equal to or longer than rachis. Siphonozooids not

aligned in longitudinal rows. Sclerites are bone-shaped, not flattened, with medial

constrictions and rounded ends, up to 0,1 mm in length.

The species is known only from the type locality at Inhaca Island, southern

Mozambique, in the western Indian Ocean; depth not recorded.

Genus Cavernulina Kikenthal & Broch, 1911

Cavernularia J. A. Thompson & Simpson, 1909: 302 (partim).

Cavernulina Kiikenthal & Broch, 1911: 172. Kiikenthal, 1915: 9.

Diagnosis

Colonies clavate. Axis present. Sclerites of the rachis are mainly rods with

bilobed or branched ends, usually <0,4 mm in length. Peduncle sclerites are

smooth rods and spindles, most of which are not bilobed at the ends, and are

mostly <0,2 mm in length. Polyps without sclerites.

Three species from the Indo-West Pacific.

Type species. Cavernularia cylindrica Kikenthal & Broch, 1911; Malay

Archipelago.

Remarks

Cavernulina cylindrica Kiikenthal & Broch, 1911, was listed without

description or figures by Day et al. (1970: 17) from 33-36 m in False Bay, near

Cape Town, as Cavernularia cylindrica. Since the specimen upon which this

record was based is not available for examination and since this is the only False

Bay record of the species known, it must be considered a dubious record. It is

probable that small specimens from the Cape region, previously identified as

SOUTHERN AFRICA PENNATULACEA 49

species of Cavernularia, may in fact be young colonies of Actinoptilum molle in

which the bifurcate calyces are neither prominent nor conspicuous, but present

nonetheless. Two species were considered valid by Ktikenthal (1915: 9): Cavern-

ulina cylindrica and C. orientalis (Thomson & Simpson, 1909), both from the

Indian Ocean. A third species, C. grandiflora D’Hondt, 1984, has recently been

described from New Caledonia. See D’Hondt (19846: 629) for a comparison of

these three species.

Cavernulina sp.

Fig. 38

Remarks

A partial colony, in which only a 30 mm long rachis is present, recently

collected from 62 m depth off the Durban region of Natal, possesses bilobed

sclerites 0,1—0,25 mm in length. These sclerites are similar in shape and size to

those figured by Kiikenthal (1915: 10) for C. cylindrica and by D’Hondt (19845:

628) for C. grandiflora. | therefore believe that this specimen belongs to the genus

Cavernulina, but a proper identification to species is not possible due to the

fragmentary nature of the material, in which the peduncle is entirely missing. The

present study establishes the genus Cavernulina as a new record for southern

Africa.

Genus Lituaria Valenciennes in Milne Edwards & Haime, 1850

Pennatula: Pallas, 1766: 179 (partim).

Veretillum: Lamarck, 1816: 420 (partim).

Lituaria Valenciennes in Milne Edwards & Haime, 1850: 84. Gray, 1870: 33. Kélliker, 1872a:

135. J. A. Thomson & Simpson, 1909: 311. Balss, 1910: 78. Kiikenthal, 1915: 7.

Clavella Gray, 1870: 33. K6élliker, 1872a: 144. J. A. Thomson & Simpson, 1909: 311. Balss,

1910: 78.

Diagnosis

Colonies clavate. Axis contained in rachis, length variable, four-angled.

Siphonozooids very numerous between the autozooids. Sclerites are small

biscuit-shaped plates, many of which possess short branches or denticles at the

ends. Other sclerites may be crosses or rods with warty ends.

Nine species of the Indo-Pacific. See Light (1921: 248) for a key to the

species, and D’Hondt (1984b: 636) for a discussion of the type species.

Type species. Pennatula phalloides Pallas, 1766; Indian Ocean.

Lituaria valenciennesi D’Hondt, 19846

Lituaria phalloides (non Pallas, 1766) Valenciennes in Milne Edwards & Haime, 1850: 84.

Kolliker, 1872a: 313. Ktkenthal, 1915: 8, fig. 6.

Lituaria valenciennesi D’Hondt, 1984b: 636, fig. 9.

Material

Material was not available for examination.

50 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

According to Kukenthal (1915: 8), colonies are clavate with rounded distal

ends. Polyps mostly without sclerites. Sclerites of the rachis and stalk are small flat

plates up to 0,1 mm long, many with terminal denticles.

The species is recorded from Mozambique (undesignated station), the Malay

Archipelago, the Andaman Islands, and Sri Lanka; shallow sublittoral.

Family Echinoptilidae Hubrecht, 1885

Colony digitiform or cylindrical. Axis absent. Rachis generally longer than

peduncle. Colonies radially (Fig. 11B) to bilaterally symmetrical (Fig. 9B).

Autozooids with distinctive, non-retractile, bifurcated calyces that are heavily

armed with sclerites (Fig. 9A, E). Sclerites mainly smooth three-flanged rods or

spindles, ovals and platelets (Fig. 13).

Two genera from the Indo-West Pacific to south-western Africa; shallow

sublittoral to 835 m.

Genus Echinoptilum Hubrecht, 1885

Echinoptilum Hubrecht, 1885: 512. Balss, 1910: 37. Kikenthal & Broch, 1911: 196. Kikenthal,

1915: 18.

Actinoptilon Kikenthal, 1910: 54 (partim).

Diagnosis

Colony digitiform, commonly curved. Concave side with a longitudinal

groove or furrow, and inverted V-shaped region at the base of the rachis devoid

of polyps. This furrow may extend full length of rachis in small specimens (less

than 40 mm). Proximal portion of rachis therefore bilaterally symmetrical. Distal

portion of rachis may be radially symmetrical with polyp-bearing region con-

tinuous in larger specimens (over 40 mm). Autozooids with prominent bifur-

cated calyces, giving rachis a spiny or prickly appearance. Two terminal teeth of

calyx deltoid to needle-like. Larger autozooids may be concentrated in two lateral

series of longitudinal rows in colonies less than 40 mm long. Autozooids capable of

total retraction into calyces. Siphonozooids numerous at base of autozooids; also

with non-retractile, bifurcated calyces. Sclerites numerous; those of calyx are long

three-flanged spindles, peduncle with ovals or three-flanged rods and spindles.

A genus of six species, Indo-West Pacific— Hawaii to eastern Africa.

Type species. Echinoptilum macintoshii Hubrecht, 1885; Japan.

Echinoptilum macintoshii Hubrecht, 1885

Figs 1H-I, 5, 6, 7A—C, 8D-F, 39

Echinoptilum macintoshii Hubrecht, 1885: 512, text-figs 1-3, pl. 30 (figs 1-8), pl. 31 (figs 9-14).

Echinoptilum macintoshi: Nutting, 1908: 561; 1912: 48. Balss, 1910: 38, pl. 2 (figs 1-3), pl. 5

(figs 1, 3-5). Kikenthal, 1915: 19. Utinomi, 1961: 221.

Echinoptilum Macintoshii: Kikenthal & Broch, 1911: 197.

Echinoptilum M’Intoshii: Hickson, 1916: 57-62.

Echinoptilum mackintoshi: J. A. Thomson & Rennet, 1927: 121, pl. 9 (fig. F). Bayer, 1956: 226.

SOUTHERN AFRICA PENNATULACEA ai

Material

SAM-H3683, off Mbotyi, Transkei (31°32’S 29°50’E), 150-170 m, August

1981, 2 colonies, dredge; Natal Museum, R.V. Meiring Naude. SAM—H3685, off

Port St. Johns, Transkei (31°39’S 29°35’E), 50-70 m, 14 August 1981, 1 colony,

dredge; Natal Museum, R.V. Meiring Naude. SAM-—H3819, off Mgazi River,

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Oe y AVA

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A mn Wi Hl 2 $2 48

Waal File ee Sch Eee

al dtd dhe ge ue as

ve ieee

PAY ‘va

PEE 6 8 oo Doers

ate DEE eS a

Fig. 5. Echinoptilum macintoshii. A. Entire colony, lateral view. Scale = 5mm. B. Five

autozooids with eight siphonozooids; sclerites shown in calyx of top autozooid only.

Scale = 1mm. C. Single tentacle (length 1,3 mm) showing alignment of sclerites.

52 ANNALS OF THE SOUTH AFRICAN MUSEUM

Transkei (31°43,7'S 29°31,9'E), 140-145 m, 4 July 1985, 8 colonies, dredge;

G. C. Williams, R.V. Meiring Naude.

Description

Specimens examined are 10-84 mm in length. Colonies curved with ventral

groove on concave side. Calyx teeth conical to deltoid, 0,4—0,8 mm in length.

Calyx length approximately twice the width. Entire calyces 1,2—2,3 mm in length.

Siphonozooids have two broad teeth, often with obtuse tips.

Sclerites of the calyces are smooth three-flanged spindles 0,17—1,07 mm in

length (Fig. 6A); aboral surface of each tentacle with a conspicuous longitudinal

medial band of red-orange three-flanged rods, which extend into the base of the

Fig. 6. Echinoptilum macintoshii. A. Calyx sclerites. B. Sclerites from surface and interior of

rachis. C. Tentacular sclerites. D. Sclerites from surface of peduncle. E. Sclerites from

interior of peduncle. Scales: A-B, D (left) = 0,2 mm; C, E (right) = 0,1 mm.

SOUTHERN AFRICA PENNATULACEA

;

: :

:. | 4

i A Gt OR CR i esc ibe

Fig. 7. Scanning electron micrographs of sclerites. A-—C. Echinoptilum macintoshii. From

surface of peduncle: 0,33 mm; 0,31 mm; 0,16 mm.D-H. Echinoptilum echinatum. D-E. From

calyx: 0,17 mm; 0,55 mm. F-—H. From surface of peduncle: 0,11 mm; 0,11 mm; 0,09 mm.

54 ANNALS OF THE SOUTH AFRICAN MUSEUM

anthocodia. These are all arranged parallel to the axis of each tentacle and form

a band 2-3 sclerites in width. Similar but colourless rods are found at various

oblique angles to the axis, near the bases of the pinnules (Fig. 5C). The rods of

the tentacles vary in length from 0,13—0,17 mm (Fig. 6C). Bare areas of the rachis

surface, aS well as the rachis interior, possess three-flanged spindles and rods

0,13-0,50 mm in length (Fig. 6B). Sclerites from the surface of the peduncle are

longitudinally grooved (but not three-flanged) spindles or ovoid rods, 0,07-

0,47 mm in length, often with undulate margins and tapering ends (Figs 6A—C,

6D). The tissues of the peduncular interior possess scattered sclerites of minute

ovals 0,008—0,020 mm in length (Fig. 6E).

Rachis rust-orange to light reddish-brown. Peduncle and naked region of

rachis whitish or cream. Calyces reddish-brown, often with margins edged in

white. Colour of colony permanent, originating within the sclerites.

Distribution (Fig. 39)

Transkei coast, Japan and Hawaii. This study establishes a new record for the

African coast and extends the range of the species to the western Indian Ocean.

Colonies from south-eastern Africa are known from 50-170 m depth. Hubrecht

(1885: 512) reported the type locality as the Sea of Japan, 130 m in depth. Nutting

(1908: 561) recorded the species from the Hawaiian Archipelago, 225-258 m, and

again off Japan, 150 m (Nutting 1912: 48).

Remarks

Echinoptilum macintoshii is distinguished from other members of the genus

by having sclerites in the tentacles, and triangular calyx teeth <0,8 mm long.

Echinoptilum echinatum (Kikenthal, 1910)

Figs 1G, 7D—H, 8G-H, 9, 10, 39

Actinoptilon echinatum Kiukenthal, 1910: 57.

Echinoptilum echinatum Kikenthal & Broch, 1911: 197, pl. 13 (fig. 3), pl. 18 (figs 38, 39), pl. 19

(figs 40-43). Kikkenthal, 1915: 19 (fig. 21). Hickson, 1916: 57-62.

Material

SAM-—H1595, off Durban (30°20’S 30°51’E), 406 m, 23 July 1964, 1 colony,

dredge; Division of Sea Fisheries. SAM—H3680, off Mendu Point, Transkei

(32°23'S 29°00’E), 450 m, 13 July 1984, 1 colony; G. C. Williams, R.V. Meiring

Naude. SAM-H3681, off Qolora River, Transkei (32°48’S 28°37'E), 510 m,

14 July 1904, dredge; G. C. Williams, R.V. Meiring Naude. SAM—H3682, off

Qora River, Transkei (32°35’S 28°49’E), 450-460 m, 14 July 1984, dredge; G. C.

Williams, R.V. Meiring Naude. SAM—H3684, off Qora River, Transkei (32°34’S

28°50'E), 400-420 m, 12 July 1984, dredge; G.C. Williams, R.V. Meiring

Naude. SAM-—H3820, off Mbashe River, Transkei (32°23,6’S 28°59,2’E), 350 m

maximum, 6 July 1985, 1 colony, dredge; G. C. Williams, R.V. Meiring Naude.

SOUTHERN AFRICA PENNATULACEA 55

Description

Specimens examined are 30—80 mm in length. Colonies curved with ventral

groove on concave side. Calyx teeth lanceolate to needle-like, 1,0-1,6 mm in

length. Total calyx length (approximately 3 mm) is more than twice and often

three times the width. Siphonozooids possess two narrow teeth with acute tips.

Sclerites of the calyces are narrow, three-flanged spindles 0,14—0,86 mm long

(Figs 7D-E, 10A). The aboral surface of each tentacle has a medially placed

longitudinal row of 1-20 sclerites, arranged more-or-less parallel to the axis of the

tentacles (Fig. 9C-D). These are three-flanged rods 0,10—-0,16 mm in length

(Fig. 10C). Bare regions of the surface of the rachis, as well as the interior of the

rachis, possess elongate three-flanged rods 0,06—0,35 mm in length (Fig. 10B).

The peduncular surface sclerites are oval finger-biscuits with longitudinal striation

(but not three-flanged) 0,04—0,15 mm in length (Figs 7F—H, 10D). The tissues of

the interior of the peduncle possess minute ovals 0,009-0,014 mm in length

(Fig. 10E).

Calyces and rachis are brick red or deep reddish-purple to cream. Peduncle,

as well as the naked longitudinal track on the concave side of the rachis are

whitish to cream, often with reddish longitudinal lines. Colour permanent due to

pigmentation of the sclerites.

Distribution (Fig. 39)

Transkei and Natal coasts to Somalia. South-east African specimens from

350-510 m depth. Kiikenthal & Broch (1911: 197) reported the type locality as off

the Somalia coast at 628 m depth. The present study extends the range of the

species southward along the coast of east Africa, and represents a new record

from southern Africa.

The two south-east African species of Echinoptilum apparently show

divergent bathymetric ranges: E. macintoshii has a shallow-water distribution,

being encountered at depths of 130-258 m, whereas E. echinatum is presumably

a deeper-water form, known only between 350 m and 628 m in depth.

Remarks

Echinoptilum echinatum is distinguished from other species of the genus by

having sclerites in the tentacles, and lanceolate calyx teeth up to 1,6 mm long.

Remarks on the genus Echinoptilum

Four other species of Echinoptilum were described by Hickson (1916) from

the Malay Archipelago: E. elongatum, E. minimum, E. roseum, and E. asperum.

These species have not been recorded outside their respective type localities.

Hickson (1916: 62) presented a key to the six described species. Echinoptilum

macintoshii is characterized by having pronounced curvature, spicules in the

tentacles, and small calyxes, whereas E. echinatum is also described as being

curved with spicules in the tentacles, but with large calyces. The southern African

material agrees with Hickson’s descriptions of these two species, as well as the

56 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 8. A-C. Actinoptilum molle colonies. A. 70 mm. B. 65 mm. C. 85 . D-F. Echinoptilum

macintoshii. D.25 mm. E.45 mm. F. 35 mm. G-H. Echinoptilum echinatum. G. 40 mm.

H. 55 mm.

SOUTHERN AFRICA PENNATULACEA 57

SSS

~—:

Fig. 9. Echinoptilum echinatum. A. Entire colony, lateral view. Scale = 5mm. B. Ventral

view of colony showing naked tract. Scale = 5 mm. C-—D. Two tentacles, showing variation in

sclerite density and placement. Each tentacle is 2,0 mm in length. E. Single autozooid, partially

retracted; calyx sclerites shown, five siphonozooids at base. Scale = 1 mm. F. Single bifurcated

calyx of a retracted autozooid; sclerites omitted. Scale = 1 mm.

58 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 10. Echinoptilum echinatum. A. Calyx sclerites. B. Sclerites from surface and interior of

rachis. C. Tentacular sclerites. D. Sclerites from surface of peduncle. E. Sclerites from

interior of peduncle. Scales: A, B (left) = 0,2 mm; C, D, E (right) = 0,1 mm.

detailed descriptions of Hubrecht (1885: 512) and Kiikenthal & Broch (1911:

197).

The distinctions between the various groups may not be consistent or marked

enough to warrant specific separation. Hickson (1916: 58) raised the possibility

that the genus Echinoptilum is monospecific and that the six described species

may in fact represent one highly variable and widespread species. A comparative

examination of type specimens and other known material is needed. However,

the differences between E. macintoshii and E. echinatum, based upon compari-

son of the available southern African material, are marked and consistent.

Genus Actinoptilum Kikenthal, 1911

Cavernularia (non Valenciennes in Milne Edwards & Haime, 1850) Hickson, 1900: 89, 92.

Actinoptilon Kiikenthal, 1910: 56.

Actinoptilum Kikenthal & Broch, 1911: 201. Kiikenthal, 1915: 20.

Actinoptinum Day et al., 1970: 17.

SOUTHERN AFRICA PENNATULACEA 59

Diagnosis

Colony cylindrical with rachis radially symmetrical. Polyps evenly distributed

and completely covering surface of rachis. Calyces of autozooids usually less than

3 mm in length, bifurcate. Terminal teeth short and conical to deltoid or broadly

lanceolate and acute. Inside of each tooth often with a fleshy rounded pro-

tuberance near the apex.

A monospecific genus restricted to southern Africa and possibly Madagascar.

Type species. Actinoptilon molle Kikenthal, 1910; South Africa.

Actinoptilum molle (Kikenthal, 1910)

Figs 1A-C, 8A-C, 11-13, 39

Cavernularia obesa (non Valenciennes in Milne Edward & Haime, 1850) Hickson, 1900: 92, pl. 3

(figs A—B). |

Cavernularia elegans (non Herklots, 1858) Hickson, 1900: 89.

Actinoptilon molle Kiikenthal, 1910: 57 (partim). Tixier-Durivault, 1960: 366. Tixier-Durivault &

D’Hondt, 1974: 259.

Actinoptilum molle Kikenthal & Broch, 1911: 201, pl. 13 (figs 3-4), pl. 18 (fig. 37). Kiikenthal,

1915: 20, figs 22-24. J. S. Thomson, 1915: 3. 1924: 81. Hickson, 1916: 41. Molander, 1929:

15. Tixier-Durivault, 1954: 628. Broch, 1958b: 251.

Actinoptilum molle var. maculatum J. S. Thomson, 1915: 5.

Actinoptilum molle var. intermedium J. S. Thomson, 1915: 7.

Actinoptilum molle var. zonatum J. S. Thomson, 1915: 8.

Actinoptinum molle Day et al., 1970: 17.

Material

SAM-H3254, Port Elizabeth Harbour (33°58’S 25°38’E), 20—25 m, February

1984, 2 colonies, small dredge; W. R. Liltved. SAM—H3263, Algoa Bay (33°59'S

25°42'E), 20m, February 1984, 2 colonies, SCUBA; W. R. Liltved. SAM-

H3164, off Jeffrey’s Bay (34°00’S 25°10’E), 30m, August 1980, 6 colonies,

dredge; W. R. Liltved, R.V. T.B. Davie. SAM-—H3272, False Bay (34°09’S

18°27'E), 26 February 1984, 12m, 1 colony, small dredge; W. R. Liltved.

SAM-—H3339, Algoa Bay (33°59’S 25°42'E), 16m, 17 May 1984, 3 colonies,

SCUBA; G. C. Williams.

Description

Colonies examined range in length between 12 mm and 240 mm. Colony

cylindrical, plump, sausage-shaped. Peduncle thick, tapering gradually, usually

one-fifth to one-third total colony length. Rachis tapering gradually to a rounded

apex. Rachis entirely covered with dimorphic polyps; thus the polyp-bearing

surface of the rachis is radially arranged. Siphonozooids crowded between bases

of autozooids with minute bifurcated calyces. Rachis surface often with longi-

tudinal lines between columns of autozooids. Calyces of autozooids usually not

conspicuous, bifurcate, non-retractile. Autozooids capable of total retraction into

calyces. Calyx teeth variable in shape and length.

60 ANNALS OF THE SOUTH AFRICAN MUSEUM

a 2 Met

SEES SSF

BSR ACS S

6 X89

23 FRPRERIEFIERS oy = BS ese

APY PAA WVATWNHY_Y 7U® WINNT IIE LSS

ep ceed Ra Sree iabe tak saben apts

Fig. 11. Actinoptilum molle. A. Expanded colony in situ. Scale = 30 mm. B. Single autozooid

and five siphonozooids. C. Three autozooids retracted into calyces, and numerous siphono-

zooids. D. Two autozooid calyces with four smaller siphonozooids. Scales: B-D = 0,8 mm.

E. Expanded autozooid. Scale = 1,8 mm. F. Preserved colony with retracted polyps.

Scale = 10 mm.

SOUTHERN AFRICA PENNATULACEA 61

Sclerites of the calyx are smooth three-flanged spindles 0,18—0,85 mm long

(Figs 12A, 13A—D). Anthocodia (distal part of polyp that contains the tentacles,

mouth, and neck zone) with small ovals or platelets in neck zone and base of

tentacles, 0,02—0,04 mm in length (Figs 12B, 13E). The surface of the rachis has

three-flanged spindles and rods 0,22—0,42 mm in length (Figs 12C, 13F, H-I).

The tissues of the interior of the rachis possess similar sclerites 0,22—0,41 mm in

length (Fig. 12D). The surface of the peduncle has ovoid finger biscuit-shaped

sclerites with longitudinal surface striations 0,03-—0,20 mm long (Figs 12E, 13G).

The tissues of the peduncular interior possess minute ovals 0,005—0,020 mm in

length (Fig. 12F).

Po()O0

20860

09 AQO

Fig. 12. Actinoptilum molle. A. Calyx sclerites. B. Sclerites from anthocodial wall. C. Scle-

rites from rachis surface and interior. D. Sclerites from surface of peduncle. E. Sclerites from

interior of peduncle. Scales: A, C (left) = 0,2 mm; B, D (right) = 0,1 mm; E = 0,02 mm.

62 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 13. Scanning electron micrographs of sclerites of Actinoptilum molle. A-—D. From calyx:

0.41 mm; 0,21 mm; 0,45 mm; 0,44 mm. E. From tentacle: 0,04 mm. F,H,I. From surface of

rachis: 0,2 mm; 0,15 mm; 0,2 mm. G. From surface of peduncle: 0,07 mm.

SOUTHERN AFRICA PENNATULACEA | 63

Colour highly variable; individual colonies may be entirely yellow, white to

cream, reddish-brown, pinkish or mauve; or colonies with rachis deep-purple to

reddish-purple, and peduncle yellow, white, pinkish or brownish. Colour

permanent, originating within the sclerites.

Distribution (Fig. 39)

This species is presumably a southern African endemic, known from off Cape

Columbine on the west coast of South Africa to Inhaca Island, Mozambique, at

depths ranging from 12 m to 333 m. However, Tixier-Durivault & D’Hondt

(1974: 259) recorded, without description or figures, seven colonies from north-

eastern Madagascar between 200 m and 605 m in depth. The species is perhaps

the most frequently encountered pennatulacean in South African coastal waters.

Observations

A 60-mm long live colony from False Bay (SAM-—H3272), which was kept in

a bucket of sea-water at night, was observed to luminesce vigorously when

agitated; the rachis sparkled throughout with minute pinpoints of green light.

SCUBA divers have observed this species inhabiting rocky reefs off Cape

Agulhas at 48 m depth. A colony was observed in a small depression in rocky

substratum that was filled with fine sediment (W. R. Liltved pers. comm.). Divers

have also observed extensive sandy areas at 20 m depth in False Bay with a

density of 4—5 colonies per square metre (P. Williams pers. comm.).

Remarks

Thomson (1915: 1-13) named three varieties of Actinoptilum molle based

primarily on differences in external coloration: A. molle var. maculatum,

A. molle var. intermedium, and A. molle var. zonatum. Because of the wide

range of colour variability observed throughout the geographic extent of the

species and many gradations between various colonies or populations, I believe

that formal subspecies or varieties should not be designated. This species can be

regarded as yet another example of the extreme range of variability exhibited by

sedentary benthic organisms such as octocorallians. Thomson (1915: 8) observed

a slight trace of a groove on the rachis of four white colonies that he referred to

A. molle var. intermedium. A conspicuous groove or furrow on the concave side

of the rachis is considered diagnostic of the genus Echinoptilum. However,

according to Thomson, the four specimens show typical characteristics of

Actinoptilum, such as smaller inconspicuous calyces and numerous polyps

arranged almost completely radially, as well as the form, dimensions, and

distribution of sclerites being similar to those described by Kukenthal for

A. molle. Calyx length is also quite variable, ranging from 1 mm to 3 mm. In

some colonies the calyx teeth are only visible with the aid of a hand lens or

dissecting microscope, whereas in others they are very conspicuous and easily

seen with the unaided eye.

64 ANNALS OF THE SOUTH AFRICAN MUSEUM

Kiikenthal (1915: 20) stated that the polyps are without sclerites. The density

of sclerites in the anthocodiae is variable; in a single colony some anthocodiae

may contain sclerites, whereas others may contain none. The number of ovals

present in a particular anthocodia is also quite variable.

Actinoptilum molle is distinguished from other southern African sea pens by

having radial symmetry of the rachis, in which the polyps are distributed evenly

on all sides, and the possession of generally inconspicuous calyx teeth.

Family Kophobelemnidae Gray, 1860

Colonies clavate (club-shaped). Axis present. Polyps arranged bilaterally

along rachis; rachis may show a tendency toward radial symmetry. Autozooids

large, without calyces. Siphonozooids with calyces, at base of autozooids. Rachis

with sclerites of three-flanged or tuberculate spindles, or plate-like, biscuit-shaped

or bone-shaped rods.

Four genera from the Atlantic, Mediterranean and Indo-Pacific: Mesobelem-

non Gravier; 1907; Sclerobelemnon KO6lliker, 1872a; Kophobelemnon Asbjgrn-

sen, 1856; and Malacobelemnon Tixier-Durivault, 1966a. The present study

represents the first record of the family for southern Africa.

Genus Kophobelemnon Asbj@rnsen, 1856

Kophobelemnon Asbj@rnsen, 1856: 81. Kukenthal & Broch, 1911: 219. Kiikenthal, 1915: 29

(Lit.!).

Bathyptilum Kolliker, 1872: 200.

Gunneria Danielssen & Koren, 1884: 58.

Diagnosis

Colony clavate, elongate, stiff. Distal end slightly expanded and rounded,

knob-like, or slightly pointed at the tip. Rachis and stalk about equal in length.

Axis thin, often slightly quadrangular in transverse section. Sclerites are mostly

three-flanged spindles that may possess tubercles.

A genus of eight to ten species. Atlantic, Mediterranean, and Indo-Pacific;

mostly deep sea, 40—4 400 m.

Type species. Pennatula stellifera Muller, 1776; northern Atlantic.

Kophobelemnon stelliferum (Miller, 1776)

Figs 14-16, 40

Pennatula stellifera Miller, 1776: 255.

Kophobelemnon stelliferum Kikenthal & Broch, 1911: 224, figs 54-60. Kiikenthal, 1915: 29,

figs 40-41 (Lit.!).

Material

SAM-—H3707, off Skeleton Coast, Namibia (20°59’S 12°23'E), 486-605 m,

20 August 1982, 1 colony, bottom trawl; Sea Fisheries Research Institute, Crab

Survey. SAM-—H4026 off Skeleton Coast, Namibia (c. 21°00'S 12°00’E),

SOUTHERN AFRICA PENNATULACEA 65

nO a is Sis

ea aa PDP Aye

a =A _ 3 ¥ > Ss

crate a. ss weeteas a gtk = a

th aad -

PATO IG a ae BPR A

See

SVs.

Rea

ot. SS

WAIT ST

WN.

‘>

Th.

AA?

tm OLE

aie

Fig. 14. Kophobelemnon stelliferum. A. Entire colony. Scale = 50mm. B. Sclerites from

surface of rachis. C. Sclerites from wall of an autozooid. D. Sclerites from surface of peduncle.

Scale: B—D = 0,2 mm. E. Detail of single autozooid and seven siphonozooids. Scale = 8 mm.

ANNALS OF THE SOUTH AFRICAN MUSEUM

wien dba pipe:

Libre

iitpepiilliin

tists wsthjpuilelgey

dle

Fig. 15. Scanning electron micrographs of sclerites of Kophobelemnon stelliferum. A-—C. From

rachis: 0,32 mm; 0,33 mm; 0,59 mm. D. Tip of rachis sclerite: 0,08 mm. E—F. From peduncle:

0,26 mm; 0,24 mm. G. Tip of peduncle sclerite: 0,1 mm.

SOUTHERN AFRICA PENNATULACEA 67

Fig. 16. Kophobelemnon stelliferum. Scanning electron micrographs of sclerites from base of

peduncle: A. 0,14 mm; B. 0,14 mm; C. 0,16 mm; D. 0,13 mm; E. 0,19 mm.

450-650 m, February 1987, 10 colonies, bottom trawl, R. Melville Smith, Sea

Fisheries Research Institute, Crab Survey.

Description

The colony, SAM—H3707, is 340 mm in length, tip of peduncle missing.

Colony elongate, slender, curved. Rachis expands gradually toward distal end,

which is obtuse and knob-like. Autozooids 10-15 mm long, c. 50 in number, in

two longitudinal rows along the concave face of the rachis. Convex side of rachis

is free of autozooids but is quite rough and has a granular appearance due to the

presence of the spiculated calyces of numerous siphonozooids. These zooids are

also found at the base of the autozooids. Colony densely spiculated throughout.

Sclerites are tuberculated three-flanged spindles varying in length from

0,1 mm to 0,6 mm. The longer sclerites from the polyps and rachis are tuberculate

only at the ends. The shorter spindles of the peduncle may have tubercles

throughout the entire length but mostly concentrated at the ends. Tubercles

rounded and prominent. The appearance, size, and distribution of these sclerites

agrees with those figured by Ktkenthal & Broch (1911: 228) for the species.

Colour in alcohol is brownish-grey.

Distribution (Fig. 40)

Atlantic Ocean from the Norwegian Sea to Namibia, and at scattered

locations in the Pacific. In southern Africa it is known only from off the Skeleton

68 ANNALS OF THE SOUTH AFRICAN MUSEUM

Coast of Namibia (450-650 m). This study establishes a new record for southern

Africa. The type locality is the Grand Banks, south-east of Newfoundland.

Remarks

The form of the sclerites of this specimen agrees with those figured by

Kikenthal & Broch (1911: 228). They are slender and many have knobby

protuberances at each end. Kophobelemnon heterospinosum Kikenthal, 1910,

has similarly shaped sclerites but without prominent tuberculation. Kophobelem-

non affine Studer, 1894, has tubercles on the peduncular sclerites, but the colonial

sclerites as a whole are more stout, being proportionally wider than those of

K. stelliferum.

Kophobelemnon stelliferum is distinguished from other southern African sea

pens by its elongate clavate appearance with rachis that gradually widens distally,

20-40 polyps arranged biserially, and peduncular sclerites that are rods with

knobby tubercles.

Kophobelemnon sp.

Fig. 38

Remarks

A 50-mm long colony, possessing only two antozooids, has recently been

collected from off Durban, Natal, at 820m in depth. Four species are

characterized by having a reduced number of autozooids: Kophobelemnon

hispidum Nutting, 1912; K. pauciflorum Hickson, 1916; K. molanderi Pasternak,

1975b nom. nov. (= K. pauciflorum Molander, 1929); and K. macrospinosum

J. A. Thomson, 1927 (= K. biflorum Pasternak, 1960). The unavailability of

pertinent literature sources precluded the making of necessary comparisons.

Family Anthoptilidae Kolliker, 1880

Colonies long and whip-like, bilaterally symmetrical with polyps in two

longitudinal series. Polyps in oblique rows, non-retractile, calyces absent.

Siphonozooids are numerous on the rachis situated between the oblique rows of

autozooids. Sclerites are absent except for minute ovals of the stalk interior. Axis

present; round to somewhat quadrangular in cross section. Peduncle stout to

robust, not more than one-fifth total colony length.

A monogeneric family of the Atlantic Ocean and Arctic region 238-2 744 m.

Genus Anthoptilum Kolliker, 1880

Virgularia (non Lamarck, 1816) Verrill, 1879: 239.

Anthoptilum Kolliker, 1880: 13. Jungersen, 1904: 65. Kikenthal & Broch, 1911: 232. Kiikenthal,

1915: 32 (Lit!). Hickson, 1916: 138. Grasshoff, 1982b: 954.

Benthoptilum Verrill, 1885: 510.

Stephanoptilum Roule, 1905: 455.

Thesioides J. A. Thomson & Henderson, 1906: 91.

SOUTHERN AFRICA PENNATULACEA 69

Diagnosis

The characters of the family.

Two species of the Atlantic and Arctic.

Type species. Virgularia grandiflora Verrill, 1879; northern Atlantic.

Anthoptilum grandiflorum (Verrill, 1879)

Figs 17, 40

Virgularia grandiflora Verrill, 1879: 239.

Anthoptilum thomsoni KOlliker, 1880: 13, pl. 4 (figs 16-18). J. A. Thomson & Rennet,

UGS NEB).

Anthoptilum grandiflorum: Jungersen, 1904: 66. Hickson, 1904: 233, pl. 8 (fig. 14). Kiikenthal &

Broch, 1911: 233, pl. 14 (figs 5-7), pl. 19 (figs 44-46) (partim). J. S. Thomson, 1915: 17.

Kikenthal, 1915: 32, fig. 44 (partim) (Lit!). Tixier-Durivault, 1954: 629. Grasshoff, 1982a:

741, 750; 1982b: 954, fig. 35 (Lit!).

Anthoptilum sertum: Kikenthal, 1915: 33.

Material

SAM-H1213, off Lion’s Head, Cape Peninsula (33°55’S 17°55’E), 313 m,

16 March 1900, 2 colonies, large dredge, S.S. Pieter Faure survey, station

JE AASKO

Description

Colonies examined are 400-1 050 mm in length. Polyps 5-10 per row, 8-

25 mm in length. Adjacent polyps may be joined proximally, forming a common

base for an entire oblique row, or only two or three polyps in a row may be joined,

this group being free from the rest of the polyps in the row. Colour often bright red

in life, yellowish brown in alcohol.

Distribution (Fig. 40)

In southern Africa, known from off the west coast of the Cape Peninsula, at

238-348 m. The species is also recorded from the western, eastern, and southern

Atlantic, as well as the Arctic region. Recorded depth range is 238-2 500 m.

Remarks

This species is distinguished from other southern African pennatulaceans by

its whip-like appearance, lack of calyces and sclerites, and with adjacent polyps

united at their bases into groups of 5-10 polyps forming short oblique rows.

Family Funiculinidae Gray, 1870

Elongate, whip-like colonies with polyps arranged biserially, often covering

the lateral and ventral surfaces of the rachis. Polyps are retractile into tubular,

eight-toothed calyces, which may be arranged irregularly or in somewhat oblique

rows. Siphonozooids few. Sclerites numerous; three-flanged spindles, rods, and

ovoid plates.

A monogeneric family of cosmopolitan distribution.

ANNALS OF THE SOUTH AFRICAN MUSEUM

res

———

ee a fi

; sls

A oS ae

}

7 —~ /

' Y Ths 7

; ‘ LLG ee, C= eee Lm

NS SS ee reget

= 50 mm. B. Detail of group of

autozooids with numerous siphonozooids at base. Scale = 10 mm.

Fig. 17. Anthoptilum grandiflorum. A. Entire colony. Scale

SOUTHERN AFRICA PENNATULACEA 71

Genus Funiculina Lamarck, 1816

Funiculina Lamarck, 1816: 422. Herklots, 1858: 8. Kolliker, 1872: 250. Kiikenthal & Broch,

1911: 241. Ktikenthal, 1915: 34 (Lit!). Manuel, 1981: 53.

Leptoptilum Kolliker, 1880: 27.

Trichoptilum Kolliker, 1880: 29.

Diagnosis

The characters of the family.

Three species are recognized worldwide.

Type species. Pennatula quadrangularis Pallas, 1766.

Funiculina quadrangularis (Pallas, 1766)

Fig. 38

Pennatula quadrangularis Pallas, 1766: 372.

Funiculina quadrangularis Jungersen, 1904: 49. Kikenthal & Broch, 1911: 243, pl. 15 (fig. 12)

(Lit.!). Kiikenthal, 1915: 34, figs 45-46. J. S. Thomson, 1915: 15, figs 45-46. Pasternak,

1975b: 104. Manuel, 1981: 53, fig. 14. Grasshoff, 1981: 227.

Material

Material was not available for examination.

Distribution (Fig. 38)

Cosmopolitan, recorded up to 2 300 m in depth.

Remarks

Thomson (1915: 15) recorded one 490-mm specimen from 146-183 m off

Cape Vidal, Natal (28°10’S 32°40'E). According to Manuel (1981: 53), Kikenthal

(1915: 34), and the above reference, the species is characterized by having a dis-

tinctly quadrangular axis covered with a thin coenenchyme. Colonies may reach

2 199 mm in length. Distal portions of colonies are flexible and often tend to

curve or coil spirally. Pedunclular interior with ovoid plate-like sclerites. Calyces

with elongate, three-flanged spindles (up to 9,63 mm in length) forming

eight, pointed terminal teeth and a transverse band of similar sclerites sur-

rounding the base of the teeth. The surface of the rachis possesses three-

flanged rod-like sclerites.

Family Protoptilidae Kolliker, 1872

Colonies elongate, slender, bilateral. Autozooids arranged in one to three

longitudinal series, retractile into spiculate calyces. Sclerites are numerous,

three-flanged, mostly spindles. Axis prominent and rounded in cross-section.

Three genera of the Atlantic and Indo-Pacific: Protoptilum Kolliker, 1872a;

Distichoptilum Verrill, 1882, and Helicoptilum Nutting, 1912. The present study

represents the first records of the family for southern Africa.

WD ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus Distichoptilum Verrill, 1882

Distichoptilum Verrill, 1882: 362. Kiikenthal, 1915: 39. Hickson, 1916: 101.

Juncoptilum J. A. Thomson & Henderson, 1905: 555.

Diagnosis

Polyps arranged in two longitudinal series. No polyps on ventral surface.

Axial side of calyces appressed to lateral margin of rachis. The siphonozooids are

two per autozooid, placed on the rachis directly above the mouth of each

autozooid, one on the dorsal side and one on the ventral side of the calyx.

A monospecific genus.

Type species. Distichoptilum gracile Verrill, 1882; eastern North America.

Distichoptilum gracile Verrill, 1882

Figs 18, 19H-I, 40

Distichoptilum gracile Verrill, 1882: 362. Jungersen, 1904: 62. J. A. Thomson & Henderson,

1906: 87. Kikenthal, 1915: 39 (Lit.!). Pasternak, 1975b: 106. Grasshoff, 1982a: 738,

741-742, map 8; 1982b: 955, figs 38-39.

Juncoptilum Alcocki J. A. Thomson & Henderson, 1905: 555.

Material

SAM-—H3163, west of Table Bay, western Cape Province (33°49’S 16°30’E),

2744 m, 27 August 1959; many fragments of several colonies, 15-foot beam

trawl, Division of Sea Fisheries.

Description

Colonies examined are perhaps up to 1 000 mm in length and 4 mm in width.

Polyps in two longitudinal series, alternate or sub-opposite, retractile into promi-

nent calyces. Distance between polyps along rachis 4—7 mm. Calyces 2—4 mm

long. Calyces adhere to rachis along their inner proximal portions and face

upward. Calyces usually with two terminal teeth, but these are sometimes obscure

in autozooids of the lower rachis. Sclerites of the calyces are large spindles (up to

0,5 mm in length), three-flanged and smooth. Peduncular sclerites are shorter and

inconspicuously three-flanged. Calyces orange, rachis yellowish to cream.

Distribution (Fig. 40)

Previously recorded from the eastern Pacific, Indian, and northern Atlantic

Oceans, 793-4 300 m. This work represents the first record of the genus and

species in southern Africa, and extends the range in the Atlantic southward from

Europe and eastern North America.

Remarks

A considerable amount of variation is evident in this species. Kukenthal

(1915: 40) stated that the calyces possess six terminal teeth and the length of calyx

sclerites is up to 1,0 mm. Jungersen (1904: 62) regarded number of calyx teeth as

SOUTHERN AFRICA PENNATULACEA 73

Fig. 18. Distichoptilum gracile. A. Entire colony. Scale = 50 mm. B. Detail of colony from

three different parts of rachis showing calyces. Scale = 2mm. C. Sclerites from calyx.

Scale = 0,2 mm. D. Sclerites from peduncle. Scale = 0,1 mm.

74 ANNALS OF THE SOUTH AFRICAN MUSEUM

Wty Sec 2

“gail

Fig. 19. Scanning electron micrographs of sclerites. A-—G. Halipteris africana. A-D. From

peduncle: 0,07 mm; 0,05 mm; 0,04 mm. E. From tentacle: 0,11 mm. F. From calyx: 0,58 mm.

G. Tip of calyx sclerite: 0,2 mm. H-I. Distichoptilum gracile. From calyx: 0,36 mm; 0,44 mm.

J-N. Umbellula thomsoni. J-L. From anthocodial neck zone: 0,05 mm; 0,05 mm; 0,06 mm.

M-N. From tentacle: 0,2 mm; 0,24 mm.

SOUTHERN AFRICA PENNATULACEA WD

a matter of the degree of contraction of the polyps, as some highly contracted

polyps show two lobes, whereas partially contracted polyps often exhibit six

distinct teeth. He also stated that the length of calyx sclerites varies between 0,528

and 0,800 mm. The calyces of the southern African material show two large teeth

only and the sclerites vary from 0,28—0,50 mm in length.

Distichoptilum gracile is distinguished from other southern African sea pens

by its thin whip-like appearance, small polyps retractile into permanent calyces

that are appressed close to the surface of the rachis and are arranged biserially in

two longitudinal rows.

Family Scleroptilidae Jungersen, 1904

Polyps usually situated in pairs or groups of three along rachis, separated by

intervals of bare rachis. Axis thin and delicate, extending entire length of colony.

Siphonozooids few and scattered on rachis. Calyces absent. Sclerites present or

absent.

A family of two genera from the North Atlantic and Indo-Pacific.

Genus Scleroptilum Kolliker, 1880

Scleroptilum KO6lliker, 1880: 30. Balss, 1910: 24. Kikenthal & Broch, 1911: 265. Kiikenthal,

1915: 43.

Diagnosis

Sclerites (rods or three-flanged spindles) are present in the walls of the

autozooids.

A genus of at least two or three species from the northern Atlantic and

Indo-Pacific.

Type species. Scleroptilum grandiflorum Kolliker, 1880; Japan.

Scleroptilum sp.

Fig. 38

Remarks

Five colonies, 35-50 mm in length, have recently been collected from off the

Transkei coast at 510 m in depth. These colonies can be assigned to the genus

Scleroptilum by the arrangement of the polyps in whorls or clusters of 2—3 (each

cluster separated by bare areas of rachis), lack of calyces, and the possession of

sclerites in the walls of the polyps. The sclerites differ from those of Scleroptilum

grandiflorum KOlliker, 1880, and the material has not been identified to species.

This represents the first record for the family and genus in southern Africa.

Genus Calibelemnon Nutting, 1908

Calibelemnon Nutting, 1908: 562. Balss, 1910: 70. Kiikenthal, 1915: 44.

Prochunella Balss, 1909: 426.

76 ANNALS OF THE SOUTH AFRICAN MUSEUM

Diagnosis

Sclerites completely absent or with small bodies restricted to the peduncle.

A genus of perhaps two species of the Indo-Pacific.

Type species. Protocaulon indicum J. A. Thomson & Henderson, 1906; Bay

of Bengal.

Calibelemnon sp.

Fig. 38

Remarks

Several colonies were collected in 1975 from near Cape Vidal, Natal, at

740 m in depth. Unlike Calibelemnon indicum (J. A. Thomson & Henderson,

1906), in which sclerites are completely absent, these colonies possess small

sclerites in the peduncle, and therefore probably represent an undescribed

species. This represents a new record of the genus in southern Africa and extends

the range to the south-western fringe of the Indo-Pacific region.

Family Chunellidae Kikenthal, 1902

Rachis slender. Axis quadrangular. Polyps in well-separated whorls. One to

four polyps per whorl. Polyps without calyces. Siphonozooids few and incon-

spicuous, on rachis between autozooids. Sclerites absent except for minute oval

bodies in the interior of the peduncle.

Two genera of the Indian Ocean. The present study represents the first

record of the family in southern Africa.

Genus Chunella Kiukenthal, 1902

Chunella Kiikenthal, 1902: 302; 1915: 45. Kikenthal & Broch, 1911: 271.

Diagnosis

With terminal autozooid highly reduced or absent. Whorls with 1-4

autozooids.

A genus of three nominal species, from the eastern coast of Africa to the

Malay Archipelago.

Type species. Chunella gracillima Kiikenthal, 1902; East Africa.

Chunella gracillima Kikenthal, 1902

Figs 20, 38

Chunella gracillima Kikenthal, 1902: 302; 1915: 45, fig. 53. Kiikenthal & Broch, 1911: 272, pl. 15

(fig. 13). Bayer, 1956: 227.

Material

SAM-—H3735, off Port Durnford, Natal (28°37,8’S 32°38,4’E), 1 000-1 200 m,

25 May 1976, 8 whole colonies, beam trawl, South African Museum R.V. Meiring

Naude cruises. SAM—H4029, one whole colony, same data as SAM-—H3735.

SOUTHERN AFRICA PENNATULACEA I

Fig. 20. Chunella gracillima. A. Entire colony. Scale = 30 mm. B. Transverse section of axis.

Scale = 0,7 mm. C. Two groups of three autozooids with minute siphonozooids on rachis. Scale

= 5mm. D. Minute sclerites from lower peduncle. Scale = 0,02 mm. E. Rudimentary polyp

from distal tip of colony. Scale = 1,2 mm.

78 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

Colonies examined vary in length from 320-550 mm. Number of autozooids

per whorl varies from one to four (most commonly three). Whorls 3-7 per colony

separated by 30-60 mm intervals of bare rachis. Autozooids vary from 3 mm to

15 mm in length in a preserved state. Siphonozooids few and inconspicuous on the

rachis at each autozooid whorl, consistent in form and size throughout the colony.

Axis conspicuously quadrangular, 0,5—1,5 mm in width. Peduncle up to 50 mm in

length and 3-4 mm in width. Sclerites absent except for minute oval bodies up to

0,02 mm long in the interior of the lower peduncle of all colonies examined.

Autozooids greyish with brown tentacles, rachis and peduncle cream.

Distribution (Fig. 38)

Equatorial eastern Africa and Natal, 818-1 200 m. This is the first record of

the genus and species in southern Africa. The type locality is Pemba Island,

border of Kenya and Tanzania, 818 m in depth.

Remarks

This species is distinguished from other southern African sea pens by its 3-7

demarcated clusters of 2—4 polyps and possession of a single rudimentary terminal

polyp.

Remarks on the genus Chunella

Kikenthal & Broch (1911: 275) described Chunella quadriflora for colonies

with lower whorls of four autozooids, from the same locality as C. gracillima.

Hickson (1916) described C. biflora (from the Malay Archipelago) for colonies

with two autozooids per whorl. Kiikenthal (1915: 45) characterized C. gracillima

as having three autozooids per whorl.

Of the eight colonies from the same trawl sample that I have examined, one

has four autozooids in a lower whorl and three in an upper whorl. Another colony

has two autozooids in upper whorls and three in lower whorls. Some whorls in

other colonies may have one, two, or three autozooids per whorl. It therefore

seems that, since the number of autozooids per whorl is not consistent in a given

colony, the character can hardly be used to separate species. As a result, I suspect

that the three species may in fact be conspecific but the pertinent type material is

presently unavailable for comparative examination.

Kikenthal & Broch (1911, pl. 20 (fig. 49)) showed a photograph labelled

‘Chunella quadriflora’. The specimen appears to possess three whorls of two

autozooids each and a single well-developed terminal autozooid. The photograph

therefore appears to be that of a colony of Amphiacme abyssorum and not

Chunella quadriflora.

Genus Amphiacme Kikenthal, 1903

Amphianthus Kikenthal, 1902: 303.

Amphiacme Kikenthal, 1903: 275; 1915: 46. Kiikenthal & Broch, 1911: 276.

SOUTHERN AFRICA PENNATULACEA 719

Diagnosis

With a single terminal autozooid that is well-developed and somewhat

asymmetrical. Autozooids in pairs. Siphonozooids present on the keel of the

terminal autozooid as well as on the rachis between the individual autozooids of

each pair.

A monospecific genus of the eastern African coast.

Type species. Amphianthus abyssorum Kikenthal, 1902; East Africa.

Amphiacme abyssorum (Kikenthal, 1902)

Figs 21, 38

Amphianthus abyssorum Kikenthal, 1902: 302.

Amphiacme abyssorum: Kiikenthal, 1903: 275; 1915: 47, fig. 54. Kikenthal & Broch, 1911: 276,

pl. 15 (fig. 14). Tixier-Durivault, 1972: 52. Tixier-Durivault & D’Hondt, 1974: 260.

Material

SAM-H3736, off Jesser Point, Natal (27°38,6'S 32°52,6’E), 860 m, 21 May

1976, 1 whole colony, beam trawl, South African Museum R.V. Meiring Naude

cruises. SAM-—H3737, off Port Durnford, Natal (28°37’S 32°38’E), 1 000-

1 200 m, 25 May 1976, 2 whole colonies, beam trawl, South African Museum

R.V. Meiring Naude cruises.

Description

The colonies examined are 230-250 mm in length. Axis conspicuously

quadrangular, usually <1 mm in width. Autozooids in two or three pairs along

rachis, at intervals of 30-50 mm. Autozooids about 10 mm in length preserved.

Terminal autozooid bilaterally symmetrical or somewhat asymmetrical, 8-10 mm

long with a ventral keel. Peduncle 10-12 mm in length and 1-2 mm in width.

Siphonozooids few, located on the rachis at each whorl and on the keel of

terminal autozooid. Sclerites absent except for oval bodies (<0,02 mm) in

peduncle. Polyps brownish-grey, rachis cream or tan, peduncle brownish.

Distribution (Fig. 38)

The Indian Ocean coast of Africa, from the equator to Natal, and

Madagascar; 760-1 200m. The present study represents a new record for

southern Africa and extends the range of the genus and species south to South

Africa. The type locality is the coastal region near the border of Kenya and

Tanzania, 818-1 019 m in depth.

Remarks

Amphiacme abyssorum is distinguished from other southern African

pennatulaceans by the arrangement of the polyps in two or three demarcated pairs

along the rachis, and the possession of a single modified terminal polyp.

80 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 21. Amphiacme abyssorum. A. Two entire colonies. Scale = 40mm. B. Terminal

autozooid with ten minute siphonozooids on keel. Scale = 10 mm. C. Single tentacle from

autozooid. Scale = 1 mm. D. Sclerites from peduncle. Scale = 0,02 mm. E. Detail of rachis

with a pair of autozooids and several siphonozooids. Scale = 10 mm. F. Transverse section of

axis. Scale = 0,5 mm.

SOUTHERN AFRICA PENNATULACEA 81

Family Umbellulidae Lindahl, 1874

Rachis long and slender. Axis quadrangular to round in cross section.

Autozooids restricted to the distal terminus, usually forming an umbellate cluster.

Siphonozooids are present on the rachis at the base of the autozooids or on bare

parts of the rachis. Sclerites present in peduncle and terminal cluster, or totally

absent. When present, sclerites are rods, spindles or needles, three-flanged round

in cross-section.

A monogeneric family.

Genus Umbellula Cuvier, 1798

Isis Linnaeus, 1758: 800 (partim).

Pennatula Pallas, 1766: 335 (partim).

Vorticella Linnaeus, 1767: 1317.

Ombellula Cuvier, 1798: 675.

Umbellularia Lamarck, 1816: 436. Kolliker, 1872a: 203.

Umbellula Gray, 1870: 39. Kolliker; 1875: 11. Kikenthal & Broch, 1911: 282 oe Broch, 1957:

350; 1958a: 267; 1958b: 251. Grasshoff, 1982b: 956.

Diagnosis

Characters of the family.

A genus of perhaps 12 valid species. Cosmopolitan distribution; deep sea,

250 m to over 6 100 m.

Type species. Isis encrinus Linnaeus, 1758; Arctic Ocean.

Umbellula thomsoni (Kolliker, 1874)

Figs 19J-N, 22, 23, 40

Umbellularia Thomsonii Kolliker, 1874: 13.

Umbellula Thomsonii: Kolliker, 1875: 11.

Umbellula thomsonii: Broch, 1958: 253, figs 2, 4 (Lit!). Grasshoff, 1972: 2, fig. 1.

Umbellula Guntheri Broch, 1913: 3, pl. 1 (fig. 1).

Umbellula aciculifera J. S. Thomson, 1915: 20 (and text-fig. on p. 21), pl. 2 (figs 1-2).

Umbellula thomsoni: Kolliker, 1880: 19. Kiikenthal, 1915: 54. Pasternak, 1970: 240, fig. 2; 1975b:

106. Grasshoff, 1982a: 753, map 11; 1982b: 958.

Material

SAM-H1166, south-west of Cape Point (34°40’S 17°45’E), 1 650 m, 21 July

1903, 1 colony, shrimp trawl, S.S. Pieter Faure survey, station PF 17026. (Holo-

type of Umbellula aciculifera Thomson, 1915.)

Description

The colony examined is about 475 mm in length. The axis is 3 mm in

diameter, rounded quadrangular near the terminal cluster. Umbellate cluster

composed of 10 autozooids, each 15-20 mm long. Sclerites conspicuous and

numerous. Terminal end of rachis, polyp walls, and proximal portions of tentacles

with knobby, three-flanged rods and spindles (<0,3 mm). Aboral surface of

tentacles and pinnules with spindles (0,5—1,3 mm); round in cross-section, not

82 ANNALS OF THE SOUTH AFRICAN MUSEUM

were a ee ene Neem ee

et rn eens

Scope SAO. ; pee

ee re EE

ae an nrg pet pag gr en pags mene

Sete

Rene a5

Se LL.

awe

Fig. 22. Umbellula thomsoni. A. Entire colony. Scale = 20mm. B. Transverse sections

through three different areas of axis. Scale = 2 mm. C. Single tentacle. Scale = 3 mm. D. Small

sclerites from proximal portion of tentacle. Scale = 0,1 mm. E. Sclerites from body wall of

autozooid. Scale = 0.02 mm. F. Large sclerites from proximal portion of tentacle. G. Sclerites

from pinnule. F—G. Scale = 0,5 mm. H. Sclerites from peduncle. Scale = 0,04 mm.

SOUTHERN AFRICA PENNATULACEA. 83

three-flanged. Peduncle with rough ovals or rods (0,08—0,11 mm in length),

having numerous low, rounded knobs or tubercles.

Distribution (Fig. 40)

Apparently cosmopolitan, 1 300-6 200m. This specimen is the only known

southern African record, first described by J. S. Thomson (1915) as Umbellula

aciculifera (type specimen). The type locality of U. thomsoni is the north Atlantic.

Fig. 23. Scanning electron micrographs of sclerites of Umbellula thomsoni. A-D. From

peduncle: 0,08 mm; 0,09 mm; 0,1 mm; 0,11 mm. E. Detail of surface of D; length of

micrograph 0,04 mm. F-H. Large sclerites from tentacle: 0,7 mm; 0,8 mm; 0,7mm.

84 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

The colony has five solitary epizoic zooanthids (species indeterminate)

attached to bare areas of the upper rachis. They vary from 5—15 mm in diameter

and 10-12 mm in length (zooanthids not shown in Fig. 22A).

This species is distinguished from other southern African sea pens by having

the autozooids restricted to a cluster at the distal end of an elongate colony, and

the possession of sclerites in the autozooids.

Umbellula lindahli Kolliker, 1874

Figs 24, 40

Umbellula lindahlii K6lliker, 1874: 12. Jungersen, 1904: 75, pl. 3 (figs 37-46). Kiikenthal,

1915: 48.

Umbellula miniacea Lindahl, 1874: 12, pls 1-2.

Umbellula pallida Lindahl, 1874: 13, pl. 3.

Umbellula lindahli: Broch, 1958: 262 (Lit.!). Grasshoff, 1982a: 741-742, 753; 1982b: 958, fig. 43.

Pasternak, 1975b: 109.

Material

SAM-—H3738, off the Cape of Good Hope Peninsula (34°37'S 17°03’E),

2 890-2 963 m, 8 December 1959, 2 colonies, beam trawl, Division of Sea

Fisheries Survey, station A315. SAM—H3739, west of Dassen Island, western Cape

Province (33°36’S 16°15’E), 2 780-2 872 m, 26 August 1959, 2 colonies, beam

trawl, Division of Sea Fisheries Survey. SAM—H4028, south of False Bay

(35°22,3'S 18°45,4’E), 490 m, 6 July 1986, 22 colonies, beam trawl, Rob Lesley

(Sea Fisheries Research Institute, West Coast Biomass Survey, station A4305046).

Description

Colonies examined are 770-900 mm in length. Axis conspicuously quad-

rangular, 1-2 mm in width. Peduncle up to 150 mm long by 6 mm in width.

Umbellate terminal cluster with 8—10 autozooids, each 20-30 mm long. Siphono-

zooids numerous on keel-like terminus of rachis. Sclerites totally absent.

Distribution (Fig. 40)

Cosmopolitan up to 6 100 m in depth. In southern Africa, the species is

presently known only from the south-western Cape (490-2 963 m). This study

represents a new record of the species to southern Africa. The type locality is the

Arctic region of Greenland.

Remarks

This species is differentiated from other southern African sea pens by the

restriction of the autozooids to a cluster at the terminal end of an elongate colony,

and the complete absence of sclerites.

SOUTHERN AFRICA PENNATULACEA 85

Fig. 24. Umbellula lindahli. A. Entire colony. Scale = 50 mm. B. Transverse section through

peduncle. Scale = 1,5 mm.

86 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks on the genus Umbellula

Grasshoff (1972) recognized three species of the genus that have sclerites

in the terminal autozooid region. Umbellula monocephalus Pasternak, 1964

(jun. syn. U. thieli Grasshoff, 1972) has only a single autozooid; the other two

species both have multiple autozooids—U. durissima KOlliker, 1880, has both

axis and sclerites that are round in cross section, whereas U. thomsoni has

three-flanged sclerites and a quadrangular axis. I have found that shape of the

axis in cross-section is an invalid differentiating character with regards to

U. thomsoni, since different parts of the axis of the colony examined

(SAM-H1166) are round or rounded quadrangular, but never sharply

quadrangular (Fig. 22B). Also, the large sclerites of the tentacles and pinnules

(>0,5 mm) are round in cross-section, whereas the small sclerites (<0,3 mm)

of the proximal regions of the tentacles and of the entire polyp body wall are

conspicuously three-flanged (Figs 19J-N, 22D—-E, 23F—H). Thus, this specimen

(which is the type upon which the des-cription of U. aciculifera Thomson, 1915, was

based) possesses characteristics intermediate between U. durissima and U. thom-

soni. In addition, Grasshoff (1972, fig. 2B) showed three-flanged spindles from the

polyp walls of U. durissima. These facts tend to negate distinctions between the two

species. Examination of type material and a variety of specimens is necessary to

determine if we are dealing with two distinct species or a single variable one. This

material is presently unavailable to me for comparison.

Broch (1958b) revised the genus Umbellula and recognized five species

without sclerites in the terminal cluster. Umbellula hemigymna Pasternak,

1975a, was subsequently added from the Caribbean Sea. Of these six,

U. lindahli is characterized by having quadrangular axis, large autozooids on

a tassel not especially crowded, and full-grown colonies >400 mm in length.

Broch (1961) compared specimens of U. encrinus (Linnaeus, 1758) with those

of U. lindahli and considered them conspecific, thereby making the latter a

junior synonym of U. encrinus. Subsequent workers, including Pasternak

(1961b, 1964, 1975b, 1980) and Grasshoff (1982a, 1982b), did not recognize

this synonymy and considered the two species as distinct. I have retained this

distinction here. Grasshoff (1982b: 957-958) distinguished the two species as

follows: U. encrinus—stalk thick, polyps stocky; U. lindahli—stalk thin,

polyps slender.

Family Virgulariidae Verrill, 1868

Colonies bilateral, long and slender to whip-like. Axis well-developed,

present throughout length of colony. Adjacent autozooids fused proximally,

forming polyp leaves or low ridges that are arranged in transverse or oblique

rows along rachis. Sclerites three-flanged spindles, minute plates, or entirely

absent.

The family is of cosmopolitan distribution, with two subfamilies and six

genera.

SOUTHERN AFRICA PENNATULACEA 87

Subfamily Virgulariinae Verrill, 1868

Autozooids form low to well-developed polyp leaves (flattened expansions)

by fusion of their proximal portions.

Five genera, two of which are included below, in addition to Acanthoptilum

K6lliker, 1870, Scytalium Herklots, 1858, and Stylatula Verrill, 1864.

Genus Virgularia Lamarck, 1816

Virgularia Lamarck, 1816: 429. Kiikenthal & Broch, 1911: 323. Kitikenthal, 1915: 71 (Lit!).

Lygus Herklots, 1858: 11.

Halisceptrum Herklots, 1863: 33.

Sceptonidium Richiardi, 1869: 63.

Cladiscus Koren & Danielssen, 1877: 101.

Protocaulon Kolliker, 1880: 26.

Svava Danielssen & Koren, 1884: 57.

Deutocaulon Marshall & Fowler, 1888: 453.

Svavopsis Roule, 1908: 181.

Diagnosis

Colonies long and slender, often vermiform. Peduncle and rachis usually

abruptly delimited. Autozooids form polyp leaves that are arranged in two

longitudinal series along rachis. Autozooids retractile into fleshy calyces.

Siphonozooids are present on polyp leaves below the free parts of the autozooids

or on the rachis between the polyp leaves. Sclerites absent except for minute ovals

and plates of the peduncle interior. Axis present throughout entire length of

rachis and upper portion of peduncle, often projecting beyond apex of rachis.

A genus of about 20 species, in need of revision. Cosmopolitan.

Type species. Pennatula mirabilis Linnaeus, 1758; North Atlantic.

Virgularia schultzei Kikenthal, 1910

Figs 1D-F, 25, 28C-D, 41

Virgularia schultzei Kiikenthal, 1910: 53; 1915: 72, figs 72, 73 (Lit.!). Kikenthal & Broch, 1911:

326, figs 124-125, pl. 17 (figs 21-22). J. S. Thomson, 1915: 16. Broch, 1939: 30. Tixier-

Durivault, 1954: 630.

Virgularia Reinwardti (non Herklots, 1858) Hickson, 1900: 87. Balss, 1910: 47 (partim).

Virgularia spec. (juv.) Broch, 1910: 231.

Material

SAM-H3387, off Danger Point, Cape Province (34°37’S 19°24'E), 19 m,

11 April 1984, 11 colonies, SCUBA, W. R. Liltved. SAM—H3159, off Koeberg

Power Station, western Cape Province (33°40’S 18°26’E), 6-20m, 24-26

November 1981, 8 colonies, SCUBA, W. R. Liltved. SAM—H3338, Algoa Bay,

eastern Cape Province (33°50’S 25°40’E), 16 m, 17 May 1984, 1 colony, SCUBA,

G. C. Williams.

ANNALS OF THE SOUTH AFRICAN MUSEUM

a3 4

a :

Fig. 25. Virgularia schultzei. A. Entire preserved colony. Scale = 20 mm. B. Expanded colony

in situ. Scale = 20 mm. C. Polyp leaf from a 120 mm long colony. Scale = 1 mm. D. Peduncle

sclerites. Scale = 0,005 mm. E. Lower surface of a polyp leaf from middle of rachis of a 400-mm

long colony. Scale = 2mm. F. Two retracted polyps from a polyp leaf showing siphonozooids

at base. Scale = 0,5 mm.

SOUTHERN AFRICA PENNATULACEA 89

Description

Colonies examined ranged in length from 85-500 mm, and 5-12 mm in

width. Peduncle long and flexible. Axis up to 1,5 mm in diameter, round in

cross-section. Surface of axis often with numerous perforations (canals of

Kolliker) 0,003—0,004 mm in diameter. Polyp leaves well developed, numerous,

transparent. Autozooids 15-35 per polyp leaf. Siphonozooids are present on the

lower surface of the polyp leaves, usually one per autozooid in a band below the

calyx-like free portions of the autozooids. Sclerites absent except for minute ovals

of the peduncle interior (c. 0,005 mm in length). Colour variable, cream or white

to yellowish or orange; uniformly coloured or distinctly mottled.

Distribution (Fig. 41)

Apparently endemic to southern Africa, from Lideritz Bay, Namibia, to at

least as far as Kosi Bay, littoral to 222 m in depth. Day (1974b: 35) listed the

species from Inhambane, Mozambique. This species is one of the most frequently

encountered sea pens in South Africa.

Observations

SCUBA divers have observed this species with patchy or localized dis-

tribution and high population density. An extensive patch about 50 m in depth off

Hout Bay, Cape Peninsula, has numerous colonies that are spaced relatively

uniformly (about 0,5 m apart) in what appears to be a relatively regular pattern

of spacing (P. A. Hulley pers. comm.).

Remarks

Virgularia schultzei is distinguished from other southern African species of

Virgularia by having siphonozooids on the lower surfaces of the polyp leaves,

below the free distal ends of the autozooids, and 15—35 autozooids per polyp leaf.

Virgularia mirabilis (Linnaeus, 1758)

Figs 26, 41

Pennatula mirabilis: Linnaeus, 1758: 819. Miiller, 1776: 255.

Virgularia mirabilis Lamarck, 1816: 431. Kikenthal & Broch, 1911: 329, fig. 126. Kikenthal,

1915: 73, fig. 76 (Lit.!). Manuel, 1981: 54, fig. 15. Tixier-Durivault, 19616: 254, figs 17-18;

1963: 72. Tixier-Durivault & D’Hondt, 1974: 262.

Material

SAM-H3240, mouth of False Bay, western Cape Province (34°22,7’'S

18°43,1’E), 78m, 10 September 1953, 1 colony, dredge, University of Cape

Town Ecological Survey. SAM-—H3746, off Stony Point, Transkei (32°38,9'S

28°45,0’E), 360 m , 17 July 1984, 1 partial colony, dredge; G. C. Williams, R.V.

Meiring Naude. SAM—H3869, 67 km west of Cape Fria, Namibia (18°25’S

11°15’E), 274 m, May—June 1982, many colonies, trawl.

90 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 26. Virgularia mirabilis. A. Entire colony. Scale = 10mm. B. Detail of rachis with

retracted polyps. Scale = 1 mm. C. Part of rachis from colony with expanded polyps. Scale =

10 mm. D. Detail of rachis showing expanded polyps. Scale = 1 mm.

SOUTHERN AFRICA PENNATULACEA 91

Description

The colonies examined are up to 95 mm long, vermiform, very slender and

delicate. Polyp leaves small and not crowded. Autozooids 5-12 per polyp leaf.

Autozooids globular or long and narrow, about 1,0 mm long and 0,2 mm wide.

Siphonozooids inconspicuous on the sides of the rachis below the polyp leaves and

do extend on to the dorsal surface of the rachis. Peduncle about as long as rachis.

Sclerites absent except for minute ovals (approximately 0,005 mm in length) from

interior of peduncle. Colour yellowish to cream.

Distribution (Fig. 41)

Atlantic Ocean, including the Arctic region, Mediterranean Sea, northern

and western Africa, and the south-western Indian Ocean, 9—400 m in depth.

Known in southern Africa from Cape Fria and Walvis Bay, Namibia; the Cape of

Good Hope region, Cape Province, and the Transkei coast, 78—360 m in depth.

Tixier-Durivault (1963: 73) recorded the species from just south of Walvis Bay,

Namibia. Tixier-Durivault & D’Hondt (1974: 262) reported the species from

Tulear, Madagascar. The type locality is the northern Atlantic.

Remarks

Virgularia mirabilis is distinguished from other southern African species of

the genus by the often limp and vermiform appearance of the colonies, with 5—12

autozooids per polyp leaf. Kikenthal (1915: 34) stated that up to 16 autozooids

per polyp may be present.

Virgularia gustaviana (Herklots, 1863)

Figs 27, 28A-B, 41

Halisceptrum gustavianum Herklots, 1863: 31.

Virgularia gustaviana Kikenthal & Broch, 1911: 334. Kiikenthal, 1915: 74, fig. 77 (Lit.!).

Tixier-Durivault, 1954: 630; 1960: 366.

Material

SAM-H1193, Durban Bay (29°50’S 31°00’E), 23 May 1929, 3 colonies.

SAM-H1199, off Salisbury Island, Durban Bay, Natal (29°50’S 31°00’E), 4 July

1927, 1 colony. SAM—H1509, Durban Bay (29°50’S 31°00’E), 1 colony.

Description

Colonies examined range between 140-419 mm in length and 10-20 mm in

width. Colonies are robust and firm, tapering gradually or abruptly at the apical

end. Polyp leaves large, numerous and crowded, placed opposite each other.

Autozooids highly congested, more than 50 per leaf (often up to 100 or more).

Siphonozooids are inconspicuous on the lateral and dorsal surfaces of the rachis

between the polyp leaves. Peduncle length varies from one third to one half of

total colony length. Peduncle may have a conspicuous end bulb. Sclerites are

minute ovals (c. 0,01 mm) restricted to the peduncle interior. Colour yellowish.

92 ANNALS OF THE SOUTH AFRICAN MUSEUM

SiG

“fnew yeurtes

ii aeie

Fig. 27. Virgularia gustaviana. A. Entire colony. Scale = 20 mm. B. Single entire polyp leaf.

Scale = 3 mm. C. Four adjacent polyps from a polyp leaf. Scale = 1 mm.

SOUTHERN AFRICA PENNATULACEA 93

Fig. 28. A-B. Virgularia gustaviana. A. Dorsal view of rachis. B. Ventral view of rachis;

length of photographs = 80 mm. C-D. Virgularia schultzei. C. Broken end of axis; diameter =

0,36 mm. D. Detail of axial surface; width of axis = 0,36 mm.

94 ANNALS OF THE SOUTH AFRICAN MUSEUM

Distribution (Fig. 41)

Indo-West Pacific. Known in southern Africa only from Inhaca Island,

Mozambique, and Durban Bay, Natal; littoral and shallow sublittoral.

Remarks

Virgularia gustaviana is distinguished from other southern African species of

Virgularia in having 50-200 autozooids per polyp leaf.

Virgularia sp.

Fig. 41

Remarks

Four colonies, 40-80 mm in length, have recently been collected from off the

eastern Cape and Transkei (60—410 m in depth). These colonies are fragmentary,

therefore assignment to species is untenable at present. The thick and firm

appearance of the polyp leaves clearly separates this species from the other

southern African forms of Virgularia.

Other species of Virgularia

Tixier-Durivault & D’Hondt (1974: 262) listed Port Natal (Durban) as a

locality of the Indo-Pacific species Virgularia juncea (Pallas, 1766). Since no

description or figures of colonies from this locality are given, and since there are

no other records or material of V. juncea known from southern Africa, I here

consider this as a dubious record.

Tixier-Durivault (1963: 73) recorded Virgularia tuberculata Marshall, 1887,

from northern Angola. This is outside the geographical limit of the present work,

but is mentioned, here nonetheless as a species that may possibly be encountered

further south. It is distinguished by having tubercles or small teeth on the terminal

rim of the calyx and only 3-6 autozooids per polyp leaf.

Genus Scytaliopsis Gravier, 1906

Scytaliopsis Gravier, 1906: 293; 1908: 237. Kiikenthal, 1915: 67.

Diagnosis

Colony firm. Axis circular in cross-section. Gradual transition between rachis

and peduncle. Small polyp leaves in pairs, separated by areas of bare rachis.

Autozooids 4—5 per polyp leaf. Innermost autozooids of each polyp leaf smaller

than adjacent ones. Anthocodiae are retractile into the globular, fleshy bases of

autozooids. Siphonozooids few and inconspicuous, situated on rachis between

leaves. Sclerites absent.

A monospecific genus of the western Indian Ocean and Red Sea.

Type species. Scytaliopsis djiboutiensis Gravier, 1906; Djibouti (Red Sea).

SOUTHERN AFRICA PENNATULACEA 95

Scytaliopsis djiboutiensis Gravier, 1906.

Figs 29, 41

Scytaliopsis djiboutiensis Gravier, 1906: 293; 1908: 237. Kiikenthal, 1915: 67 (figs 66, 67).

Material

SAM-H3711, off Qora River, Transkei (32°35,4’S 28°49,2'E), 450-460 m,

14 July 1984, 1 colony, dredge, G. C. Williams, R.V. Meiring Naude.

Description

The colony is 92 mm in length; rigid, straight, and shaft-like. Peduncle

shorter than rachis. Polyp leaves arranged in 15—20 pairs, each leaf with not more

than five autozooids. Adjacent autozooids gradually increase in size from

innermost to outermost. Intervals of bare rachis up to 3 mm. Siphonozooids are

very sparse and inconspicuous, usually not more than two are present on the

rachis below each pair of polyp leaves. Sclerites entirely absent. Colour brownish.

Distribution (Fig. 41)

East African coast, littoral to 460 m in depth. This is the first record of the

genus and species for southern Africa and extends the range southward from the

Red Sea to the Transkei coast. The type locality is Djibouti, Red Sea.

Remarks

This species is differentiated from other southern African pennatulaceans by

having 5 polyps per polyp leaf, which grade in size from the smallest on the dorsal

side of each leaf to the largest on the ventral side.

Subfamily Balticininae Gray, 1870

Fusion of proximal portions of adjacent autozooids forms raised ridges or

swellings that emanate laterally from the rachis. Siphonozooids few and incon-

spicuous between autozooids.

One genus of the Atlantic and Indo-Pacific oceans.

Genus Halipteris Kolliker, 1869

non Virgularia Lamarck, 1816: Koren & Danielssen, 1847: 269.

Halipteris Kolliker, 1869: 124.

Pavonaria K6lliker, 1869: 123 (non Schweigger, 1820; see Bayer, 1956: F228). Kukenthal &

Broch, 1911: 305. Kiikenthal, 1915: 60 (Lit!).

Balticina Gray, 1870: 13.

Norticina Gray, 1870: 13.

Osteocella Gray, 1870: 40.

Lygomorpha Koren & Danielssen, 1877: 99.

Stichoptilum Grieg, 1887: 15.

Diagnosis

Colonies long and slender. Axis conspicuous, extending entire length of

colony, round in cross-section. Autozooids arranged in numerous oblique rows in

96 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 29. Scytaliopsis djiboutiensis. A. Entire colony. Scale = 20mm. B. Detail of rachis

showing two polyp leaves. Scale = 2mm. C. Single polyp leaf showing gradation in size of

adjacent autozooids. Scale = 2 mm.

SOUTHERN AFRICA PENNATULACEA 97

two longitudinal series. Adjacent autozooids sometimes fused at their bases, often

forming ridges or swellings that emanate laterally from the rachis. Autozooids

with two-toothed calyces. Anthocodia retractile into calyces. Sclerites numerous;

three-flanged spindles in calyx and tentacles, smooth rods or spindles in peduncle.

A genus of five species; Atlantic, Pacific and Western Indian oceans.

Type species. Virgularia finmarchica Sars, 1851; northern Atlantic.

Halipteris africana (Studer, 1879) comb. nov.

Figs 19A-G, 30, 40

Halipteris sp. Studer, 1878: 139.

Pavonaria africana Studer, 1879: 672, pl. 5 (fig. 41). Kiikenthal, 1915: 61.

Material

SAM-—H3237, north-west of Lamberts Bay, western Cape Province

(31°29,7’'S 16°03,4’E), 459 m, 15 August 1947, 1 colony, dredge, University of

Cape Town Ecological Survey, AFR730G. SAM-—H4027, off Namaqualand

(30°00,5’S 14°53,6’E) 461 m, 20 July 1986, 8 colonies, bottom trawl, Rob Leslie

(Sea Fisheries Research Institute, West Coast Biomass Survey, station

A4373046).

Description

Colonies whip-like, up to 1 550 mm in length and averaging 10 mm in width.

Axis round to rounded quadrangular, approximately 5 mm in diameter. Peduncle

stout, approximately 200 mm long and up to 15 mm wide. Peduncle distinctly

quadrangular. Autozooids arranged in numerous oblique rows, 3-7 (usually 4—6)

per row. Some rows form basal ridges by fusion of proximal regions of autozooids.

In some rows, circular or elliptical walls of the basal ridge surround individual

autozooids or groups of two or three. Aboral surface of tentacles with densely set

three-flanged spindles, 0,10—0,15 mm in length, arranged longitudinally. Calyx

with numerous three-flanged spindles 0,12—0,60 mm long. Terminal teeth of calyx

sometimes blunt or indistinct. Peduncle with smooth spindles or rods 0,04—

0,07 mm in length. Rachis and peduncle pale red to white, calyx light blue, polyps

brown.

Distribution (Fig. 40)

Atlantic coast of Africa, 459-659 m in depth. This study represents a new

record from southern Africa and extends the range of the species south from the

Senegal—Guinea region to the Namaqualand coast, South Africa. The type

locality is tropical west Africa.

Remarks

Gray (1870: 13) described two genera: Balticina for B. finmarchica (Sars,

1851) and Norticina for N. christii (Koren & Danielssen, 1848), presumably after

ANNALS OF THE SOUTH AFRICAN MUSEUM

SCRE

ca.

eec eee

g <

wazre

iH t iG

a Lh LO ae eee

oe ene

Ay \\ hee

j (

SU IS

—

eA = 78) —— Sacee

Nig as

7,5mm. D. Group of six autozooids with

Fig. 30. Halipteris africana. A. Entire colony. Scale = 30 mm. B. Detail of rachis. Scale =

Js “>

7mm. C. Transverse section of axis. Scale

common basal ridge. Scale = 2 mm. E. Two autozooids with ten siphonozooids at their bases.

Scale = 2 mm. F. Single tentacle with sclerites. Scale = 0,5 mm. G. Sclerites from tentacle.

Scale = 0,1 mm. H. Sclerites from peduncle. Scale = 0,04 mm. I. Sclerites from calyx. Scale =

0,25 mm.

SOUTHERN AFRICA PENNATULACEA — 99

the Baltic and Nordic (North) seas, respectively. Both genera are considered

synonyms of Pavonaria Kolliker, 1869, by Kikenthal (1915: 60). However,

Pavonaria was found by Kikenthal (1915: 60) and Bayer (1956: 228) to be

preoccupied by Pavonaria Schweigger, 1820, a synonym of Funiculina. Therefore

the name Halipteris Kolliker, 1869, which is synonymous with Pavonaria Kolliker,

1869, is the valid generic name (M. Grasshoff pers. comm.).

Tixier-Durivault made the original determination for this specimen and

identified it as Funiculina armata Verrill, 1879. However, the possession of ridges

in oblique rows, which unify the fused bases of adjacent autozooids, and the

autozooids with calyces of two terminal teeth prove this to be a misidentification.

Tixier-Durivault (1954: 629) also listed F. armata from South Africa based on two

specimens collected on the same day and in the same vicinity as the present

specimen. I consider this a dubious record until these specimens can be examined.

Kiukenthal (1915: 60-61) distinguished B. africana by the possession of five

or six autozooids per row, and the presence of contiguous distal portions of the

basal swellings of the polyp leaves that may surround the proximal regions of

several adjacent autozooids. This latter feature is common in autozooids of the

colony examined but is not universal (Fig. 31D).

Jungersen (1904: 43) maintained that B. africana is probably not a distinct

species but rather a southern form of the common Atlantic species, B. fin-

marchica (Sars, 1851). However, he did not make a formal synonymy. The

distinguishing characters recognized by Studer (4—6 autozooids per row and low

ridges with wall-like extensions around individual autozooids) may indeed not be

sufficient for specific differentiation. A comparison of many specimens from

various localities is necessary.

Remarks on the genus Halipteris

Three additional colonies assignable to the genus Halipteris are present in the

South African Museum’s collection. They are from the Natal south coast

(900-625 m), the Transkei coast (300 m), and the Namaqualand coast (457 m).

The three colonies are relatively small (105-220 mm in length) and apparently

represent at least two species, which have not been identified at present.

Family Pennatulidae Ehrenberg, 1834

Colonies bilateral with large polyp leaves. Autozooids in one or more

rows on margin of polyp leaves. Anthocodia retractile into calyces with marginal

teeth. Siphonozooids confined to rachis. Mesozooids may be present. Sclerites

numerous; mostly three-flanged needles, rods, plates, or ovals.

Three genera: Pennatula Linnaeus, 1758, of cosmopolitan distribution; and

Leioptilus Gray, 1860, and Ptilosarcus Gray, 1860, both restricted to the Pacific

coast of North America.

100 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus Pennatula Linnaeus, 1758

Pennatula Linnaeus, 1758: 818. Kikenthal & Broch, 1911: 348. Kikenthal, 1915: 81 (Lit.!).

Grasshoff, 19826: 959.

Penna Bohadsch, 1761: 98 (partim).

Diagnosis

Calyces usually with eight terminal teeth. Calyces tubular with numerous

sclerites, which are three-flanged needles. Inconspicuously three-flanged spindles

and plates occur in peduncle.

A cosmopolitan genus of probably 12 to 15 valid species and many uncertain

or dubious ones.

Type species. Pennatula phosphorea Linnaeus, 1758.

Pennatula inflata Kikenthal, 1910

Figs 31A-B, 32, 33A-C, 34, 35, 40

Pennatula inflata Kikenthal, 1910: 52; 1915: 83, figs 81-83. Kikenthal & Broch, 1911: 350, pl. 17

(fig. 25), pl. 22 (figs 61, 62).

Fennatula phosphorea (non Linnaeus, 1758) Tixier-Durivault, 1954: 630.

Material

SAM-—H3238, north-west of Lambert’s Bay (31°43,6’S 16°13,2’E), 457 m,

9 June 1947, 2 colonies, trawl, University of Cape Town Ecological Survey.

Fig. 31. Scanning electron micrographs of sclerites. A-B. Pennatula inflata. From surface of

peduncle: 0,1 mm; 0,08 mm. C-—D. Pteroeides isosceles. From surface of peduncle: 0,08 mm;

0,1 mm.

SOUTHERN AFRICA PENNATULACEA 101

Fig. 32. Pennatula inflata. A. Entire colony, lateral view. Scale 40 mm. B. Two mesozooids

with sclerites. Scale = 0,3 mm. C. Group of siphonozooids. Scale = 0,2 mm. D. Single polyp

leaf showing positions of autozooids (a), mesozooids (m), and siphonozooids (s). Scale =

8,8 mm. E. Group of three retracted autozooids. Scale = 1,6 mm. F. Two autozooids showing

sclerite arrangement. Scale = 1 mm. G. Three sclerites from calyx. Scale = 0,3 mm. H. Seven

sclerites from surface of peduncle. Scale = 0,1 mm. I. Terminal tip of a calyx sclerite. Scale =

0,035 mm. J. Sclerites from interior of peduncle. Scale = 0,03 mm.

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 33. Scanning electron micrographs of sclerites. A-C. Pennatula inflata. From calyx:

0,8 mm; 0,6 mm; 0,8 mm. D-E. Pteroeides isosceles. D. Needle from supporting ray of polyp

leaf: 7 mm. E. Terminal tip of needle from supporting ray: 1,3 mm.

SOUTHERN AFRICA PENNATULACEA 103

Fig. 34. Pennatula inflata. Entire colony; length 220 mm. A. Dorsal view. B. Ventral view.

104 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 35. Pennatula inflata. Two views of distal portion of rachis. Length of rachis portion shown

is 150 mm.

SOUTHERN AFRICA PENNATULACEA 105

Description

The colonies examined are 220 mm and 400 mm in length. Peduncle stout,

rigid, one-third total length of colony, oval in cross-section. Prominent bulb about

midway on peduncle. Rachis spongy and flexible, with naked dorsal tract up to

10 mm in width. This tract is widest in the distal half of the rachis. Polyp leaves

well developed, up to 40 mm in length and 16 mm in width, 26-37 pairs of polyp

leaves per colony. Autozooids 29—42 per polyp leaf, in usually 2 or 3 ill-defined

rows. Calyces tubular, up to 10 mm long, eight-toothed, with teeth equal to

subequal; many calyces with one tooth more strongly developed than the others.

Zooids dimorphic, dense and numerous on two narrow marginal tracts bordering

dorsal face of rachis. Mesozooids with two tooth-like points composed of sclerites,

common in a row along the basal dorsal margin of each polyp leaf. Siphonozooids

without sclerites, crowded in narrow band on rachis above each polyp leaf.

Colonies with 26—37 rows of siphonozooids on each side, representing one row

per polyp leaf. Sclerites numerous; three-flanged needles in calyx, up to 1,2 mm

in length; inconspicuously three-flanged spindles of peduncular surface up to

0,2 mm in length; minute oval plates in peduncular interior 0,015—0,025 mm long.

Colour of peduncle, rachis and leaves yellowish-white to reddish-orange, sclerites

of autozooids and mesozooids orange, siphonozooids white.

Distribution (Fig. 40)

East African coast and south-west coast of South Africa, 457-741 m in

depth. The present study establishes a new record for southern Africa and extends

the range south-west from Somalia to the Namaqualand coast, South Africa. It is

the first record of the species for the Atlantic Ocean. The type locality is Somalia

(Indian Ocean), 628-741 m in depth.

Remarks

Tixier-Durivault (1954: 630) identified these specimens as P. phosphorea.

However, the possession of mesozooids on the dorsal edge of the polyp leaves

proves this to be a misidentification. According to Kikenthal (1915: 82),

P. phosphorea does not possess mesozooids and does not possess more than

20 polyps per mature polyp leaf.

Day (1974b: 35) recorded several unidentified species of the genus Pennatula

from Delagoa Bay, Mozambique.

Of the twelve species of Pennatula considered to be valid by Kikenthal

(1915: 81), six possess between 3 and 20 polyps per polyp leaf—P. aculeata

Danielssen, 1860; P. indica J. A. Thomson & Henderson, 1906; P. murrayi

Kolliker, 1880; P. pearceyi KGlliker, 1880; P. phosphorea Linnaeus, 1758; and

P. prolifera Jungersen, 1904. Six additional species have 30-100 polyps per polyp

leaf—P. fimbriata Herklots, 1858; P. grandis Ehrenberg, 1834; P. inflata

Kiikenthal, 1910; P. moselyi KGlliker, 1880; P. naresi Kolliker, 1880; and

P. rubra (Ellis in Bohadsch, 1761). Of these, only P. inflata possesses a wide

106 ANNALS OF THE SOUTH AFRICAN MUSEUM

naked dorsal tract of the upper rachis with a spongy texture, and many calyces

have one tooth more strongly pronounced than the others.

A certain amount of variation occurs with regard to the calyx teeth in

P. inflata. In the present specimens, the development of the calyx teeth is

variable—many autozooids examined possess eight equal or subequal teeth.

However, in a few polyps, one, two, three, or four of the teeth may be somewhat

larger in size than the remaining teeth; and in many polyps one tooth 1s slightly or

distinctly larger than the other seven teeth.

Family Pteroeididae Kolliker, 1880

Colonies bilateral, possessing well-developed and rigid polyp leaves with long

stout sclerites that are not three-flanged. Siphonozooids crowded in proximal

region of leaves. Autozooids retractile into fleshy distal region of polyp leaves, or

into rounded calyces that are not composed of dense sclerites.

Three genera of the Atlantic and Indo-Pacific: Pteroeides Herklots, 1858;

Sarcoptilus Gray, 1848; and Crassophyllum Tixier-Durivault, 1961a.

Genus Pteroeides Herklots, 1858

Pteroeides Herklots, 1858: 19. K6lliker, 1872a: 124. Gray, 1870: 24. Kiikenthal & Broch,

1911: 395. Ktikenthal, 1915: 97.

Godeffroyia Kolliker, 1869: 222.

Argentella Gray, 1870: 22.

Pteromorpha Gray, 1870: 23.

Crispella Gray, 1870: 25.

Diagnosis

Polyp leaves with supporting rays composed of long needle-like sclerites.

Needles of supporting rays smooth, except for terminal ends that are somewhat

tuberculated. Sclerites of peduncle are rods or spindles.

A genus of over 80 named species, including many uncertain or dubious ones;

Atlantic, Mediterranean, and Indo-Pacific.

Type species. Penna grisea Bohadsch, 1761 (= Pennatula spinosa Ellis, 1764);

Europe.

Pteroeides isosceles Thomson, 1915

Figs 31C—D, 33D-E, 36, 37, 38

Pteroeides isosceles J. S. Thomson, 1915: 17, pl. 1.

Material

SAM-—H1207, off East London (33°00’S 28°00’E), 58-69 m, 28 November

1906, 1 colony, large trawl, S.S. Pieter Faure Survey, P.F. 19061 (holotype).

SOUTHERN AFRICA PENNATULACEA 107

Redescription of the holotype

The colony is 170 mm long and up to 40 mm in width. Peduncle 18—20 mm

in diameter and slightly more than half the length of the colony. Rachis with

22 pairs of polyp leaves. Larger leaves 20 mm in width by 35 mm in length,

15-18 rays per leaf in leaves from the middle region of the rachis. Rays extend

through the entire length of a particular leaf and may extend 1-2 mm beyond the

margin. Rays not visible in the basal plate of siphonozooids. Autozooids

restricted to the outer margin of each leaf to 5 mm below the margin on both the

upper and lower surfaces of each leaf; retractile directly into pockets of the fleshy

leaf margin. Autozooid zone relatively free of smaller spindles. The autozooids

are not arranged in distinct rows, but 5—8 autozooids are usually scattered

between two adjacent rays. Mesozooids not apparent. Siphonozooids are densely

arranged and restricted to a plate about 12 mm broad in the proximal region of

each leaf. Needles of rays up to 10 mm long. Spindles of dorsal surface of polyp

leaves 0,2—0,4 mm in length. Rods and spindles of peduncular surface are

longitudinally grooved but not distinctly three-flanged, 0,10—0,17 mm long.

Sclerites of peduncular interior absent. Peduncle gray, leaves cream with

yellowish siphonozooids and white rays.

Distribution (Fig. 38)

Known only from the type locality off East London, South Africa, 58-69 m

depth.

Remarks

This species can be differentiated from other southern African sea pens by

having autozooids without calyces united into thick, fleshy polyp leaves,

11-18 conspicuous rays per leaf, and siphonozooids densely set but restricted to

a plate near the base of each leaf.

Remarks on the genus Pteroeides

Recent collections from northern Zululand have yielded several colonies of

a species distinct from P. isosceles, which has not been identified. Several

damaged or presumably young colonies of Pteroeides from Natal are also present

in the South African Museum collection that can not presently be identified to

species but seem to be distinct from P. isosceles. It is likely that several species of

the genus can be expected to occur in southern African waters. The genus is in

much need of revision; well over 80 specific names can be found in the literature

at present.

Broch (1958: 278) recorded Pteroeides griseum (Bohadsch, 1761) from off the

Angola/Namibia border region at 80-91 m. This is just north of the geographic

range of the present work, but is mentioned since the species may be encountered

further south. It is distinguished from P. isosceles by having numerous spindles in

the autozooid zone, independent of the ray needles. In P. isosceles the ray

108 ANNALS OF THE SOUTH AFRICAN MUSEUM

TN . ; :

Fig. 36. Pteroides isosceles. A. Entire colony, ventral face. Scale = 60 mm. B. Detail of

margin of polyp leaf, upper surface. Scale = 3 mm. C. Entire polyp leaf, lower surface. Scale =

10 mm. D. Peduncular surface sclerites. Scale = 0,1 mm. E. Ray needles from polyp leaf.

Scale = 2 mm.

SOUTHERN AFRICA PENNATULACEA — 109

Fig. 37. Pteroeides isosceles. Entire colony, length 170 mm. A. Ventral view. B. Dorsal face.

needles are pointed and similar at both ends; the autozooid margins may have

only scattered sclerites that differ from the ray needles; the siphonozooid and

autozooid zones have a distinct boundary.

J. S. Thomson (1915: 17) described a damaged colony from the type locality

of Pteroeides isosceles as Pteroeides sp. An examination of this specimen has

shown that it closely resembles the holotype of P. isosceles and thus the two

specimens are very likely conspecific.

110 ANNALS OF THE SOUTH AFRICAN MUSEUM

MAPUTOA RE

INDIAN

OCEAN

EAST LONDON,

PORT ELIZABETH

Fig. 38. Map of south-eastern Africa showing known collecting stations for twelve pennatu-

lacean species: Amphiacme abyssorum (@); Calibelemnon sp. (@); Cavernularia dayi (A);

Cavernularia lutkenii (¥); Cavernulina sp. (*); Chunella gracillima (+); Funiculina quadran-

gularis (©); Kophobelemnon sp. ((@); Pteroeides isosceles ({_}); Scleroptilum sp. (®); Veretillum

leloupi and V. cynomorium (§).

ATLANTIC SOUTHERN AFRICA

OCEAN

Fig. 39. Map showing known distribution of the family Echinoptilidae: Actinoptilum molle (@);

Echinoptilum echinatum (&); Echinoptilum macintoshii (¥); Echinoptilum asperum, E. elong-

atum, E. minimum, E. roseum (x).

SOUTHERN AFRICA PENNATULACEA 111

ANGOLA

NAMIBIA

Walvis Bay

®)\Luderitz

SOUTH AFRICA

ST. HELENA INDIAN

BAY OCEAN

LANGEBAAN

LAGOON

35 ATLANTIC OCEAN

Fig. 40. Map of south-western Africa showing known collecting stations of eight pennatulacean

species: Anthoptilum grandiflorum (A); Cavernularia elegans ((@); Distichoptilum gracile ({_]});

Halipteris africana (A); Kophobelemnon stelliferum (+); Pennatula inflata (J); Umbellula

lindahli (@); Umbellula thomsoni (CE); Veretillum cynomorium (Q).

MOZAMBIQUE

NAMIBIA

MAPUTOS @

e@) LUDERITZ

DURBAN/@

ATLANTIC INDIAN

OCEAN SOUTH AFRICA OCEAN

EAST 6

LONDON¢

D) eo

PORT

ELIZABETH

t )

)

PCAPE TOWN

19 23

Fig. 41. Map showing known collecting stations for the family Virgulariidae (subfamily

Virgulariinae) in southern Africa: Scytaliopsis djiboutiensis (QO); Virgularia gustaviana ()

Virgularia mirabilis (A); Virgularia schultzei (@); Virgularia sp. ((_]).

fA ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks on the genus Crassophyllum

Tixier-Durivault (1961a: 428) described a new genus and species, Crasso-

phyllum cristatum, from northern Angola. It is mentioned here since it may

possibly be encountered further south. This form is distinguished by the lack of

supporting rays in the polyp leaves, but possesses scattered spindles and needles

in the region between the siphonozooid plate and the polyp leaf margin.

DISCUSSION

ZOOGEOGRAPHY

The littoral and sublittoral benthic regions of southern Africa constitute a

remarkably diverse and biogeographically unique area. The region is made up of

cold-water and warm-water temperate elements, as well as a tropical component.

In particular, the southernmost coast (the region between the Cape Peninsula and

Port St. Johns) has a high degree of endemism in the invertebrate fauna (Ekman

1953: 189; Day et al. 1970: 2). This region is referred to as the ‘Agulhas Province’

by Briggs (1974: 149), the ‘Cape Fauna’ by Stephenson & Stephenson (1972: 101),

and the ‘South Coast’ fauna by Ekman (1953: 187) and Millard (1978).

The peculiarities of this region and the composition of the fauna are due to

a number of factors. Most important of these are: (1) the isolated, peninsular

nature of southern Africa, which is bounded by two oceans, and the extensive

area of continental shelf known as the Agulhas Bank, which is a probable centre

of adaptive radiation (Millard 1978, fig. 1); and (2) the convergence of oceanic

waters of divergent temperatures and origins in the vicinity of the Cape Peninsula,

via the colder Benguela Current of the South Atlantic and the warmer Agulhas

Current of tropical Indian Ocean influence (Branch & Branch 1981: 14).

A zoogeographical analysis of the southern African octocoral fauna will be

dealt with in a subsequent paper.

AFFINITIES OF THE FAUNA

Shallow-water benthic invertebrates of southern Africa are generally

recognized to have four major zoogeographical components: temperate Atlantic,

Subantarctic, Indo-Pacific, and endemic (Millard 1978; Gosliner 1987). Many

deep-water species have more scattered or cosmopolitan distributions. Several

species of the west and south coasts, such as Lituaria valenciennes, extend further

north into the Mediterranean Sea. Similarly, a number of species of the east coast,

such as Calibelemnon indicum, have wider distributions in the tropical. Indo-

Pacific. At least 20 species of hydroids from southern Africa have Antarctic or

Subantarctic affinities (Millard 1978: 171-172). Gosliner (1987), in considering

the sister relationships in opisthobranch molluscs, regarded the Subantarctic

influence as perhaps the oldest influence, which may be associated with the

break-up of Gondwanaland. In regard to sea pens, the cosmopolitan Umbellula

lindahli is known from throughout Antarctic waters (Grasshoff 1982a: 753,

SOUTHERN AFRICA PENNATULACEA 113

map 11). However, no truly Subantarctic component is presently recognized

within the southern African fauna. Endemic species, such as Virgularia schultzei

and Actinoptilum molle, are found throughout the region, with the centre of

distribution along the south coast (the region between the Cape Peninsula and the

Transkei coast).

SUMMARY AND CONCLUSION

The present study is the first in a two part series of a comprehensive survey

of pennatulaceans from the subcontinent of southern Africa. It has increased in

the number of identified species in the region from 12 to 23, the number of genera

from 10 to 20, and the number of families from 8 to 12.

The families recorded from southern Africa for the first time are the

Kophobelemnidae, Protoptilidae, Scleroptilidae, and Chunellidae. The eleven

species newly recorded for the region are Cavernularia elegans, Echinoptilum

macintoshii, Echinoptilum echinatum, Kophobelemnon stelliferum, Disticho-

ptilum gracile, Chunella gracillima, Amphiacme abyssorum, Umbellula lindahli,

Scytaliopsis djiboutiensis, Halipteris africana, and Pennatula inflata. The genera

Cavernulina, Sclerobelemnon, and Calibelemnon are recorded for the first time

from the region.

ACKNOWLEDGEMENTS

I extend my appreciation to Dane Gerneke and Klaus Schultes of the Uni-

versity of Cape Town Electron Microscope Unit for assistance in the use of the

Cambridge S180 and S200 scanning electron microscopes.

I am grateful to William Liltved (South African Museum), Philip Coetzee

(University of Port Elizabeth), Richard Kilburn, Ruth Fregona, and David

Herbert (Natal Museum), and Roy Melville-Smith, Robert Leslie, and Paul

Williams (Sea Fisheries Research Institute) for field assistance and collection of

specimens.

Elizabeth Hoenson and Michelle van der Merwe provided curatorial assist-

ance. The late Dr A. H. B. De Decker assisted in the translation of some

literature. Marcelle Scheiner and Sandra Saven typed the manuscript. Elizabeth

Louw and Sheryl Ozinsky aided in correcting the manuscript.

I thank Professor G. M. Branch and Dr J. A. Day of the Department of

Zoology, University of Cape Town; Dr M.-J. D’Hondt of the Museum National

d’Histoire Naturelle, Paris; and Dr M. Grasshoff of the Forschungsinstitut Sen-

ckenberg, West Germany, for their helpful comments.

Partial financial support for this work was provided by the Council for

Scientific and Industrial Research, Research Grants Division.

Part of the material examined was procured by the South African Museum

and Natal Museum during dredging and trawling cruises on board the

R.V. Meiring Naude between 1975 and 1985.

114 ANNALS OF THE SOUTH AFRICAN MUSEUM

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118 ANNALS OF THE SOUTH AFRICAN MUSEUM

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Royal Indian Marine Survey Ship Investigator in the Indian Ocean. Part I. The alcyonarians

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Indian Marine Survey Ship Investigator in the Indian Ocean. II. The alcyonarians of the

littoral area. Calcutta: Trustees of the Indian Museum.

Tuomson, J. S. 1915. The Pennatulaceae of the Cape of Good Hope and Natal. Memoirs and

Proceedings of the Manchester Literary and Philosophical Society 59 (1): 1-26.

THomson, J. S. 1917. The occurrence of Cavernularia Liitkenii KOll. in the Seas of Natal.

Memoirs and Proceedings of the Manchester Literary and Philosophical Society 62 (3): 1-5.

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Transactions of the Royal Society of South Africa 11 (1): 45-84.

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TIXIER-DURIVAULT, A. 1960. Les octocoralliaires de I’Ile Inhaca. Bulletin du Muséum national

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(Alcyonacea, Pennatulacea). Campagne da la ‘Calypso’: Golfe de Guinée. Annales de

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1945-1946 7: 63-76.

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national dhistoire naturelle (2) 37 (4): 705-716.

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de Madagascar 21: 1-456.

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avoisinantes. Téthys, supplément 3: 11-68.

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Palaeontographical Society.

SOUTHERN AFRICA PENNATULACEA 119

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to other institutions in exchange, with annotations. Bulletin of the Museum of Comparative

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Expedition, under Commodore C. Ringgold and Captain John Rogers, U.S.N., from 1853

to 1856. Collected by Dr. Wm. Stimpson, naturalist of the Expedition. With descriptions of

some additional species from the west coast of North America. Proceedings of the Essex

Institute, Salem, Mass. 4: 181-196.

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corals and polyps of the west coast of America. Transactions of the Connecticut Academy

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North America. No. 3. American Journal of Science and Arts (3) 17: 239-243.

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southern coast of New England, No. 7, and of some additions of the fauna of Vineyard

Sound. American Journal of Science and Arts (3) 24: 360-364.

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northern coast of the United States in 1883. Report United States Commissioner of Fisheries

1883: 503-699.

WEINBERG, S. 1976. Revision of the common Octocorallia of the Mediterranean circalittoral.

I. Gorgonacea. Beaufortia 24 (313): 63-104.

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Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

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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.

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GARY C. WILLIAMS

THE PENNATULACEA OF

| SOUTHERN AFRICA

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‘OF THE SOUTH AFRICAN |

KUM

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

FiscHER, P. H. 1948. Données sur la résistance et de 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 99 Band

May 1990 Mei

Part 5 Deel

FORAMINIFERA FROM THE LATE

PLEISTOCENE (LATEST EEMIAN TO

EARLIEST WEICHSELIAN)

SHELLY SANDS OF CAPE TOWN CITY

CENTRE, SOUTH AFRICA

I. K. McMILLAN

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

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Gay tai), HEA), 8 TAD, 7), 1OM=9),, ING, 5, 7, tia),

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33, 36(2), 43(1), 45(1), 67(5), 84(2)

Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 111 3

Printed in South Africa by In Suid-Afrika gedruk deur

The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk.,

Court Road, Wynberg, Cape Courtweg, Wynberg,

C632

FORAMINIFERA FROM THE LATE PLEISTOCENE

(LATEST EEMIAN TO EARLIEST WEICHSELIAN)

SHELLY SANDS OF CAPE TOWN CITY CENTRE, SOUTH AFRICA

I. K. McMILLAN

Department of Geology, University College of Wales,

Aberystwyth, Dyfed, United Kingdom*

(With 20 figures and 4 tables)

[MS accepted 29 March 1989]

ABSTRACT

Samples from shelly sands exposed during deep trenching in Cape Town city centre have

yielded diverse and relatively well-preserved foraminifera faunas. Fifty-three species are

recognized, of which four are reworked from older deposits; none are new. The total assemblage

is very similar to ones from off the mouth of the Orange River, substantially further north. From

foraminiferal and geological considerations, a latest Pleistocene (latest Eemian to earliest

Weichselian) age is proposed for the sequence, which is regarded as having acccumulated during

the fall in sea-level at the end of the Eemian. The foraminifera lived in a wave-dominated, littoral

environment, and the clean sands in which they occur are considered to be part of an extensive,

regressive sheet sand that originally covered almost the entire continental shelf off all three coasts

of South Africa, and which is still widely preserved. Similar shelly sands, also attributable to the

latest Eemian—earliest Weichselian sea-level fall, occur in the coastal parts of both the

‘Bredasdorp Formation’ and the ‘Alexandria Formation’ (both termed Unit III), as well as along

the Gamtoos coast (Unit III), at Durban (the Bluff Sandstone Formation), and extensively over

the Zululand coastal plain (the Uloa Formation). Variable diagenesis has resulted in considerable

cementing of these deposits at some localities. The foraminifera of the same sequence at several

localities on the west coast are given in three appendices: for the Llandudno—Sandy Bay coast,

Cape Peninsula, the Berg River mouth, and the Hoedjiespunt Peninsula, Saldanha.

CONTENTS

PAGE

TNCKOGIC HO Meni rarer a eet ree ncaa « leales Ws WPetdnin tus 122

JPRREAIOIOS WHOS oreo G.c dhe ei 0.5 CRO ONG Ee UCI Re a a 122

hcworaminiieraassemblAGeSine amass ese) sles Sele wot sede anor 125)

EVIL One ntzOMmdepOstiOnesi we. is oe dae oe eee ee oes See 125

Stratigraphy andeconmelatiomene: see sei acide Se Cac e ean as 126

Synchronous deposits of the South African continental margin..... 126

Companisoniwitholder Pleistocene deposits .44-.4.55-6- 54045000" 129

Companisoniwith Holocene depositsinn sn.) ons ssse5 ase ea se 3h

INC WOK Speman yeny ter. er en tnta uve PG ees Ahan Gan Wate clea uaa aes 1132

EALOCESS TMS MMIC LAOS Agta rae cei pet keels ecaiestin era tuniresos Soi Jiang, Sake ue oes 133

Rhotographysandsdepositioniotmatenialtes s+ ee eoe ano see oe 1133)

FOKAMMMILCEAM AKON OMe ne emcee eee oh ikk jase tah a staels ee cole on 135

NCKMOWLe AS CINE MUS ameristar ids eth, claasv etal alee SRS 173

INC IC KCNC SMB rye atte race (ke Menace ac Gade weno gai ath Nea ee a 174

* Present address: SOEKOR (Pty) Ltd, P. O. Box 307, Parow 7500, South Africa.

WA).

Ann. S. Afr. Mus. 99 (5), 1990: 121-186, 20 figs, 4 tables.

Oy ANNALS OF THE SOUTH AFRICAN MUSEUM

PAGE

Appendix A: The Llandudno-Sandy Bay coast, Cape Peninsula....... 179

Appendix B: Latest Pleistocene Foraminifera from Berg River mouth . . 182

Appendix C: Latest Pleistocene Foraminifera of Hoedjiespunt Peninsula,

Salama. 5252 se oe Ree he EN ct ns oe eo eee 184

INTRODUCTION

During deep trenching by the Municipality along several of Cape Town’s city

centre streets in late 1986, clean white, rather bleached, shelly quartz sands were

exposed. These deposits appear to be confined to the more seawards parts of the

city (Fig. 1). The sands are distinctly bedded, and undisturbed; human

interference is confined to a distinct zone overlying the sands. Despite the clean

and somewhat coarse nature of the sands, studied samples yielded a fairly

well-preserved and diverse foraminifera fauna that contains a number of

age-diagnostic species. The whole assemblage is almost identical to that described

by McMillan (1987b) from the Late Pleistocene (latest Eemian to earliest

Weichselian) deposits occurring on the inner Orange Shelf, just to the north of the

Orange River mouth. A summarized stratigraphic column, based on observaticns

by Mr V. H. Valicenti of SOEKOR (Pty) Ltd, Cape Town, at the time the trench

was open, is given in Figure 2. The total thickness of the Cape Town sequence is

not known. Two samples each were collected at the corner of Hertzog Boulevard

and Oswald Pirow Street, and at the corner of Oswald Pirow and Martin

Hammerschlagweg. Results of the foraminifera study are given in Table 1.

PREVIOUS WORK

Studies of Quaternary foraminifera in South Africa were initiated by

Chapman (1907), who examined samples from a drill hole in the bed of the

Buffalo River, East London. The precise age of the sequence, studied to a depth

of 33,2 m below the bed of the river, is probably partly of Holocene, but mostly

of latest Eemian—earliest Weichselian age. The latter age is indicated by the

presence of Elphidium crispum (Linné) (given as Polystomella crispa by

Chapman) through most of the studied sequence, suggesting that the Holocene is

less than 4,57 m thick. Further comments on the use of Elphidium crispum as a

zone fossil for the latest Eemian to earliest Weichselian deposits of South Africa

are given below in the section on stratigraphy (see p. 129).

Most later work has involved lists of foraminifera species only, but Biesiot

(1957) illustrated a number of the in situ foraminifera of the Uloa Formation at its

type locality (see McMillan 1987a); these include the species of Amphistegina,

Elphidium, and perhaps some of the miliolids, together with Eponides zuluensis

Biesiot, ?Eponidella sp. sensu Biesiot, Rotalia beccarii sensu Biesiot, Cibicides

lobatulus (Walker & Jacob), and possibly Nonion elongatum sensu Biesiot.

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 123

18°25'E

(e}

cae

«za

(0) J 5

CAPE TOWN

City Centre s 33°56'S

Fig. 1. Location of studied samples from the latest Eemian—earliest Weichselian regressive

shelly sands (Unit III) beneath Cape Town city centre. Samples 1 and 2 are from the corner of

Oswald Pirow Street and Hertzog Boulevard; 3 and 4 from the corner of Oswald Pirow Street

and Martin Hammerschlagweg. Based on 1 : 50 000 topographic sheet 3318CD (Cape Town),

4th edition.

Street level

Rubble and soil

later than

1693 A.D.

Latest Eemian- earliest

Weichselian Bedded and Shelly Sands

Fig. 2. The stratigraphy of exposures in deep trenches in Oswald Pirow Street, Cape Town city

centre, as measured in November 1986, by Mr V. H. Valicenti, SOEKOR (Pty) Ltd. Sampled

horizons are indicated (CTC 1 to 4).

124 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 1

Distribution of foraminifera in studied samples from the latest Eemian—earliest Weichselian

sands of Oswald Pirow Street, Cape Town city centre.

Sample number Sp. Species

CTC-1 CTC-2 CTC-3 CTC-4 no.

1 1 1 = Spiroplectammina sp.

1 1 2 Quinqueloculina contorta

8 6 18 28 3. Quinqueloculina dunkerquiana

Z 4 Quinqueloculina isabellet

2, 2, 5 Quinqueloculina cf. Q. lata

1 1 1 >) 6 Quinqueloculina seminulum

11 7 Quinqueloculina triangulans

1 Z 8 Quinqueloculina cf. Q. undulata

4 6 24 34 9 Quinqueloculina spp. (broken)

y 4 10 Siphonaperta sp.

2 1 7 3) 11 Milolinella subrotunda

1 12 ?Dentalina sp. (reworked)

1 13. Lagena semilineata var.

1 14 Lenticulina sp.

1 15. = Oolina squamososulcata

1 3. 16 Oolina sp. A

1 17 _—‘-Fissurina lucida

1 18 = Fissurina cf. F. marginata

2 19 Fissurina spp. (reworked)

1 1 20 Guttulina spp. (broken)

1 21 Bulimina marginata

1 22 Uvigerina sp. (reworked)

1 23 Trifarina angulosa

1 24 ~Bolivina sp.

1 25. Brizalina spathulata

16 8 26 Cassidulina laevigata

1 27 ~~ Cassidulina crassa s.1.

1 28 ?Gavelinopsis sp. (reworked)

1 29 Planulinoides biconcavus

1 1 30 = @Rosalina bradyi

1 31 =Rosalina sp.

2 dip) 10 32 Glabratella australensis

3) 33 Cibicides fletchen s.1.

5 a, 24 13. 34 ~=©Cibicides lobatulus

4 4 35 Cibicides spp. (reworked)

y 5 y) 3. 36 = Cibicidoides sp.

1 37. Ayalinea balthica

1 1 1 38 Sphaerogypsina globulus

1 39 = ?Acervulina sp.

14 3. 40 Elphidium advenum

6 1 41 Elphidium sp. A

1 4 42 — Elphidum articulatum

54 26 131 137 = 43, —Ss Elphidium crispum s.1.

2 6 130 44 -~— Elphidium macellum s.1.

1 45 Elphidium sp: B

1 46 Astrononion echolsi

9 2 47 Ammonia japonica

1 i 3 6 48 Ammonia parkinsonia s.1.

11 16 105 146 49 Pararotalia nipponica

1 50 indeterminate reworked rotalid

51. Globigerina (G.) bulloides

Globorotalia (G.) inflata

53 Neogloboquadrina cf. N. pachyderma

91 V2 429 452 Total number of tests

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 125

All the remaining fossils detailed by Biesiot (1957) are reworked from the Early

and Middle Miocene (McMillan 1987a).

Lists of foraminifera from the coastal ‘Bredasdorp Formation’ and the coastal

‘Alexandria Formation’ were given by Rtmke, in Spies etal. (1963) and

Engelbrecht et al. (1962). Some of the samples described by Parr (1958),

including those from the Bluff at Durban, are also of this age.

It is now certain that the Pararotalia cf. P. inermis (Terquem) emend. Le

Calvez, and the Ammonia beccarii (Linné) var. 2 sensu McMillan (1974), from

the Agulhas Bank, are also of latest Eemian—earliest Weichselian age. The

former can now be referred to Pararotalia nipponica (Asano), whereas the latter

are poorly preserved tests of the Ammonia sp. C (Alexandria) detailed by

McMillan (1987a).

McMillan (1987a) illustrated Ammonia species from a variety of South

African Pleistocene, Holocene and Pliocene localities as a preliminary attempt to

correlate the many, geographically disparate deposits of these ages. McMillan

(1987b) described in detail the stratigraphy and foraminiferal assemblages of the

latest Eemian-—earliest Weichselian and the Holocene silts and sands of the

Orange Shelf, southern Namibia continental margin. Diagnostic zone foramini-

fera from the different aged rock units that comprise the ‘Alexandria Formation’

have been illustrated by McMillan (in press).

THE FORAMINIFERA ASSEMBLAGES

All four Cape Town samples yielded faunas dominated by the benthonic

species Elphidium crispum (Linné) s.l. and Pararotalia nipponica (Asano). As is

the case with all of the Pleistocene marine sand deposits preserved on the South

African continental margin, planktonic foraminifera are rare. Species of the

benthonic genera Elphidium, Ammonia, Pararotalia and Cibicides predominate,

again in keeping with other Pleistocene marine sequences.

ENVIRONMENT OF DEPOSITION

A littoral, wave-influenced environment of deposition is indicated from both

the foraminifera and the sediments. The clean nature of the sands, the abrasion

and breakage of some foraminifera tests, and the presence of a number of

foraminifera species that in life attach to a substrate, all clearly reflect a turbulent

sea-floor, with highly oxygenated conditions both on the sea-bed and in the water

column. A water depth of 10 m or less is suggested. The attached species include

Cibicides lobatulus (Walker & Jacob), Rosalina sp., Rosalina bradyi (Cushman),

Glabratella australensis (Heron-Allen & Earland) and Sphaerogypsina globulus

(Reuss), and possibly Cibicides fletcheri Galloway & Wissler s.1. The macrofaunal

remains in the sands, mainly bivalve and echinoid debris, are substantially

comminuted.

126 ANNALS OF THE SOUTH AFRICAN MUSEUM

It appears likely that the foraminifera tests, while evidently in situ in a

stratigraphical sense, were much transported before coming to their final rest. It

seems likely that these foraminifera lived in a sublittoral environment, possibly a

kelp-bed, and were swept shorewards after death by wave action, to be

incorporated into truly littoral deposits accumulating as the shoreline retreated.

The retreat of the sea at this time would seem to have occurred over a fairly short

interval (perhaps well within 5 000 years, if an analogy with the Holocene

transgression holds good), so that the littoral sands laid down at any one locality

may have soon no longer been subject to wave- and swell-induced abrasion, lying

instead in coastal sand flats or dunes.

The presence of Ammonia parkinsoniana (d’Orbigny) s.1. in small numbers

in the Cape Town samples suggests that the assemblage, if regarded as from one

environment, derives from an area of slightly reduced salinity conditions. Thus,

river output or other run-off affected the habitats of the studied foraminifera, but

the lack of land-derived plant debris, seeds, charophyte oogonia and so on,

together with the relatively small numbers of Ammonia parkinsoniana, clearly

indicate the freshwater influence to have been an insubstantial one, perhaps as is

the case in the Cape Town part of Table Bay at the present day. It seems that a

comparison of numbers of Ammonia japonica (Hada), typical of shelf

environments, with numbers of Ammonia parkinsoniana, characteristic of

estuarine and littoral environments, may provide a simplified environmental

indicator for these latest Pleistocene accumulations around the west and south

coasts of South Africa.

Since most of the foraminifera species encountered in the Cape Town sands

are known to live off the western Cape coast at the present day, it would appear

that sea temperatures at the end of the Eemian were much the same as they are

at present. However, until the full geographic ranges of the various species

around the South African coast are better known, an improved understanding of

water temperatures cannot be attempted.

STRATIGRAPHY AND CORRELATION

The foraminifera of the Cape Town sands show varying similarities to those

of older Pleistocene deposits, those of deposits of the same age, and those of the

Holocene. Generalized locations are given in Figure 3.

SYNCHRONOUS DEPOSITS OF THE SOUTH AFRICAN CONTINENTAL MARGIN

The Cape Town shelly sands can be correlated with ease to the sands and silts

occurring to a considerable thickness (about 70 m according to Hoyt et al. 1969)

off the mouth of the Orange River, based on the very close similarity of their

foraminiferal faunas (see McMillan 1987b). Correlation is also possible with a

number of deposits preserved about 8 m above high sea-level at the Hoedjiespunt

Peninsula, Saldanha, along the Sandy Bay to Llandudno coastline, Cape

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 27

MAPUTO

% WA

-4~—=Jc holes

Bluff Sst.

Formation

Aliwal Shoal

HOEDJIESPUNT

CAPE TOWN

LLANDUDNO- ‘Alexandria Formation’

=o a a ~ PLETTENBERG BAY

Bredasdorp Formation ~___

Fig. 3. Distribution of latest Eemian—earliest Weichselian deposits around the South African

coast, of equivalent age to the shelly sands of Cape Town centre. The areas shown for the

Varswater Formation, the ‘Bredasdorp Formation’, and the ‘Alexandria Formation’ are for the

entire sequence, Unit I to Unit III. In all these areas Unit III occurs as a narrow strip along the

present coast. The major foraminiferal facies, based mainly on Ammonia and Elphidium species

are also shown. Species listed distinguish the Unit III sands from both older Pleistocene and

Holocene deposits.

Peninsula, and below the level of the present Berg River mouth (see Appendices

A, B, C).

Deposits of the same age also occur along the south coast of South Africa: the

latest, most coastal portion of the ‘Bredasdorp Formation’ between Cape Agulhas

and Mossel Bay, below the mouth of the Keurbooms and Bietou rivers at

Plettenberg Bay and onshore adjacent to the estuary, and the latest, most coastal

portion of the ‘Alexandria Formation’ between Port Elizabeth and Port Alfred, as

well as the whole of the smaller area of ‘Alexandria Formation’ along the

Gamtoos coast. Deposits of this age have been termed Unit III informally by the

present author (see McMillan in press). It is now known that Unit III and Unit IV

of the ‘Alexandria Formation’ are slightly different facies of the same rock

sequence, and are of the same age (see McMillan in press). Unit III in the eastern

Cape is characterized by polished foraminifera tests, with many normal marine

shelf species, and the distinctive Ammonia sp. C (Alexandria) is common and

widespread. Unit IV, seen only at Woody Cape, contains foraminifera assem-

blages dominated by miliolids, together with Poroeponides lateralis (Terquem),

that derive from a more exclusively littoral environment.

Off the south coast, Unit II] sediments are extensively preserved as a thin

veneer over the majority of the Agulhas Bank. The sequence has been frequently

128 ANNALS OF THE SOUTH AFRICAN MUSEUM

encountered in borehole tops as far offshore as the outermost shelf, down to a

water depth of about 200 m. Foraminifera faunas remain uniform and distinctive

throughout, and are easily correlated with those deposits occurring onshore at the

present day. At borehole F—A 13, the latest Eemian—earliest Weichselian shelly

deposits, together with the overlying Holocene interval (the two are separated by

an unconformity), are no more than 0,5 m thick. Ammonia species from F—A 13

and E-B 1 have been previously discussed by McMillan (1987a). Where the latest

Eemian-—earliest Weichselian Unit III deposits unconformably overlie the shelly

white limey clays of the Early Miocene (Burdigalian) over most of the middle and

outer shelf on the Agulhas Bank, much reworking of the Miocene has occurred,

so that the latest Pleistocene sediments are often white clays, with abundant

Miocene foraminifera in some cases.

The thin sliver of coastal deposits preserved along the East London to

Durban coast have only been examined for foraminifera by Parr (1958); the

species listed by him from near Bats Cave, East London, are insufficient to

determine what part of the Pleistocene (but presumably Unit III) is represented

here. It seems certain now that most, possibly all, of the Bluff Sandstone

Formation at Durban and the Uloa Formation, extensively developed over the

Zululand coastal plain, are of the same age—latest Eemian to earliest

Weichselian. Although very much thinner offshore, the sequence also occurs in

the borehole tops of Jc-A 1, Jc-B 1 and Jc—C 1, where it lies unconformably

between the Holocene and Middle or Early Miocene, and consists of a loose

shelly sand. The eastern offshore sequence thus exactly duplicates that seen on

the Agulhas Bank. Again, there is little change in the littoral, benthonic

foraminifera of this unit, and Ammonia sp. C (Uloa) (possibly referable to

Challengerella persica Billman, Hottinger & Oesterle), Elphidium crispum

(Linné) s.1., Planorbulinella larvata (Parker & Jones) and Cellanthus craticulatus

(Fichtel & Moll) have proved useful local indicators for Unit HI from Durban

northward.

Earlier comments on the age of the Uloa Formation have been made by

McMillan (1987a), who suggested reasons for a probable middle Pleistocene age.

However, the presence of Elphidium crispum s.1., together with feathered-suture

Ammonias in the east-coast and the south-coast sequences, as well as their

unconsolidated nature on the outer shelf off both the east and south coasts, clearly

argues for the two to be of the same age. Since the offshore deposits remain

unconsolidated for the most part, it is clear that they have never been subjected

to a major regression of the sea and the surf-zone, and must thus date from the

last fall in sea-level at the end of the Eemian. It is clear from study of the

Holocene sequences around South Africa that the early Holocene transgression,

in strong contrast, was a very mild event in terms of its erosive power.

On the basis of the presence of Elphidium crispum (Linné), the Aliwal Shoal,

south of Durban (Carter 1966; McCarthy 1967), would appear to be also of

Unit III age.

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 129

In all of the above rock sequences, correlation of this youngest regressive

event can be achieved in part by its contained littoral foraminifera, particularly

the species of Elphidium and Ammonia, and partly by its stratigraphical

relationships. Although, on following Unit III around the continental margin

of South Africa, the foraminifera faunas change substantially, nevertheless

with sufficient density of samples, key species can be traced until they overlap

with other key species, and correlation is thus maintained. Elphidium crispum

(Linné) s.l. has proved the most reliable species for identifying Unit III,

though there are subtle differences in test morphology between east- and

west-coast examples that remain to be defined. The major foraminiferal facies

recognized in Unit III deposits so far, together with Jocal age-diagnostic

species, are shown in Figure 3.

McMillan (1987a) has proposed the use of different Ammonia species to

identify the various latest Cainozoic (Pliocene, Pleistocene and Holocene)

marine deposits of southernmost Africa. Subsequent unpublished work on the

Elphidium species has indicated that they are morphologically more diverse

and often more distinctive than the Ammonia species, and thus would seem to

be a more useful group for correlation purposes. Perhaps in response to the

changing environments available for occupation during high (glacial minima)

and low (glacial maxima) sea-level stands by species of the two genera, it

would seem that appearances of new species are more frequent than in the

planktonic foraminifera (Globigerinacea) over the timespan of the Quater-

nary. Particularly in Ammonia, but also in Elphidium, the historical view has

been that species of these genera show much morphological variation in their

tests. On the contrary, intraspecific variation is as limited as in most other

rotalid species, but the number of species in both genera is unusually large,

and most appear to have survived for relatively short time periods, often

markedly less than one million years in duration.

COMPARISON WITH OLDER PLEISTOCENE DEPOSITS

The regressive littoral sands of the end of the last interglacial can be

identified as such in a number of ways. From a foraminiferal point of view, they

contain the greatest number of species in common with extant littoral assemblages

around South Africa. The foraminifera of the earlier Pleistocene and Pliocene

deposits show a progressive decline in similarity with increasing age. A typical

example of this is given by McMillan (in press) for the ‘Alexandria Formation’. In

both the oldest, Pliocene part of the ‘Alexandria Formation’, and the Pliocene of

the Orange Shelf (McMillan 1987b), the benthonic foraminifera show no

similarity with living South African species, though in deeper-water assemblages

some conservative forms persist (e.g. Pullenia, Gyroidina).

Secondly, the latest Eemian—earliest Weichselian deposits can be recognized

because they consist of unconsolidated or poorly cemented shelly sand, up to

86 m thick in the coastal ‘Alexandria Formation’, but often less than 1 m, that

130 ANNALS OF THE SOUTH AFRICAN MUSEUM

covers most of the South African continental margin, though there are large areas

where it is missing off the west coast. This sheet sand has suffered relatively little

from later erosion. In contrast, relatively litthe now remains of the earlier

Pleistocene regressive deposits, the remnants usually being unusually thick

because of local, mild subsidence prior to deposition, and uplift after deposition

has removed them from the erosive activities of subsequent regressions. Earlier

Pleistocene deposits of this type are known from the west coast, especially in the

vicinity of Saldanha and Langebaan, and in the older ‘Bredasdorp Formation’ and

‘Alexandria Formation’, inland from the present coast. It seems probable that the

older, higher terraces (A to C) of the Oranjemund to Chameis Bay coast in

southern Namibia should be included here, but foraminiferal study of them has

not yet been attempted.

Thirdly, there is a tendency for the Unit III shoreline at maximum advance

in the Eemian to be similar to the present shoreline, with the only major differ-

ence occurring on the Zululand coastal plain, which was entirely inundated during

the Eemian. With the older Pleistocene and Pliocene deposits, a progressive

divergence away from the present shoreline can be seen, which by Pliocene times

was quite substantial, notably in the eastern Cape.

Off the south and west coasts of South Africa, the marine deposits of the

latest Eemian to earliest Weichselian can be distinguished from older units by the

particular presence of Elphidium crispum (Linné) s.l., as well as Elphidium

macellum (Fichtel & Moll) s.l., Ammonia japonica (Hada), and Ammonia

parkinsoniana (d’Orbigny) s.l. (see McMillan 1987a, 1987b). In addition,

Astrononion echolsi Kennett, Quinqueloculina isabellei d’Orbigny, Oolina sp. A,

Elphidium cf. E. advenum (Cushman) and Cassidulina crassa d’Orbigny s.1., as

well as the very distinctive Quinqueloculina cf. Q. undulata d’Orbigny, are all

unique to the latest Eemian to earliest Weichselian of the west coast (McMillan

1987b). Also characteristic of these youngest marine Pleistocene deposits is the

widespread presence of mollusc shell retaining its life colours: purples, blues and

pinks seem the most commonly preserved. In earlier Pleistocene deposits, all the

mollusc shell is white.

Preliminary examination of the Varswater Formation (here regarded as later

Early Pleistocene) at a number of localities, as well as the phosphatic lithofacies

termed the ‘Saldanha Formation’ by Tankard (1975), has revealed the following

foraminifera species: Ammonia sp. (similar to Ammonia japonica), Elphidium sp.

(within the Elphidium crispum—macellum group), Glabratella australensis (Heron-

Allen & Earland) locally in abundance, Planulina cf. P. ariminensis d’Orbigny and

?Gavelinopsis sp. Most of these are unknown in the latest Eemian to earliest

Weichselian of both the Orange Shelf (McMillan 1987b) and Cape Town city

centre. The Pelletal Phosphorite Member of the Varswater Formation in phosphate

exploration borehole Q12 from near the New Varswater Quarry, Langebaanweg,

has yielded a somewhat different assemblage, with Cibicides lobatulus (Walker &

Jacob), Pararotalia nipponica (Asano), Elphidium advenum (Cushman), Pseudo-

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 131

nonion cf. P. chiliensis (Cushman & Kellett), Nonion boueanum (d’Orbigny),

Trifarina angulosa (Williamson), Oolinasp., Elphidium spp., Rosalina cf.

R. bradyi (Cushman), Planorbulina mediterranensis d’Orbigny, ?Gavelinella spp..,

Cibicides spp., Ammonia spp., ?Pseudononion sp., and Globigerina cf. G. bul-

loides d’Orbigny. More work is necessary on these older Pleistocene deposits but,

from the abundance of Glabratella australensis at many localities, a correlation of

the Varswater Formation with Unit IIA of the ‘Bredasdorp Formation’ seems

likely.

It must be emphasized that the majority of the foraminifera species employed

in correlation of the southern African latest Cainozoic are of local value only.

Ammonia japonica, Ammonia parkinsoniana, Elphidium crispum and Elphidium

macellum, for example, range back to the Early Pleistocene or Pliocene in the

Mediterranean or Japan. In the case of Ammonia and Elphidium, both were late

arrivals to southern Africa. Ammonia is first seen in the Early Miocene of

Mediterranean Europe, but in South Africa it first appears in the Early Pliocene.

Elphidium is known from the Early Eocene (Loeblich & Tappan 1964), but first

appears in South Africa in the Middle Miocene.

COMPARISON WITH HOLOCENE DEPOSITS

The bedded shelly sands exposed in the Cape Town municipal trenching lie

seaward of the Holocene shoreline as it was in 1693, before the construction of the

modern harbour works. The location, relative to the site of the Castle, is evident

on old maps. There thus exists the possibility that the sands are Holocene in age,

having accumulated prior to commencement of the land reclamation schemes

associated with harbour development. However, a Holocene age can be

discounted on a number of points.

The latest Pleistocene Unit III sands are rich in tests of Elphidium crispum

(Linné) s.1., and this species is absent everywhere in the Holocene, except rarely

where it has been reworked. Similarly, Quinqueloculina cf. Q. undulata

d’Orbigny and the other species listed on page 139 are not known from the

Holocene (see Fig. 4 for interpreted stratigraphic ranges). The absence of

Elphidium magellanicum Heron-Allen & Earland, widespread in shallow marine

environments off the west coast during the later Holocene (McMillan 1987b), also

supports a pre-Holocene age. Elphidium magellanicum occurs in the present-day

sands washed into the canalized mouth of the Soutrivier, just a little distance

north-east of the study area. Finally, nearly all the tests from the trench sands are

bleached and lack their life colours, whereas tests from Holocene samples include

a high proportion that retain their natural golden-brown coloration. This is

probably due to the destruction of the inner tectin lining of the test by bacteria

and percolating groundwater.

Since the latest Eemian—earliest Weichselian sands are overlain by a mix of

rubble and soil, the fate of any Holocene deposits at the site of the trenching, laid

down prior to land reclamation, is unclear.

132 ANNALS OF THE SOUTH AFRICAN MUSEUM

_TORANGE SHELF

Hil STRATIGRAPHY

S P Ey Gages

Elphidium magellanicum

Discammina compressa

Ammonia japonica

Ammonia parkinsoniana s.l.

Hyalinea balthica

Elphidium macellum s.l.

Elphidium articulatum

Elphidium advenum

Pararotalia nipponica

Cibicides lobatulus

Glabratella australensis

Elphidium

Quinqueloculina cf. undulata

crispum s.l.

Quinqueloculina isabellei

Oolina sp. A

Elphidium cf. advenum

Astrononion echolsi

Cassidulina crassa s.l.

Fig. 4. Range chart of selected time-restricted benthonic foraminifera species from the latest

Quaternary littoral of the west Cape and Namibia coasts. Those species marked with an asterisk

range back to near the Pliocene—Pleistocene boundary (base of Unit II) in the “Alexandria

Formation’; that marked by a cross ranges back in abundance to the later Early Pleistocene (base

of Unit ITA) of the ‘Bredasdorp Formation’.

REWORKING

A small number of the Cape Town tests are rather more poorly preserved

than the majority, and are regarded as having been reworked. One, ?Gavelinopsis

Sp., appears to be derived from the local equivalent of the Varswater Formation

(later Early Pleistocene) sandy limestones, as it is creamy in colour and cemented;

the same species occurs in the limestones at Saldanha. Occasional angular lumps

of creamy sandy limestone present in the Cape Town sands are probably also from

an equivalent of the Varswater Formation.

The remaining reworked tests may prove to be from the Early Miocene,

though this cannot be confirmed from the species present, which are

?Dentalina sp., Cibicides spp. and Uvigerina sp. (Fig. 9B). All show darker

infillings to the chambers (and dark test calcite) than is typical for later Neogene

or Quaternary foraminifera tests. An Early Miocene age seems most likely,

because of the extensive development of very fossiliferous limy clays of this age

that outcrop or subcrop below the Quaternary on the continental shelf off the

west coast. Intensive reworking of Early and Middle Miocene foraminifera tests

into latest Pleistocene regressive sands is known from the Orange Shelf (McMillan

1987b) and the Uloa Formation of Zululand (McMillan 1987a). The species listed

above are not typical of the Varswater Formation.

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 139

PROCESSING METHODS

All of the samples studied, including those described in the three appendices,

consist of clean or muddy, unconsolidated sands. These were easily washed with

a cold-water spray over a 240 mesh/63 micron sieve. Those samples with

substantial present-day plant debris were further cleaned by standing the washed

residue in a beaker of water and picking off the floating debris using a pair of

tweezers.

PHOTOGRAPHY AND DEPOSITION OF MATERIAL

Scanning electron microscope work was performed using the JEOL

JSM240 microscope of the University College of Wales, Aberystwyth. The

line drawings were executed with a camera lucida attachment to a light

microscope.

All illustrated specimens were returned to their relevant assemblage slides;

they can be distinguished by their gold coating. The eleven assemblage slides have

been deposited with the Ph.D. thesis material of McMillan (198765) in the

foraminifera collections of the Department of Geology, University College of

Wales, Aberystwyth. The slides possess the following McMillan collection

numbers:

CTC-1: Slide 118

C€TC—2: - Slide 119

CTC-3: Slide 120

CTC-4: Slide 121

BR-1: Slide 115

BR-2: Slide 116

SB-1: Slide 111

SB-LL1: Slide 112

SB-LL2: Slide 113

SB-LL3: Slide 114

H=1: Slide 117

A representative set of species slides has also been deposited in the Micro-

palaeontology Unit of the South African Museum.

FORAMINIFERA TAXONOMY

The taxonomic scheme followed is essentially a combination of that of

Loeblich & Tappan (1964, 1974), as revised by Haynes (1981). More detailed

discussions and full descriptions of many of the species listed below can be found

in McMillan (1987b).

134 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Textulariidae Ehrenberg, 1838

Subfamily Spiroplectammininae Cushman, 1927

Genus Spiroplectammina Cushman, 1927

?Spiroplectammina sp.

Fig. SA-B

Remarks

Two badly broken tests, which seem more likely to be referable to

Spiroplectammina than Textularia from their test shape. The specimens differ

from Spiroplectammina atrata (Cushman), prevalent in the Holocene deposits

of the middle and outer Orange Shelf (McMillan 1987b), in being less

compressed and with a more rounded test periphery. The latter feature,

however, is poorly preserved. The grain size of the test wall is also coarser than

either Spiroplectammina atrata from off the west coast or Spiroplectammina

wrightii (Silvestri), known from the Holocene of the Agulhas Bank (McMillan

1974).

Family Milliolidae Ehrenberg, 1839

Subfamily Quinqueloculininae Cushman, 1917

Genus Quinqueloculina d’Orbigny, 1826

Quinqueloculina contorta d’Orbigny, 1846

Fig. 6A—C

Quinqueloculina contorta d’Orbigny, 1846: 298, pl. 20 (figs 4-6). McMillan, 1974: 33, pl. 2

(fig. la—c); 1987b: 164, pl. 2 (figs 8-11).

Remarks

Some variation in the degree of indentation of the rectangular test margins is

evident in southern African individuals. Those from the Agulhas Bank (McMillan

1974) display strong indentations, but examples from the west coast are flatter

(McMillan 1987b, present study) and more in keeping with the type illustrations

of the species (D’Orbigny 1846). Some of the Orange Shelf tests display distinctly

inflated margins, and the rectangular feature is almost lost (McMillan 19875, pl. 2

(figs 8-9)). The toothplate of the Cape Town tests is perhaps a little more bladed

and more elongate than that originally figured by D’Orbigny.

The species occurs in small numbers in the more nearly littoral intervals of

the latest Eemian—earliest Weichselian of the Orange Shelf (McMillan 19875),

and over the middle and outer shelf on the Agulhas Bank in the Holocene

(McMillan 1974). This discrepancy in environments between the latest Pleis-

tocene and the Holocene is mirrored by several other species found in the Cape

Town shelly sands.

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 135

Fig. 5. A-B. ?Spiroplectammina sp. A. Side view, CTC 4. x 130. B. Apertural view, same

specimen. X 141. C-—E. Quinqueloculina isabellei dOrbigny. C. Side view, CTC 4. xX 67.

D. Apertural view, same specimen. X 520. E. Edge view, same specimen. xX 60. F. Quin-

queloculina cf. Q. lata Terquem. Side view, CTC 4. x 117. G—H. Quinqueloculina triangu-

laris d’Orbigny. G. Side view, CTC 4. xX 97. H. Apertural view, same specimen. xX 113.

I. Quinqueloculina cf. Q. undulata d’Orbigny. Apertural view, CTC 2. xX 108.

136 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 6. A-C. Quinqueloculina contorta d’Orbigny. All same specimen, CTC 3. All x 67.

A. Side. B. Opposing side. C. Apertural view. D-F. Quinqueloculina dunkerquiana (Heron-

Allen & Earland). All same specimen, CTC 4. All x 67. D. Side. E. Opposing side.

F. Apertural view. G—I. Quinqueloculina seminulum (Linné). All same specimen, CTC 3. All

x 67. G. Side. H. Opposing side. I. Apertural view.

Quinqueloculina dunkerquiana (Heron-Allen & Earland, 1930)

Fig. 6D-F

Miliolina dunkerquiana Heron-Allen & Earland, 1930: 56, pl. 12 (figs 9-11).

Quinqueloculina dunkerquiana (Heron-Allen & Earland) Haynes, 19736: 2, pl. 1 (fig. 8).

McMillan, 1987b: 166, pl. 2 (figs 12-13).

Quinqueloculina cf. Q. dunkerquiana (Heron-Allen & Earland) McMillan, 1974: 33, pl. 2

(fig. la—c).

Remarks

A number of small Quinqueloculina specimens with test width, height and

outline, and chamber shape much in keeping with the north-west European

shallow marine species Quinqueloculina dunkerquiana. As with the shell figured

by Heron-Allen & Earland (1930) from present-day sediments of the Plymouth

district, south-western England, the maximum height of tests is little greater than

their maximum width. The distinctive indentations on the two last-formed

chambers give a slightly sigmoiline appearance to the test if seen in apertural

view, but the internal arrangement is always clearly quinqueloculine.

The species has been regarded as close to Quinqueloculina seminulum

(Linné) by British authors (Heron-Allen & Earland 1930; Haynes 1973a, 19735),

but the gradation of forms recognized by Haynes (19735, pl. 1) between the two

species is far less evident in southern African late Quaternary assemblages.

Specimens referable to Quinqueloculina dunkerquiana occur widely at the present

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 137

day in shallow marine, innermost shelf deposits around the entire South African

coastline. The full stratigraphic range of the species remains unclear but on the

Orange Shelf it extends throughout the latest Eemian to earliest Weichselian

interval, while it is rare in the Holocene (McMillan 1987b).

Quinqueloculina isabellei d Orbigny, 1839

Fig. 5C-E

Quinqueloculina isabellei d Orbigny, 1839b: 74, pl. 4 (figs 17-19).

Quinqueloculina seminulum (non Linné) Boltovskoy et al., 1980: 47, pl. 29 (figs 7-10,

?figs 11-13).

‘Quinqueloculina plataensis’ McMillan, 1987b: 169, pl. 3 (figs 1-4).

Remarks

The nearly circular aperture and strongly bifid apertural tooth, with the

pronounced inflation of the final three chambers leading to a broad, evenly

rounded test periphery, are distinctive features. These clearly distinguish the

species from Quinqueloculina seminulum (Linné), with which it has been

confused. Quinqueloculina araucana, also described by D’Orbigny (18396) from

present-day sediments off the west coast of South America, is clearly very similar

to Qinqueloculina isabellei, and the two may be conspecific. The test outline of the

type specimen of Quinqueloculina isabellei is strongly lobate, though this feature

is not evident in the well-illustrated specimens of Boltovskoy et al. (1980), nor in

the southern African tests here referred to D’Orbigny’s species.

Boltovskoy et al. (1980: 47) considered their specimens to be part of a range

of forms that they grouped under the name Quinqueloculina seminulum. If the

neotype of Quinqueloculina seminulum from Rimini, Italy, illustrated by Loeblich

& Tappan (1964, fig. 349 (no. la—c)), is regarded as typical for Linné’s species,

then the South Atlantic shells here referred to Quinqueloculina isabellei are

clearly distinct. There is no gradation between Quinqueloculina isabellei and

Quinqueloculina seminulum in either the Orange Shelf latest Eemian to earliest

Weichselian shells (McMillan 1987b), or in those from Cape Town. See also

remarks on Quinqueloculina seminulum (p. 138).

Quinqueloculina isabellei has yet to be found in the Holocene of southern

Africa, whereas most, if not all, southern South American records are from the

Holocene.

Quinqueloculina cf. Q. lata Terquem, 1876

Fig. 5F

see Quinqueloculina lata Terquem, 1876: 82, pl. 2 (fig. 8a—c). Mathieu er al., 1971: 159, pl. 1

(figs 5-7).

Remarks

The few, rather damaged specimens from the Cape Town samples lack the

embracing, quadrate nature of the final chambers of European littoral specimens

138 ANNALS OF THE SOUTH AFRICAN MUSEUM

referred to Quinqueloculina lata. However, they are markedly more elongate

than is typical for Quinqueloculina seminulum (Linné). In this respect they appear

similar to the tests illustrated by Haynes (1973a, pl. 7 (figs 10, 12), 1973b, pl. 1

(fig. 1)) from Cardigan Bay, Wales, although the Cape Town tests lack such an

elongate aperture.

Similar forms occur in the latest Pleistocene of the Orange Shelf (McMillan

1987b) to those in the Cape Town samples but, as with the majority of Pleistocene

littoral miliolids, almost all are badly damaged and usually only the kernel of each

test has preserved. An exact comparison of the Orange Shelf and Cape Town

shells is thus not easy to achieve.

Quinqueloculina seminulum (Linné, 1758)

Fig. 6G-I

Serpula seminulum Linné, 1758: 786.

Quinqueloculina seminulum (Linné) Loeblich & Tappan, 1964: C458, fig. 349 (no. la—c).

Haynes, 1973a: 74, pl. 7 (figs 14, 19), pl. 8 (fig. 3), pl. 32 (figs 1-3), text-fig. 18 (nos 1-4).

McMillan, 1974: 35, pl. 2 (fig. Sa—c); 1987b: 172, pl. 3 (figs 5-6). Martin, 1981: 26, pl. 2

(i, 1).

Remarks

The neotype illustrated by Loeblich & Tappan (1964), from the shore sands

of Rimini, Italian Adriatic, has been relied on in establishing the identity of the

various smooth-walled Quinqueloculina tests in the Cape Town samples. The

neotype displays a rounded-triangular aperture with a rather short tooth that

exhibits a distinctly bifid tip (apparently slightly broken on one side). In contrast,

southern African tests reveal a more elongate-ovate aperture, within which lies a

long, narrow tooth that extends for about three-quarters of the aperture length.

The tooth is incipiently bifid at its free end, but this feature is never as well

developed as the pronounced Y-shaped tooth of Quinqueloculina isabellei

d’Orbigny. In keeping with the neotype, southern African tests of Quinquelocu-

lina seminulum display a general test outline and chamber morphology sharply

different to those seen in Quinqueloculina isabellei, notably in the sub-rounded,

rather than broadly rounded test margin. However, the Cape Town tests are not

as strongly triangular in cross-section as the Rimini neotype.

Quinqueloculina triangularis d’Orbigny, 1846

Fig. 5G—H

Quinqueloculina triangularis d’Orbigny, 1846: 288, pl. 18 (figs 7-9). Mathieu ef al., 1971: 159,

pl. 1 (figs 8-10).

Remarks

In terms of its test morphology, this species compares closely with

Quinqueloculina dunkerquiana (Heron-Allen & Earland), differing mainly in the

possession of a distinctly bifid tooth, and lacking the indentations of the final two

chambers. The Cape Town tests are a little more squat than those of Mathieu

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 139

et al. (1971), and more nearly as high as wide. The bifid tooth is particularly well

developed in the figured specimen.

D’Orbigny (1946) described Quinqueloculina triangularis from the Miocene

of the Vienna Basin; Mathieu ef al. (1971) figured tests from the present-day

beach sands of the Dunkerque coast, north France.

Quinqueloculina cf. Q. undulata d’Orbigny, 1852

Figs 51, 7A

see Quinqueloculina undulata d’Orbigny, 1852: 195. Rosset-Moulinier, 1972: 140, pl. 6

(figs 5-7).

Quinqueloculina cf. Q. undulata d’Orbigny: McMillan, 1987b: 175, pl. 3 (figs 7-9).

Remarks

This distinctive species, with its finely grooved surface and very elongate

aperture revealing a slender, tapering tooth, is at present known only from the

latest Pleistocene (Unit HI) off the west coast of southern Africa. D’Orbigny’s

(1852) species was partly distinguished by its undulating test margins, a feature

that was also depicted, though not to so marked a degree, by Cushman (1945).

Cushman’s figured specimen is much less intensely ornamented than the original

test of D’Orbigny (1852). The shells illustrated by Rosset-Moulinier (1972) are

not undulate; she noted that the species was close to Quinqueloculina bicornis

(Walker & Jacob), but more elongate in form.

None of the southern African tests, from Cape Town or the Orange Shelf,

possess an undulate periphery, and they thus appear closest to those described by

Rosset-Moulinier (1972) from the Brittany coast. D’Orbigny (1852) described the

species from the Pliocene of Castel-Arquato and from present-day deposits near

Rimini, both in Italy.

The illustrated test from Cape Town possesses a damaged apertural tooth;

Orange Shelf tests display a bladed tooth, which extends for about three-quarters

the length of the aperture (McMillan 1987b). The depressed area at either end of

the aperture is also typical of the Orange Shelf examples.

Quinqueloculina spp.

Remarks

Many badly broken tests of smooth-walled, unornamented Quinqueloculina that

probably mostly fall within the Quinqueloculina seminulum—dunkerquiana group.

Genus Siphonaperta Vella, 1957

Siphonaperta sp.

Fig. 7B—C

Quinqueloculina agglutinans (non d’Orbigny) Martin, 1974: 85; 1981: 25, pl. 2 (fig. 8).

Sigmoilopsis sp. McMillan, 1974: 37, pl. 2 (fig. 7a—c).

Sigmoilopsis schlumbergeri (non Silvestri) Martin, 1981: 27, pl. 2 (fig. 13).

Quinqueloculina horrida (non Cushman) Boltovskoy et al., 1980: 46, pl. 27 (figs 17-20).

140 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 7. A. Quinqueloculina cf. Q. undulata d’Orbigny. Oblique edge view, same specimen as

Fig. 51. x 65. B-C. Siphonaperta sp. B. Side view, CTC 4. X 96. C. Apertural view, same

specimen. x 132. D. Miliolinella subrotunda (Montagu). Side view, CTC 4. xX 110.

E-F. Lagena semilineata Wright var. E. Apertural view, CTC 3. x 122. F. Side view, same

specimen. X 112. G. Lenticulina sp. Side view, CTC 4. x 129. H. Oolinasp. A. Side view,

CTC 4. x 168.

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 141

Remarks

The above, rather disparate references appear to be to the same species. The

presence of an apertural neck and the rough, agglutinated test surface, as well as

a quinqueloculine chamber arrangement, are indicative of the genus Siphonap-

erta, rather than Quinqueloculina or Sigmoilopsis. Sectioning of tests from the

Orange Shelf Holocene (see McMillan 19875, pl. 3 (fig. 11), text-fig. 2) shows the

chamber arrangement to be quinqueloculine rather than sigmoiline.

These tests can be distinguished from Quinqueloculina horrida, described by

Cushman (1947) from off Charleston, South Carolina, in being substantially more

triangular in cross-section and somewhat more squat, and lacking the elongate

neck and slender bifid apertural tooth of the United States species. Quinqueloc-

ulina agglutinans, in contrast, possesses subangular chamber margins, and a large,

circular aperture with a phialine lip, but not developed on a neck (see Le Calvez

& Le Calvez 1958: 166, pl. 9 (figs 103—104)).

Analysis of wall composition of one test dissolved in dilute hydrochloric acid

(McMillan 1987b) revealed the presence of small lengths of broken sponge

spicules in the wall, a feature that is not evident in external view. The species

ranges throughout the latest Eemian to earliest Weichselian and Holocene

sequences studied by McMillan (19875), though it is substantially more abundant

in the Holocene. The species does not occur in the older Pleistocene rocks of

South Africa. Most of the Cape Town specimens are damaged, but the distinctive

arrangement of coarse- and fine-grained quartz particles over the test wall easily

distinguishes even badly fragmented tests.

Subfamily Miliolinellinae Vella, 1957

Genus Miliolinella Wiesner, 1931

Miliolinella subrotunda (Montagu, 1803)

Fig. 7D

Serpula subrotunda dorso elevato Walker & Boys, 1784: 2, pl. 1 (fig. 4).

Vermiculum subrotundum Montagu, 1803: 521.

Miliolinella subrotunda (Montagu) Haynes, 1973a: 56, pl. 5 (figs 5-6, 12-13), pl. 31 (figs 8-9),

text-fig. 11 (nos 1-4), text-fig. 12 (nos 1-11). Ponder, 1974: 201, pl. 1 (figs 1-2), pl. 2

(figs 6-11).

Remarks

British researchers have considered the wide variety of forms encountered to

be referable to one species: ‘It would seem altogether inappropriate to separate

some as Miliolinella, some as Scutuloris and to dispatch the irregular,

wild-growing, hauerinid forms into a separate genus and even subfamily’ (Haynes

1973a: 57). This concept is followed here. South African specimens also show

much variation, from tight triloculine tests to ones that are loosely and irregularly

triloculine, but the wild forms are by and large missing from the cool waters of the

west coast.

142 ANNALS OF THE SOUTH AFRICAN MUSEUM

Miliolinella subrotunda occurs widely in the South African littoral at the

present day, from cold-water west coast to warm-water east coast. The species

ranges back to the latest Eemian to earliest Weichselian on the Orange Shelf

(McMillan 1987b), but its earlier record in southern Africa is as yet unclear.

Family Nodosariidae Ehrenberg, 1838

Genus Lagena Walker & Jacob, 1798

Lagena semilineata Wright, 1886 var.

Fig. 7E—F

see Lagena semilineata Wright, 1886: 320, pl. 26 (fig. 7).

Lagena semilineata (non Wright) Earland, 1934: 161, pl. 7 (figs 19-20).

Lagena semilineata Wright var. McMillan, 1987b: 204, pl. 5 (figs 9-11).

Remarks

The single specimen is not particularly well formed, and the ornamentation

is not quite typical of the tests described by McMillan (1987b) from the Orange

Shelf. The group of short apical spines of this specimen are poorly developed, lost

perhaps through abrasion of the test. The flat-topped, tapering ribs that occupy

much of the lower half of the globular part of the test are rather more irregularly

formed, particularly in their varying width and length, than is usual. There also

are occasional tubercular or rugose developments in the depressed areas between

the ribs. However, the style of the ornamentation is so distinctive that there can

be no doubt that the Cape Town specimen and those from the Orange Shelf are

the same variety.

Genus Lenticulina Lamarck, 1804

Lenticulina sp.

Fig. 7G

Remarks

One broken specimen that appears to be in situ, may be referable to one of

the rather conservative, unornamented species typified by Lenticulina gibba

(d’Orbigny). Little can be made of the single specimen.

Family Glandulinidae Reuss, 1860

Subfamily Oolininae Loeblich & Tappan, 1961

Genus Oolina d’Orbigny, 1839

Oolina sp. A McMillan, 1987

Figs 7H, 8A-B

Oolina sp. A McMillan, 1987b: 220, pl. 6 (figs 13-14).

Remarks

This graceful, pyriform Oolina occurs rarely in the sandier, littoral deposits

of the latest Eemian to earliest Weichselian of the Orange Shelf (McMillan

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 143

Fig. 8. A-B. Oolina sp. A. A. Apertural view, CTC 4. xX 156. B. Apertural view, same

specimen as Figure 7H. X 173. C-—D. Oolina squamososulcata (Heron-Allen & Earland).

C. Side view, CTC 4. xX 206. D. Apertural view, same specimen. X 246. E. Fissurina lucida

(Williamson). Side view, CTC 3. xX 327. F-G. Fissurina cf. F. marginata (Walker & Boys).

F. Apertural view, CTC 3. xX 204 G. Side view, same specimen. xX 161.

144 ANNALS OF THE SOUTH AFRICAN MUSEUM

1987b). The species is distinguished by a vertically elongated hexagonal ribbing

pattern ornamenting the test surface. Some variation is evident in the height of

the elongated hexagons at the maximum width of the test; those from Cape Town

exhibit more nearly regular hexagons than those from further north.

The narrower part of the test adjacent to the aperture is marked with small

depressions and protrusions that give the surface a rather scaly, reptilian

appearance. The subcircular aperture is developed on a short neck, which in

well-preserved examples is almost phialine and ornamented with many short,

vertically aligned grooves.

Oolina squamososulcata (Heron-Allen & Earland, 1922)

Fig. 8C—D

Lagena squamoso-sulcata Heron-Allen & Earland, 1922: 151, pl. 5 (figs 15, 19).

Oolina squamoso-sulcata (Heron-Allen & Earland) Loeblich & Tappan, 1953: 74, pl. 12

(figs 6-7).

Remarks

The single test compares closely with Oolina squamososulcata except that the

horizontally aligned ribs ornamenting the test are more arched, and they fade

away only slowly toward the apical end of the test, rather than disappearing

abruptly about midway down. The holotype figured by Heron-Allen & Earland

(1922) displays 13 vertical ribs; Loeblich & Tappan (1953) reported that their

tests exhibit about 12 to 20; the present example possesses 15.

Genus Fissurina Reuss, 1850

Fissurina lucida (Williamson, 1848)

Fig. 8E

Entosolenia marginata (Montagu) var. lucida Williamson, 1848: 17, pl. 2 (fig. 17).

Fissurina lucida (Williamson) Murray, 1971: 97, pl. 39 (figs 1-3). Boltovskoy ef al., 1980: 32,

pl. 15 (figs 17-20).

Remarks

One specimen, rather damaged, but closely comparable to Orange Shelf

examples of this species. As with Fissurina marginata (Walker & Boys), it would

seem to be confined to cool, shallow marine environments, off the west coast of

southern Africa only, in both the latest Pleistocene and the Holocene (McMillan

1987b).

Fissurina cf. F. marginata (Walker & Boys, 1784)

Fig. 8F-G

see Serpula (Lagena) marginata Walker & Boys, 1784: 2, table 1, fig. 7.

see Fissurina marginata (Walker & Boys) Feyling-Hanssen, 1964: 315, pl. 15 (fig. 22). Haynes,

1973a: 97, fig. 20 (nos 7-8). McMillan, 1987b: 225, pl. 7 (figs 2-3).

Remarks

This single example displays several slight differences from the tests seen in

the latest Pleistocene and Holocene of the Orange Shelf (McMillan 1987b). The

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 145

peripheral ornamentation is more nearly carinate rather than a rounded

thickening; the lips on each side of the slit aperture are thicker but less elongate;

and there are two raised areas on each side of the test, a little below the aperture.

The raised features are low elongate protrusions, aligned roughly parallel to the

test periphery. The degree of inflation and the general outline of the Cape Town

test is about the same as seen in the Orange Shelf specimens.

A rounded peripheral ornamentation would seem to be closer to Fissurina

marginata in British waters (Haynes 1973a: 98), so that this Cape Town example

rather falls outside the confines of the north-west European interpretation of the

species.

Family Buliminidae Jones, 1875

Genus Bulimina d’Orbigny, 1826

Bulimina marginata d’Orbigny, 1826

Fig. 9A

Bulimina marginata dOrbigny, 1826: 269, pl. 12 (figs 10-12). Murray, 1971: 119, pl. 49

(figs 1-7). Knudsen, 1973: 181, pl. 2 (figs 10-11). Martin, 1981: 41, pl. 4 (figs 8-9).

Remarks

One specimen, showing rather poorly incised steps into the bases of the later

chambers, but with a distinct fringe of blunt spines near the lower margins of some

chambers. The test is abraded and has been subject to boring, but is clearly

comparable to the more distinctive tests detailed by Martin (1981) and McMillan

(19876) from further north. Bulimina marginata ranges throughout the latest

Eemian-earliest Weichselian and the Holocene sequences of the Orange Shelf

(McMillan 19875).

Family Uvigerinidae Haeckel, 1894

Genus Trifarina Cushman, 1923

Trifarina angulosa (Williamson, 1858)

Fig. 1OA-B

Uvigerina angulosa Williamson, 1858: 67, pl. 5 (fig. 140).

Trifarina angulosa (Williamson) Knudsen, 1971: 241, pl. 18 (figs 8-9). Haynes, 1973a: 126,

pl. 10 (figs 12-13, 16-17), pl. 11 (fig. 11).

Remarks

One example, with the terminal part rather broken. However, it clearly

shows the triserial chamber arrangement, the tricarinate test periphery, the nearly

triangular cross-section to the test, and irregular, vertically aligned ribs on the test

surface. The species occurs throughout the latest Eemian to earliest Weichselian

and the Holocene of the Orange Shelf (McMillan 1987b), where it shows a

considerable range in its surface ornamentation of vertically aligned ribs. Tests

range from being almost smooth, with only the three peripheral keels, to densely

146 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 9. A. Bulimina marginata d’Orbigny. Side view, CTC 4. x 123. B. Uvigerina sp.

(reworked). Side view, CTC 3. x 133. C. Brizalina spathulata (Williamson). Side view,

CTC 3. xX 153. D-E. Cassidulina laevigata d’Orbigny. D. Side view, CTC 4. xX 136.

E. Apertural view, same specimen. X 129. F-H. Cassidulina crassa d’Orbigny s.l. F. Side

view, CTC 3. X 187. G. Apertural view, same specimen. X 177. H. Close up of foramen,

showing serrate margins, same specimen. X 646.

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 147

Fig. 10. A-B. Trifarina angulosa (Williamson), same specimen, CTC 4. Both x 67. A. Side

view. B. Apertural view. C—D. Bolivina sp., same specimen, CTC 3. Both x 135. C. Side

view. D. Apertural view.

ribbed over almost the entire exterior. Variation also occurs in the intensity of the

ribs, some shells displaying few strongly developed ones, whereas others reveal

many closely spaced fine ribs.

Family Bolivinitidae Cushman, 1927

Genus Bolivina d’Orbigny, 1839

Bolivina sp.

Fig. 10C—D

Remarks

One small test, ornamented with fine, dense pitting over the surface. The

specimen shows some resemblance to Bolivina variabilis (Williamson), but is too

juvenile a shell to make a reliable identification.

Genus Brizalina Costa, 1856

Brizalina spathulata (Williamson, 1858)

Fig. 9C

Textularia variabilis Williamson var. spathulata Williamson, 1858: 76, pl. 6 (figs 164-165).

Brizalina spathulata (Williamson) Hedley et al., 1965: 21, pl. 6 (fig. 23a—b), text-fig. 6A—-G.

Haynes, 1973a: 135, fig. 25 (nos 10-13).

Remarks

One example, rather damaged. A narrow test when compared to examples

from off the Orange River, but the arrangement and coarseness of the test

perforations, the sub-rounded acute margin to the test, and the angle at which the

sutures decline to the margin are closely comparable. The central zig-zag suture

of the Cape Town test is particularly limbate and prominent.

Brizalina spathulata ranges throughout the latest Eemian and earliest

Weichselian, and the Holocene of the Orange Shelf (McMillan 1987b). However,

148 ANNALS OF THE SOUTH AFRICAN MUSEUM

in the latest Pleistocene, the species is widespread through the sandier, more

nearly littoral parts of the sequence; these tests lack any substantial carination of

the periphery (see McMillan 19875, pl. 11 (figs 10—-11)). The shell from the Cape

Town samples is of this type. In contrast, in the Holocene, the species appears

commonest in more offshore, shelf environments; these tests usually possess a

rather bladed carina (see Martin 1981, pl. 3 (figs 12-13); McMillan 19875, pl. 11

(fig. 12)). There is thus the possibility that two varieties are represented off the

west coast in the Late Quaternary.

Family Cassidulinidae d’Orbigny, 1839

Genus Cassidulina d’Orbigny, 1826

Cassidulina laevigata d’Orbigny, 1826

Fig. 9D-E

Cassidulina laevigata d’Orbigny, 1826: 282, pl. 15 (figs 4-5). Knudsen, 1971: 246, pl. 7

(figs 20-21), pl. 18 (fig. 12). Boltovskoy et al., 1980: 22, pl. 7 (figs 4-6).

Remarks

Few specimens, all distinguished from the similar species Cassidulina carinata

Silvestri by a rounded, non-carinate margin, coarser test perforations and thicker

walled test. The precise relationship between the two species is not clear. On the

Orange Shelf, Cassidulina laevigata occurs in abundance in the littoral and

sublittoral deposits of the latest Eemian—earliest Weichselian, but is present only

rarely in the nearshore Holocene. Cassidulina carinata, in contrast, is rare in the

latest Pleistocene, but common in the offshore (middle to outer shelf) Holocene.

It remains possible that the two species are two facies-controlled formae of one

species.

Cassidulina crassa d’Orbigny, 1839 s.1.

Fig. 9F—H

see Cassidulina crassa d’Orbigny, 1839b: 56, pl. 7 (figs 18-20).

Cassidulina crassa d’Orbigny ‘intermediate form’ Heron-Allen & Earland, 1932: 358.

Cassidulina crassa d’Orbigny forma media Lena, 1966: 316, pl. 2 (fig. 7).

Remarks

One test, which like those present in the latest Pleistocene of the Orange

Shelf, is not Cassidulina crassa sensu stricto but a smaller form, first recognized by

Heron-Allen & Earland (1932) from the seas around the Falkland Islands. This

form, termed ‘intermediate’, was believed by Heron-Allen & Earland to be a

juvenile or pauperate stage of Cassidulina crassa, as first described by D’Orbigny

(1839b).

Tests possess a rather short aperture, extending from the interio-marginal

suture, up toward, but not reaching, the test periphery. There is no extension of

the aperture along the interio-marginal suture, as is seen in Cassidulina crassa

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 149

‘small type’ of Heron-Allen & Earland (1932, pl.9 (figs 29-31)). Some

subsequent listings of the ‘intermediate form’ have been made by Boltovskoy

(1959, 1961) and Lena (1966) from the coast of Argentina.

Cassidulina crassa s.s. would seem to be absent in southern African waters,

both in the Pleistocene and the Holocene. The ‘intermediate form’ appears to be

confined to deposits of the latest Eemian—earliest Weichselian, and to the cold

waters off the west coast.

Family Discorbidae Ehrenberg, 1838

Genus Planulinoides Parr, 1941

Planulinoides biconcavus (Jones & Parker, 1862)

Fig. 11A—C

Discorbina bi-concava Jones & Parker, in Carpenter, 1862: 201, fig. 32G.

Discorbina biconcava Parker & Jones (sic) Parker & Jones, 1865: 385, 422, pl. 19 (fig. 10a—c).

Brady, 1884: 653, pl. 91 (fig. 2a—c).

Planulinoides biconcavus (Jones & Parker) Parr, 1941: 305, fig. a-c. Loeblich & Tappan, 1964:

C584, fig. 458 (nos 4a—c, 5-6). Lowry, 1987: 269, pl. 17 (fig. 1a, c, ?1b).

Planulinoides biconcava (Jones & Parker) (sic) McMillan, 1974: 58, pl. 5 (fig. 5a—c).

Discorbinella biconcava (Jones & Parker) Carter, 1978: 19, pl. 1 (figs 2-3).

Remarks

This very distinctive species, widespread in shallow marine environments

around south-east Australia at the present day (Carpenter 1862; Parker & Jones

1865; Brady 1884; Chapman 1909; Parr 1941, 1945; Collins 1974), also occurs

sporadically around southernmost Africa. Specimens from the middle shelf of the

Agulhas Bank (McMillan 1974) are always abraded and scoured, and clean tests

would seem to be lacking. Whether this indicates that Planulinoides biconcavus is

restricted to latest Pleistocene (latest Eemian—earliest Weichselian), Unit III

deposits on the Agulhas Bank, rather than the Holocene as well, and has been

reworked into the Holocene, is not fully clear at present.

Fig. 11. Planulinoides biconcavus (Jones & Parker). All same specimen, CTC 3. All x 68.

A. Dorsal view. B. Apertural view. C. Ventral view.

150 ANNALS OF THE SOUTH AFRICAN MUSEUM

Planulinoides biconcavus occurs in small numbers on the Agulhas Bank

(McMillan 1974; Lowry 1987), and Lowry recorded it between 107 m and 890 m

water depth. The species appears to be absent off the west coast in the Holocene

and Pleistocene, except for the single specimen from the present study. However,

in the Early Miocene of the Orange Shelf, a similar, perhaps identical species

occurs (McMillan 1975). A full comparison of the Miocene and Late Quaternary

specimens has not yet been attempted. In Australia, the species is known from the

Early Miocene (Heron-Allen & Earland 1924), the Middle Miocene (Mitchellian)

(Carter 1978), and the Early and Middle Pliocene (Cheltenhamian and Kalimnan)

(Parr 1939, 1941) of Victoria.

The Cape Town test is typical in its shell morphology, but as is often the case

with the Agulhas Bank specimens examined by McMillan (1974), the thinner-

walled, less-ornamented ventral side, with the small rounded flaps over the spiral

suture, is very badly broken. The elongate-ovate, oblique primary aperture is

distinct, though rather damaged. Despite showing some morphological similari-

ties to Planulina ariminensis d’Orbigny, even very badly broken tests of

Planulinoides biconcavus are easily distinguished, since they entirely lack the

coarse test perforations of D’Orbigny’s species.

Some differences in size of the flaps on the ventral side, over the spiral

suture, can be seen. Tests such as that illustrated by Carter (1978) display large

triangular flaps, whereas southern African Quaternary examples reveal only

small, rounded flaps.

Genus Rosalina d’Orbigny, 1826

Rosalina bradyi (Cushman, 1915)

Fig. 12A-—C

Discorbina globularis (non d’Orbigny) Brady, 1884: 643, pl. 86 (fig. 8a—c).

Discorbis globularis (d’Orbigny) var. bradyi Cushman, 1915: 12, pl. 8 (fig. 1a—c).

Rosalina bradyi (Cushman) Hedley et al., 1967: 42, pl. 1 (fig. 3), pl. 11 (fig. 2a—c), text-figs 50-55.

Remarks

This essentially Indo-Pacific species appears to reside preferentially in

patches of Corallina alga in the littoral (Hedley etal. 1967). It occurs in

association with Corallina debris in vibracore 620/40 on the nearshore Orange

Shelf, in sandy sediments of earliest Weichselian age (McMillan 1987b).

The illustrated test from Cape Town is closely comparable to those from the

Orange Shelf, although, being a rather juvenile shell, the limbation of the

dorsal sutures is not well developed. The ‘milled edge’ of Rosalina bradyi

(Cushman, 1915) is caused by the peripheral coarse perforations when viewed

from the ventral side, which are visible at right angles through the glassy shell.

This feature, obviously, is obscured in the scanning electron microscope

photographs.

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 151

Fig. 12. A-C. Rosalina bradyi (Cushman). A. Ventral view, CTC 4. xX 139. B. Side view,

same specimen. X 144. C. Dorsal view, same specimen. X 135. D-F. Rosalina sp. D. Dorsal

view, CTC 3. x 125. E. Side view, same specimen. xX 115. F. Ventral view, same specimen.

x 120. G-I. Glabratella australensis (Heron-Allen & Earland). G. Dorsal view, CTC 4.

x 81. H. Side view, CTC 4. xX 130. I. Ventral view, CTC 4. x 107.

152 ANNALS OF THE SOUTH AFRICAN MUSEUM

Rosalina sp.

Fig. 12D-—F

Remarks

One specimen, closely comparable in its test morphology to Rosalina

williamsoni (Chapman & Parr), but differing in a pronounced umbilical

ornament. A similar form, less ornate, occurs on the Orange Shelf (McMillan

1987b—described as Rosalina williamsoni). The Orange Shelf shells display a

variable, tuberculate ornamentation in and around the umbilicus, but the Cape

Town test exhibits a stronger ornament of radiating, elongate-ovate flat-topped

pillars and bosses.

North-west European examples of Rosalina williamsoni (see Rosset-

Moulinier 1972, pl. 9 (fig. 32); Haynes 1973a, pl. 17 (fig. 13), text-fig. 31 (no. 3))

feature a relatively unornamented umbilicus, with just occasional bosses or

tubercles on the umbilical terminations of the chambers in the final whorl, and

within the umbilicus itself. In contrast, the tests figured by Boltovskoy et al.

(1980, pl. 12 (figs 5—12)) from the coast of Argentina as Discorbis williamsoni

appear in fact more closely allied to Gavelinopsis praegeri (Heron-Allen &

Earland), since the umbilicus is infilled with a distinct plug. It seems probable that

the Discorbis cf. D. valvulatus (d’Orbigny) of Boltovskoy et al. (1980, pl. 12

(figs 1—4)) is conspecific with Rosalina williamsoni sensu McMillan (1987b). Some

similarity, in terms of style of ornamentation, exists between the Orange Shelf

tests and Rosalina malovensis (Heron-Allen & Earland) from the seas around the

Falkland Islands. However, none of the southern African shells feature such

elongate, crescentic chambers, evident on both dorsal and ventral sides of the

test, as exist in Rosalina malovensis.

The radiating ornament around the umbilicus of both the Cape Town and

Orange Shelf forms perhaps shows some similarity to Rosalina parri Collins (1974,

pl. 3 (fig. 36a—c)). However, this Australian species is only slightly bedecked with

pillars around the umbilicus; more evident are the radiating grooves and ridges that

cover nearly all the ventral side of the illustrated test. Radiating ornament around

the umbilicus has been one of the characteristics of the family Gla-

bratellidae of Loeblich & Tappan (1964: C587), but the last chamber of many of the

southern African tests of this group clearly extends a small triangular flap into the

umbilical area, a feature that is rather more typical of Rosalina than Glabratella.

Family Bagginidae Cushman, 1927

Genus Glabratella Dorreen, 1948

Glabratella australensis (Heron-Allen & Earland, 1932)

Fig. 12G-I

Discorbina pileolus (non d’Orbigny) Brady, 1884: 469, pl. 89 (figs 2—4).

Discorbis australensis Heron-Allen & Earland, 1932: 416. Parr, 1939: 68.

Glabratella australensis (Heron-Allen & Earland) McMillan, 1987b: 363, pl. 14 (figs 14-18),

pl. 15 (figs 1-2).

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 153

Remarks

A common littoral species along most of the South African coastline at the

present day (Brady 1884), extending at least from Cape Town to Mzamba,

Transkei (unpubl. studies). It has also been widely recorded along the south,

south-east and south-west coasts of Australia (Brady 1884; Chapman 1909; Parr

1945; McKenzie 1962; Albani 1968a, 1968b; Johnson & Albani 1973; Collins

1974; Albani & Johnson 1975) and off New Zealand (Mestayer 1916; Cushman

1919). Its fossil range in Australia appears to be Middle Pliocene (Kalimnan)

(Parr 1939), Pleistocene (Collins 1953), and present day. In contrast, in South

Africa, it is confined to the Pleistocene, being extremely rare or absent in the

earliest Pleistocene (Unit II) of the ‘Alexandria Formation’, and only coming into

prominence in the later Early Pleistocene (Unit IIA) of the ‘Bredasdorp

Formation’. Glabratella australensis occurs in considerable numbers at some

localities of the Varswater Formation (probably Unit IIA), particularly at

Hoedjiespunt, Saldanha.

The species is distinguished from the South American Glabratella pileolus

(d’Orbigny) in its larger number of chambers per whorl (‘eight or more’ rather

than ‘four or five—Heron-Allen & Earland 1932: 416), its shallower umbilicus,

and its somewhat compressed, low-spired test. Living specimens of Glabratella

australensis from intertidal pools at Camps Bay and elsewhere on the north-west

Cape Peninsula are a strong greenish brown in colour; perfect, unabraded tests

are spectacular when viewed under the scanning electron microscope (see

McMillan 1987b, pl. 14 (figs 14-18)). Tests from the Cape Town city centre

samples are rather abraded, and much of the detail of the ventral ornamentation

of radiating ridges and tubercles has been smoothed.

Family Anomalinidae Cushman, 1927

Subfamily Cibicidinae Cushman, 1927

Genus Cibicides De Montfort, 1808

Cibicides fletcheri Galloway & Wissler, 1927 s.1.

Fig. 13A—C

Cibicides fletcheri Galloway & Wissler, 1927: 64, pl. 10 (figs 8a—c, 9a—c). Lankford & Phleger,

1973: 117, pl. 6 (figs 1la—c).

Cibicides cf. C. fletcheri Galloway & Wissler. Boltovskoy et al., 1980: 24, pl. 8 (figs 17-21).

Remarks

Three rather damaged tests, compressed, and characterized by a small,

rounded, domed umbilical infilling on the ventral side, and a flat and wide boss,

flush with the test surface, in the dorsal umbilicus. These tests are similar to those

detailed by McMillan (19875) from the latest Eemian-—earliest Weichselian and

Holocene of the Orange Shelf.

The correct taxonomic name for this form is difficult to determine; so many

similar, though distinctly different morphotypes have been assigned to Galloway

154 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 13. A-C. Cibicides fletcheri Galloway & Wissler s.l. A. Ventral view, CTC 3. x 174.

B. Side view, same specimen. X 168. C. Dorsal view, same specimen. X 174. D-F. Cibicides

lobatulus (Walker & Jacob). D. Dorsal view, CTC 4. X 88. E. Ventral view, CTC 4. x 82.

F. Side view, CTC 4. x 146. G-H. Cibicidoides sp. G. Side view, CTC 3. X 84. H. Ventral

OY, CIE BZ, X< WY,

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 155

& Wissler’s (1927) species that it is unclear what the limits of the species should

be. A comparable species is Anomalina schmitti, described by Cushman &

Wickenden (1929) from present-day sediments off Juan Fernandez Islands, off the

west coast of South America. It is not clear how distinct are these two species.

Cibicides fletcheri was first described by Galloway & Wissler (1927) from the

Pleistocene of the Palos Verdes Hills of California.

Cibicides lobatulus (Walker & Jacob, 1798)

Fig. 13D-F

Nautilus lobatulus Walker & Jacob, in Kanmacher, 1798: 642, pl. 14 (fig. 36).

Cibicides lobatulus (Walker & Jacob) Murray, 1971: 175, pl. 73 (figs 1-7). Haynes, 1973a: 173,

pl. 20 (figs 1-2), pl. 21 (figs 3, 5-6), pl. 33 (figs 1-7), text-fig. 35 (nos 4-10). Boltovskoy

et al., 1980: 24, pl. 9 (figs 1, 3-4, ?2). Martin, 1981: 52, pl. 5 (figs 4-6).

Remarks

All tests possess narrow, elongate chamber flaps over the spiral suture, from

under which the secondary apertures are developed on the dorsal sides of the final

whorl of chambers. The shape of the flaps along the spiral suture is in keeping

with those seen on tests from the latest Pleistocene and Holocene of the Orange

Shelf (McMillan 1987b), off the west coast of southern Africa (Martin 1981), on

the Agulhas Bank (McMillan 1974—though not well drawn on pl. 8 (fig. 1c)),

and off the coast of Argentina (Boltovskoy et al. 1980). However, north-west

European tests (the type locality is Whitstable in Kent) tend to possess rather

more lobate chamber flaps (Feyling-Hanssen 1964; Knudsen 1971; Murray 1971;

Rosset-Moulinier 1972; Haynes 1973a). The difference is slight but it may prove

to be consistent between the two regions.

As with the latest Pleistocene and Holocene tests of Cibicides lobatulus from

the Orange Shelf (McMillan 19876), there are substantial differences in the

density of test perforations and in the overall test morphology of the Cape Town

shells. Spreading, more rounded-conical forms, often with a distinct peripheral

thickening, are usually densely perforate over the entire ventral side. The higher,

button-shaped, more biconvex tests, with a more rounded periphery and little

thickening, are generally only sporadically perforate on the ventral side, more so

over the surface of the earlier chambers.

Genus Cibicidoides Thalmann, emend. Loeblich & Tappan, 1955

Cibicidoides sp.

Figs 13G—H, 14A-B

Remarks

All tests of this species are rather broken. All are coarsely perforate on

both sides of the test, and exhibit ventral sutures that are more radiate and

156 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 14. A-B. Cibicidoides sp. A. Dorsal view, CTC 2. xX 112. B. Side view, same specimen

as Figure 13H. x 120. C-—D. Hyalinea balthica (Schréter). Side view, CTC 3. X 205.

D. Apertural view, same specimen. x 205. E-F. Sphaerogypsina globulus (Reuss). Different

views of the same specimen, CTC 3. E. Xx 137. F. x 144. G-H. Elphidium advenum

(Cushman). G. Side view, CTC 3. x 144. H. Apertural view, same specimen. X 146.

I. Elphidium sp. A. Side view, CTC 4. x 120.

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE SS7/

straight or weakly curved than is typical of the type species, Cibicidoides

mundulus (Brady, Parker & Jones), which is coarsely perforate only on the

dorsal side. This type of Cibicidoides is not known from the Orange Shelf

(McMillan 1987b), and would seem closer to the Cibicides cf. C. praecinctus

(Karrer) of McMillan (1974) from the Agulhas Bank and Martin (1981) from

scattered western offshore localities. These tests are distinctly more biconvex

than is typical for Cibicidoides pseudo-ungerianus (Cushman), widely found

off both the south and west coasts of southern Africa (McMillan 1974; Martin

1974, 1981).

Subfamily Planulininae Bermudez, 1952

Genus Hyalinea Hofker, 1951

Hyalinea balthica (Schréter, 1783)

Fig. 14C—D

Nautilus balthicus Schroter, 1783: 20, pl. 1 (fig. 2).

Hyalinea balthica (Schréter) Murray, 1971: 173, pl. 72 (figs 5-8).

Hyalinea balthica (Gmelin) Bremer et al., 1980: 23, pl. 3 (fig. 8).

Remarks

One specimen, rather broken and with tubular borings. The specimen is

typical for the species, which occurs widely over the western continental margin

of southern Africa at the present day (Pinto 1955; Martin 1974, 1981), as far south

as the westernmost Agulhas Bank (McMillan 1974). Hyalinea balthica ranges

throughout the latest Eemian—earliest Weichselian and Holocene deposits of the

Orange Shelf (McMillan 19876). It is known from the latest Eemian—earliest

Weichselian sediments of the Agulhas Bank (borehole F—A 13) (McMillan 1986),

and at that time ranged as far east as Algoa Bay, where it is seen in the coastal

Unit III of the ‘Alexandria Formation’ (Rimke in Engelbrecht etal. 1962;

Salmon 1981; McMillan in press).

In the latest Eemian—earliest Weichselian Unit III, Hyalinea balthica occurs

in littoral sands but, in the Holocene and at the present day, the species is more

typical of middle to outer shelf environments and deeper; Martin (1981) gave its

depth range as 92 m to 995 m off the west coast. As is the case with the Cape

Town test, latest Pleistocene shells would appear to be slightly thicker walled than

those from the Holocene, but there seems to be no distinct morphological

difference.

Hyalinea balthica has not been found in earlier Pleistocene deposits in South

Africa, though this may be due to the depositional facies of the rocks. Certainly,

though, the species cannot be considered as a marker for the Pliocene-

Pleistocene boundary on the continental shelf of southern Africa, as has been

done elsewhere (see Bayliss 1969).

158 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Acervulinidae Schultze, 1854

Genus Sphaerogypsina Galloway, 1933

Sphaerogypsina globulus (Reuss, 1848)

Fig. 14E-F

Ceriopora globulus Reuss, 1848: 33, pl. 5 (fig. 7a—c).

Sphaerogypsina globulus (Reuss) Loeblich & Tappan, 1964: C698, fig. 569 (nos 1-2).

Remarks

Several subspherical tests, although damaged, some with a flattened surface

that may have attached to a substrate, are referable to this species. One or two

examples were seen in the latest Pleistocene deposits of the Orange Shelf

(McMillan 1987b), but the species is more numerous in the Cape Town samples.

In most of the studied tests, the thin perforated walls over each chamberlet are

usually damaged or entirely worn away, and only the thicker intercameral walls

remain visible, as in the illustrated test. Similar forms, though larger and more

nearly spherical, occur in the Early Miocene (Burdigalian) shelly, white lime-

muds at a number of localities on the South African continental margin.

Family Elphidiidae Galloway, 1933

Subfamily Elphidiinae Galloway, 1933

Genus Elphidium De Montfort, 1808

Elphidium advenum (Cushman, 1922)

Fig. 14G-H

Polystomella subnodosa (non Minster) Brady, 1884: 734, pl. 110 (fig. 1a—b).

Polystomella advena Cushman, 1922: 56, pl. 9 (figs 11-12).

Elphidium advenum (Cushman) Albani, 1968a: 111, pl. 10 (fig. 6).

see Elphidium advena (Cushman) Apthorpe, 1980: 211, pl. 26 (figs 10-11).

Remarks

Most later authors studying Indo-Pacific material appear to have followed the

illustrations of Brady (1884) rather than those given by Cushman (1922) for their

identification of Elphidium advenum. The septal pits of Indo-Pacific tests of the

species are short and never as elongate as those of the holotype, which is from the

Tortugas region of the West Indies. It may be, as suggested by Cushman (1922),

that the Indo-Pacific shells assigned to this species are of a slightly different form

than the tropical Atlantic Elphidium advenum s:.s.

From studies of the ‘Alexandria Formation’ of the eastern Cape (McMillan

in press), it is clear that Elphidium advenum, as here understood, ranges from the

present day to near the base of the Pleistocene (and includes Unit II). It can thus

be considered as a useful marker species for the South African Quaternary, in

company with Pararotalia nipponica (Asano) and Cibicides lobatulus (Walker &

Jacob). The species is extensively distributed off the west, south and east coasts

of southern Africa at the present day, and tends to predominate in shallow marine

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 159

environments (Albani 1965; Martin 1981; unpubl. studies), although-off the west

coast it ranges into waters as deep as 418 m (Martin 1981), and small numbers of

specimens occur as deep as 127 m on the Agulhas Bank (McMillan 1974). It was

also widely distributed in latest Eemian-—earliest Weichselian times in shallow

marine environments around southern Africa, from the Orange Shelf (McMillan

1987b) to the Uloa Formation.

Elphidium sp. A

Fig. 141, 1I5A

Elphidium advenum (non Cushman) McMillan, 19875, pl. 17 (figs 1-2).

Remarks

A small proportion of the tests, initially assigned to Elphidium advenum by

McMillan (19875), lack the distinctive peripheral carina and wide, flat-topped and

flush umbilical boss of that species. Instead, the umbilical boss is small, almost

like a pinhead, and sited in a narrow umbilicus that is not entirely occupied by the

boss. The later chambers are slightly inflated, the test size is somewhat greater,

and the septal pits are more sharply defined, rather deeper, and more elongate

than true Elphidium advenum.

Elphidium sp. A is apparently confined to the latest Pleistocene of the west

coast, and is absent in the Holocene. The differences from true Elphidium

advenum are sufficient for this form to be considered a separate, possibly new

species but insufficient specimens occur in the Cape Town samples for a formal

taxon to be erected here.

Elphidium articulatum (d’Orbigny, 1839)

Fig. 1SB—C

Polystomella articulata d’Orbigny, 1839b: 30, pl. 3 (figs 9-10).

Elphidium articulatum (d’Orbigny) Boltovskoy, 1963: 61, pl. 6 (fig. 15). Boltovskoy et al., 1980:

29, pl. 13 (figs 1-4).

Remarks

Distinct, though slight differences exist in the style of the umbilical

ornamentation of Cape Town tests compared with those illustrated by McMillan

(1987b) from the Orange Shelf. The Orange Shelf examples reveal a rather

depressed umbilicus ornamented with 20 or more thin calcite pillars. In the

illustrated Cape Town shell, there are no more than ten pillars, and these are

often merged together, or with the umbilical terminations of the chambers; they

also tend to be larger in size than the pillars of the Orange Shelf tests. The septal

pits, both in their numbers and outline, the style of the aperture, the number of

chambers in the final whorl, the chamber morphology and the overall test

morphology of the Orange Shelf and Cape Town tests are all closely comparable.

Those from Cape Town show an umbilical arrangement somewhat closer to the

160 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 15. A. Elphidium sp. A. Oblique apertural view, same specimen as Figure 14]. x 113.

B-C. Elphidium articulatum (d’Orbigny). B. Apertural view, CTC 4. x 84. C. Side view,

same specimen. X 86. D-E. Elphidium crispum (Linné) s.l. D. Apertural view, CTC 4.

ZS Wake 1 Be Side view, CTC 4. x 83. F-G. Elphidium macellum (Fichtel & Moll) s.1. F. Oblique

apertural view, CTC 4. x 95. G. Side view, same specimen. x 95. H-I. Elphidium sp. B.

H. Side view, CTC 3. xX 178. I. Apertural view, same specimen. X 178.

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 161

individuals illustrated by Boltovskoy et al. (1980) from the coast of Argentina.

However, the Argentinian tests are more strongly biconvex than those from

southern Africa, if the apertural view (Boltovskoy et al. 1980, pl. 13 (fig. 3)) can

be regarded as typical.

D’Orbigny (1839b) figured a specimen that is substantially more lobate

around the test periphery than is the case in more recently illustrated South

Atlantic tests; his specimens were from the Rio Negro coast of Patagonia and

from off the Falkland Islands. Elphidium articulatum appears to range from the

latest Eemian—earliest Weichselian to the present day in southern Africa. It is

typical of innermost shelf, cold-water locations on the west coast, and occurs in

the present-day sands washed into the canalized mouth of the Soutrivier, just

north-east of Cape Town city centre.

Elphidium crispum (Linné, 1758) s.1.

Fig. 1SD-E

Nautilus crispus Linné, 1758: 709 (figured by Plancus, 1739, pl. 1 (fig. 2d—-f)).

Elphidium crispum (Linné) Cushman & Leavitt, 1929: 20, pl. 4 (figs 3a—b, 4a—b). Hageman,

1979: 94, pl. 5 (fig. 6a—b).

Remarks

Preliminary details of the taxonomic complexity surrounding this species and

Elphidium macellum (Fichtel & Moll), and the relationship of these two

Mediterranean species to comparable southern African forms, were given by

McMillan (19876). The following are supplementary notes.

Examination of authors’ illustrations of Elphidium crispum from the

Mediterranean Sea shows a wide range of forms. All the original figures are

somewhat stylized drawings (Plancus 1739; Gualtieri 1742), and it is upon these

that Linné based his concept of this species. The tests detailed by Gualtieri appear

to display an immense number of chambers in the final whorl, whereas those

figured by Plancus seem to contain about 20 chambers.

Cushman & Leavitt (1929), in an attempt to distinguish clearly between

Elphidium macellum and Elphidium crispum, re-examined the illustrations of

both species as figured by Fichtel & Moll (1798), and compared these with

specimens ‘from as near the type locality as possible’. In the case of Elphidium

crispum, it is worth including here Cushman & Leavitt’s full description:

‘Test of large size for the genus, about 21/2 times longer than wide in

peripheral view, lenticular, completely involute, the umbilical regions with a

medium sized, slightly projecting, rounded boss of clear shell material, the surface

of which is set with 10—12 small, shallow, rounded pits, periphery sharply angular

but not sharply keeled, sometimes becoming slightly lobulate and blunter in the

last portion of the adult coil; chambers very numerous, 20—40 in number, long,

narrow, the last few often inflated, forward part slightly raised, giving a radially

ribbed appearance; sutures somewhat sigmoid, the middle portion of each nearly

162 ANNALS OF THE SOUTH AFRICAN MUSEUM

radial, partially obscured by the retral processes, set with pores, those of the

earlier ones indistinct but later ones with large and deep pores; wall finely

perforate with about 12 relatively long retral processes extending well up onto the

side of the next added chamber, appearing as fairly widely spaced, low elevations

of the wall in the earlier chambers, but as thicker and more rod-like structures in

the later chambers; aperture, a row of small openings between the retral processes

at the base of the apertural face, which is low, sagittate, the sides often slightly

concave, the ends pointed. Diameter up to 3 mm.’

Such large-sized tests of E/phidium occur nowhere in South Africa, living or

fossil. South African tests assigned to this species possess only from 17 to

22 chambers in the final whorl of adult specimens, whereas juvenile examples

reveal rarely more than 12. The South African individuals tend to be more

biconvex rather than lenticular, normally with a small, sharply defined,

well-raised umbilical boss that exhibits a maximum of seven pits on its surface,

rather than the 10-12 described by Cushman & Leavitt (1929). Closer similarity

would seem to exist between the style and numbers of the septal pits and bridges,

though perhaps the tests figured by Cushman & Leavitt (1929) are a little stylized

in this respect.

South African shells feature a rounded to weakly sub-rounded, never acute,

test margin that reveals a hint of thickening along the earlier part of the final

whorl, whereas the periphery of the Italian form is sharply angular. However,

some latest Pleistocene tests of Elphidium crispum clearly show abrasion along

the periphery, which in severe cases may wear through to the chamber cavities. It

thus seems likely that the test periphery has been substantially amended on

occasion by post-mortem transport, probably through rotating and cartwheeling

of these shells in the littoral sands before final deposition. Nevertheless, despite

the probability of some post-mortem changes to these Elphidium crispum shells,

there are clearly distinct differences between southern African and Mediterra-

nean tests. Although the differences are not great, and the southern African

examples clearly fall within Elphidium crispum s.\., nevertheless they can be

confidently regarded as a local, distinctive form, which is easily distinguished from

Elphidium crispum s.s.

Unfortunately, this fairly simple picture is complicated by authors’

changing attitudes to species through the twentieth century. Cushman &

Leavitt (1929) evidently based their descriptions on adult specimens only, for

they noted that a considerable difference exists between the test morphologies

of microspheric and megalospheric, and juvenile and adult tests of both

Elphidium crispum and Elphidium macellum; these changes mostly remain to

be documented. Thus, for Cushman & Leavitt (1929), the two species are

easily distinguishable, but later authors have tended to see the two as

end-members with substantial variation between them. In addition, later

authors’ illustrations are of smaller-sized (?juvenile) tests, e.g. Kruit (1955,

pl. 2 (fig. 10)) from the Rhéne delta, Hageman (1979, pl. 5 (fig. 6a—b)) from

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 163

the Plio—Pleistocene of the western Peloponnesus of Greece, and Jonkers

(1984, pl. 12 (fig. 4)) from the Pliocene of Crete. Hageman (1979) reported on

a continuous gradation from lenticular tests with a large umbilical boss (typical

Elphidium crispum) to compressed tests lacking or possessing a small umbilical

boss (closely resembling typical Elphidium macellum), and Jonkers (1984)

followed similar methods of interpretation. Despite this rather confusing

picture of Mediterranean Elphidium crispum, the South African tests appear

to remain distinct from the species in a strict sense. However, to avoid

increasing the range of names available for this group of Elphidium, the

southern African tests are here referred to Elphidium crispum s.1.

Elphidium crispum s.\. occurs in large numbers in the Cape Town samples

and elsewhere around the south-western Cape coast in deposits of the same age

(see Appendices A to C). The species is not found in deposits older than latest

Eemian-—earliest Weichselian in the South African Pleistocene, though similar

species, of the Elphidium crispum—macellum group, occur in both the Varswater

Formation and in the older ‘Bredasdorp Formation’ (both Unit IA). Elphidium

crispum s.l. became extinct during the Weichselian or at the end of the

Pleistocene, and it is not found in situ in any Holocene deposits in South Africa.

This species occurs in smaller numbers in the latest Eemian—earliest

Weichselian of the Orange Shelf (McMillan 1987b) than at Cape Town. It is also

present close to the coast in the youngest parts (Unit III) of the ‘Bredasdorp

Formation’ and the “Alexandria Formation’, at Plettenberg Bay, and on the

Agulhas Bank (McMillan 1974); on the inner shelf it may occur in considerable

numbers, reworked into the Holocene. The species is also known from the Aliwal

Shoal (Carter 1966) and the Uloa Formation of Zululand (pers. obs.), but there

are slight morphological differences evident in these warmer-water tests that have

yet to be defined. Elphidium crispum s.1. thus seems to be an ideal zone fossil for

all the littoral or sublittoral deposits of the latest Eemian—earliest Weichselian

regression. It is restricted to the same time interval over which Ammonia sp. B

(Alexandria) and Ammonia sp. C (Alexandria) range (McMillan 1987a), but is

better developed in the colder waters of the west coast, where the two Ammonia

species are absent (see Fig. 3).

Elphidium crispum s.\. tests from studied localities are invariably broken or

abraded, and often the entire outer whorl of chambers is missing, leading to

strongly raised umbilical bosses, which tend to better survive the abrasion

processes. McMillan (19876: 565) considered the possibility of Elphidium crispum

s.l. and Elphidium macellum s.\. being abraded and clean tests, respectively, of

the same species. On the Orange Shelf, the former is most common in the sandier,

littoral parts of studied sections; the latter, always well preserved, is most

frequent in the clayier intervals that presumably accumulated in quieter water

conditions. There thus appears to be rather different facies controls on the two

species. However, examination of the test outlines of the two, particularly their

cross-sections, reveal distinct differences. Sectioned tests of Elphidium macellum

s.l. are far too slender and lenticuline throughout their course of growth to

164 ANNALS OF THE SOUTH AFRICAN MUSEUM

accommodate a kernel in their interior of the more strongly biconvex Elphidium

crispum s.1. (or vice versa).

Elphidium macellum (Fichtel & Moll, 1798) s.1.

Fig. 15F-G

Nautilus macellus Fichtel & Moll, 1798: 66, var. B, pl. 10 (fig. h—-k).

Elphidium macellum (Fichtel & Moll) Cushman & Leavitt, 1929: 18, pl. 4 (figs la—b, 2a—b).

Régl & Hansen, 1984: 50, pl. 14 (figs 2, S—6), pl. 15 (figs 1-2), text-fig. 18B.

Remarks

As with Elphidium crispum (Linné), local, southern African populations of

Elphidium macellum, though closely similar to those from the type area of the

Mediterranean Sea, nevertheless show some distinct and consistent differences.

Cushman & Leavitt (1929), in their examination of Elphidium macellum from

Rimini on the Adriatic coast of Italy, gave the following description:

‘Test of medium size for the genus, about 31/2 times longer than wide in

peripheral view, lenticular, planispiral, completely involute, umbilical regions

flat, periphery angular, slightly more rounded in the last-formed chambers,

slightly keeled, somewhat lobulate; chambers numerous, averaging 17 in adults in

the last-formed coil, slightly arched giving a ribbed appearance to the test; sutures

slightly depressed, those of the last-formed chambers slightly more so, partially

obscured by the retral processes, curved backward strongly toward the periphery,

the proximal half nearly radial, set with indistinct pores; wall thin, finely

perforate, usually with less than 12 relatively long retral processes, more widely

spread than in E. crispum and extending from well up on the side of the chamber

to the front of the preceding and appearing as narrow, fairly widely spaced, low

elevations of the wall; aperture, a row of small openings between the retral

processes at the base of the apertural face which is convex, sagittate, with the

sides slightly convex, the lobes sharply angled, saddling the preceding coil.

Diameter typically less than 1 mm; thickness 0,25 mm.’

Cushman & Leavitt (1929) concluded by distinguishing Elphidium macellum

from Elphidium crispum and Elphidium striatopunctatum (Fichtel & Moll) by its

keeled periphery that is often lobate, the fewer chambers in the final whorl, and

the flattened or depressed umbilicus.

More recently, the Fichtel & Moll collections have been rediscovered in

Vienna (Hansen & Roégl 1980), and a full re-appraisal of their species of 1798,

including Elphidium macellum, has been achieved (R6gl & Hansen 1984). In the

case of Elphidium macellum itself, Régl & Hansen (1984: 51) described and

illustrated a lectotype from the Fichtel & Moll collection, which thus must be

regarded as the nomenclatural type. This specimen possesses a circular,

non-lobate outline, a sharply angled and keeled periphery, 16 and a half chambers

in the final whorl, curved sutures with 8 to 15 septal bridges per suture, slightly

depressed umbilicus, lacking ornamentation or plug, apertural face low, sagittate,

with fine tubercles, and interio-marginally an equatorially placed aperture

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 165

composed of multiple openings with small protruding lips; maximum diameter

0,79 mm; maximum thickness 0,32 mm.

On comparing the South African tests with the illustrations and descriptions of

Rogl & Hansen (1984), it can be seen that the local tests differ in a number of

skeletal features. These shells are more compressed, more slender than either the

lectotype of R6gl & Hansen (1984) or the comparative Rimini tests illustrated by

Cushman & Leavitt (1929). They contain from 13 to 19 chambers in the final whorl,

with an apparent increase in numbers with increasing test size (McMillan 19875).

Most significantly, the umbilical area of the southern African tests is infilled with a

low, irregular plug of varying size that is ornamented over its surface with one to

seven pits, which in well-preserved examples can be seen to be spinose or

tuberculate interiorly. The plug is never as large or as prominent as that seen in

Elphidium crispum s.\. tests from the latest Pleistocene of south-western Africa, but

it clearly shows variation in size and height (McMillan 19875, pl. 18 (figs 1-6)). The

southern African shells are also not quite as acutely carinate on the periphery of the

test as the lectotype of Rogl & Hansen (1984). Because these differences in test

morphology appear to be consistent distinctions between the Mediterranean and

southern African tests, the latter are here termed Elphidium macellum s.1., for the

same reasons as expressed for Elphidium crispum s.1.

It should be added that the southern African shells of Elphidium macellum

s.l. compare closely, especially in their umbilical ornament, with some figured

from the coast of Patagonia (Thomson 1978, pl. 2 (fig. 1)). The test illustrated by

Brady (1884, pl. 110 (fig. 11a—b)) is of the same group, though the arrangement

of pits over the surface of the umbilical infilling has not been found in so

symmetrical a pattern in any of the southern African examples examined by the

present author, and is perhaps a little stylized. Tests of this group have also been

illustrated by Martin (1981, pl. 6 (fig. 12, and probably also fig. 10)) and

McMillan (1974, pl. 9 (fig. 7a—b)).

Elphidium macellum s.\. ranges from the latest Eemian—earliest Weichselian

(Unit III) deposits through to the present day around South Africa, and shows a

slight preference for cooler water conditions. The species is also one of the group

that occurs on the shelf during the Holocene (Martin (1981) gave its single

occurrence in her samples as at 118 m water depth, and it was recognized by

McMillan (19875) between 137 m and 183 m on the Orange Shelf), but it is more

typical of sublittoral, fairly quiet water conditions in the latest Pleistocene.

Spinose juveniles, such as are typical of north-west European waters (see Haynes

1973a: 202, pl. 24 (figs 2—-3)) do not occur in South Africa, either fossil or extant.

Elphidium sp. B

Fig. 1SH-I

Remarks

One specimen, slightly reminiscent of Elphidium gunteri Cole, as described

by McMillan (19876) from the Orange Shelf but differing on several points. There

166 ANNALS OF THE SOUTH AFRICAN MUSEUM

are 12 chambers in the final whorl, intersected by deeply incised, curved to

straight, radiate sutures that are crossed by up to five septal bridges per side, the

bridges appearing only on the later sutures. The umbilical chamber terminations

are rounded and slightly nodose, and the umbilicus is fairly wide and shallow,

being ornamented with a granular, almost sugary surface. The test wall is finely

and densely perforate; the test periphery is broadly and uniformly rounded, the

margin being sub-circular and continuous. The foramen, possibly composed of

two arched interio-marginal openings at the base of the terminal face, is overlain

by a zone of tuberculate ornamentation. Elphidium gunteri tests display a more

lobate test periphery, coarser test wall perforations, less incised sutures, and an

umbilical ornamentation of small numbers of irregularly shaped calcite pillars.

Genus Astrononion Cushman & Edwards, 1937

Astrononion echolsi Kennett, 1967

Fig. 16A—-B

Astrononion echolsi Kennett, 1967: 134, pl. 11 (figs 7a—b, 8). Fillon, 1974: 139, pl. 6 (figs 1-3).

Astrononion stelligerum (non d@’Orbigny) McMillan, 1987b: 422, pl. 19 (figs 1-3).

Remarks

A reconsideration of the Astrononion specimens described and illustrated by

McMillan (1987b) from the latest Eemian—earliest Weichselian of the Orange

Shelf suggests that they should rather be referred to the Antarctic species

Astrononion echolsi.

Le Calvez (1974) has examined the type specimen of Nonionina stelligera

d’Orbigny, found it to be unusable, and has proposed a neotype (Le Calvez 1974:

37, pl. 9 (figs 1-4)) from D’Orbigny’s Canary Island (Teneriffe) specimens. The

neotype is strongly compressed, with ten chambers in the last-formed whorl,

deeply incised sutural slits outside of the umbilical cover plate(s), and a broad,

shallow umbilicus almost entirely covered, except centrally, by the cover plate(s).

In contrast, the present specimen and those from the Orange Shelf possess

only seven or eight chambers in the final whorl, rather more inflated chambers

and a lobate test periphery, wider, more open and shorter incisions of the sutures,

and a less extensive cover plate revealing more of the central umbilicus. The Cape

Town specimen is a little more strongly biconvex than is typical for the Orange

Shelf shells. This species would seem to be confined to the cold-water influenced

latest Eemian and earliest Weichselian deposits of the west coast of South Africa.

Family Rotaliidae Ehrenberg, 1839

Genus Ammonia Brtinnich, 1772

Ammonia japonica (Hada, 1931)

Fig. 16C-E

Rotalia japonica Hada, 1931: 137, fig. 93a—c.

Ammonia japonica (Hada) Matoba, 1970: 48, pl. 5 (fig. 14a—-c), pl. 6 (fig. 1la—c). McMillan,

1987a: 37, text-fig. 7a—d.

Ammonia beccarii (non Linné) Martin, 1974: 84, fig. 14-1 (part.). Salmon, 1979: 77, fig. 3p.

Martin, 1981: 48, pl. 3 (figs 2-3).

Ammonia beccarii (Linné) var. inflata (non Seguenza) McMillan, 1974: 62, pl. 6 (fig. 2a—c).

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 167

ONS

Fig. 16. A-—B. Astrononion echolsi Kennett. A. Side view, CTC 4. x 139. B. Apertural view,

same specimen. X 157. C-—E. Ammonia japonica (Hada). C. Ventral view, CTC 4. x 111.

D. Side view, same specimen. X 124. E. Dorsal view, same specimen. Xx 110. F—H. Ammonia

parkinsoniana (d’Orbigny) s.l. F. Side view, CTC 4. Xx 135. G. Dorsal view, CTC 4. x 111.

H. Ventral view, same specimen as F. X 142.

168 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

Subsequent to the comments on Ammonia japonica by McMillan (1987a,

1987b), additional studies have been made of the Pleistocene littoral deposits of

the Saldanha—Langebaan region, the ‘Bredasdorp Formation’ and the ‘Alexan-

dria Formation’ of the southern Cape coast, and the Uloa Formation and Bluff

Sandstone Formation of Zululand/Natal. As was mentioned by McMillan (1987a),

Ammonia japonica is best developed in the Holocene off the south and west

coasts of South Africa, and is similarly distributed in the latest Eemian—earliest

Weichselian (Unit III) deposits. The species is absent in older Pleistocene and

Pliocene marine rocks of this region, although a similar form does occur in the

Varswater Formation (Unit IIA) of the Langebaan—Saldanha area, that is of the

same Ammonia-group as Ammonia Japonica.

Holocene tests tend to possess numerous, rather granular calcite pillars in the

umbilicus (Martin 1981, pl. 3 (fig. 2); McMillan 19875, pl. 19 (figs 12, 15-16)),

but latest Pleistocene shells are usually characterized by more robust umbilical

pillars, which may often merge into sinuous, irregular structures (McMillan

1987b, pl. 20 (figs 1-2)). It is probable that the difference is due to abrasion of the

Pleistocene tests but not the Holocene ones. However, variation in test

morphology through time is also a possible cause.

Rouvillois (1974) described Pseudoeponides falsobeccarii from the continen-

tal shelf (60-120 m water depth) off the west of France, and it is clear that

southern African Ammonia japonica is closely related to this North Atlantic

species. Neither show much similarity to the genotype, Pseudoeponides japonicus

Uchio (in Kawai et al. 1950). Rouvillois (1974) identified both ventral and dorsal

openings in her species, but these are so unlike those of Pseudoeponides japonicus

that, together with major differences in test and chamber morphology, it seems

best for the moment to regard the French species as referable to Ammonia. The

‘secondary openings’ of the ventral side of Ammonia falsobeccarii occur in various

forms in all species of Ammonia, but the dorsal openings, present also in southern

African Ammonia japonica, clearly separate these two species from Ammonia

beccarii (Linné) s.s., Ammonia parkinsoniana (d’Orbigny) s.1., and other species

typical of more littoral environments. It may be that the deeper-water, shelf

species of Ammonia warrant a distinct genus.

A comparison of southern African Ammonia japonica with Ammonia

falsobeccarii evinces the following similarities and differences. Rouvillois’ species

possesses usually eight chambers, southern African tests display eight or nine,

more rarely seven or ten, chambers in the final whorl. Both species possess very

strongly convex ventral sides but the dorsal side of the southern African shells is

always markedly convex, whereas that of the French species is weakly convex,

almost flat. Variation in the arrangement and the style of the umbilical filling is

similar, with well-preserved tests exhibiting a granular or tubercular ornament

over the dense array of thin calcite pillars that occupy the umbilicus. The

arrangement of incisions, bordered with granules and small nodes along the spiral

and septal sutures, is much the same in both groups of specimens.

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 169

The detail of the secondary openings at the inner junctions of the septal and

spiral sutures cannot be seen in the illustrations of Ammonia japonica given by

Hada (1931) and Matoba (1967, 1970). Their figures appear to be derived from

light microscope photography and drawings, and the depressions on the dorsal

side may thus either be obscured by reflection effects within the shell, or they are

absent in the Japanese tests. For the present, Hada’s name has been retained for

these southern African tests.

McMillan (1974) employed the name Ammonia beccarii (Linné) var. inflata

(Seguenza) for this species on the Agulhas Bank. The dorsal sides of Agulhas

Bank tests are generally less convex than those of tests from off the west coast

(McMillan 1987a). Rosalina inflata was originally described by Seguenza (1862)

from the ‘Pleistocene abbastanza recente’ clays of Catania, Sicily. Again, the

ventral side of the test is strongly convex, the dorsal side less so. Some variation

can be seen in the tests illustrated by Seguenza (1862, pl. 1 (figs 6, 6a—c)) but,

although the umbilical ornamentation and the test outline are similar to southern

African Ammonia japonica, the beaded bordered incisions of the sutures (in

fig. 6b) may extend to the test periphery, and the ventral surfaces of the chambers

may be strongly ornamented with irregular rugosities (figs 6a, 6c). Billman et al.

(1980: 86, pl. 5 (figs 1-14)) have illustrated tests referred to Ammonia inflata

(Seguenza) from northern Morocco, but noted that the species should be revised

using topotypic material. The Moroccan tests possess incised sutures with beading

over the entire ventral side; over the dorsal side the same ornamentation occurs,

but it is less well developed along both septal and spiral sutures of the early part

of the test. The umbilical infilling again consists of numerous calcite pillars. It is

felt by the present author that Ammonia inflata, as understood by Seguenza

(1862) or as differently interpreted by Billman et al. (1980), does not occur in

southern Africa. The Agulhas Bank tests are hence regarded as Ammonia

Japonica, with a local variation being the less high trochospire of the tests

compared to those seen off the west coast.

Although the species is known only as far back as the latest Eemian-—earliest

Weichselian in southern Africa, Ammonia japonica ranges back to the Early

Pleistocene in Japan (Matoba 1967).

Ammonia parkinsoniana (d’Orbigny, 1839) s.1.

Fig. 16F—H

Rosalina parkinsoniana d’Orbigny, 1839a: 99, pl. 4 (figs 25-27).

Ammonia parkinsoniana (d’Orbigny) forma tepida Poag, 1978: 397, pl. 1 (figs 1-4, 10-12,

17-18).

Ammonia parkinsoniana (d’Orbigny) forma typica Poag, 1978: 397, pl. 1 (figs 5-9, 13-16,

19-21).

Ammonia beccarii (non Linné) Martin, 1981: 48 (part.).

Ammonia parkinsoniana (d’Orbigny) s.1. McMillan, 1987a: 35, figs 3a—r, 4—S.

170 ANNALS OF THE SOUTH AFRICAN MUSEUM

Remarks

Extensively distributed around southern Africa in shallow marine environ-

ments in the Holocene and the latest Eemian—earliest Weichselian. The major

morphological variations of the species in the southern African Holocene have

been detailed by McMillan (1987a). Ammonia parkinsoniana s.1. occurs in the

quieter water, clayier and siltier intervals, as well as the more active, sandier parts

of the latest Pleistocene sections studied by McMillan (1987b) on the Orange

Shelf.

Genus Pararotalia Le Calvez, 1949

Pararotalia nipponica (Asano, 1936)

Fig. 17A-—C

Calcarina rotula (non Egger) Chapman, 1923: 3, pl. 1 (fig. 1).

Rotalia nipponica Asano, 1936: 614, pl. 31 (fig. 2a—c).

Rotalia ozawai Asano, 1951: 15, figs 115-117.

Pararotalia taiwanica (non Nakamura) Huang, 1964: 56, pl. 2 (fig. 2a—c).

Pararotalia sp. A Martin, 1974: 84, fig. 14-1 (part.).

Pararotalia cf. P. inermis (non Terquem emend. Le Calvez) McMillan, 1974: 63, pl. 6 (fig. 3a—c).

Pararotalia cf. P. nipponica (Asano) Martin, 1981: 48, pl. 9 (fig. 6).

Pararotalia nipponica (Asano) McMillan, 1987b: 443, pl. 20 (figs 13-17), pl. 21 (figs 1-3),

text-fig. 9.

Remarks

Studies of the ‘Alexandria Formation’ (McMillan in press) indicate that Para-

rotalia nipponica ranges from the earliest Pleistocene (Unit II) to the present day;

it is absent from the oldest, most inland part of the sequence (Early Pliocene,

Unit I). In this respect it has a stratigraphic range comparable to Cibicides

lobatulus (Walker & Jacob) and Elphidium advenum (Cushman).

Chapman’s (1923) record of Pararotalia nipponica in the Late Cretaceous

Mzamba Formation is clearly a contaminant from the overlying latest Pleistocene

sands; similar tests, markedly different in their preservation to the in situ

microfauna, were recognized by Makrides (1979). Neither Calcarina nor Para-

rotalia, nor any morphologically similar genera with peripheral blunt spines, are

known anywhere in the South African Late Cretaceous.

The reference to Rotalia dentata Parker & Jones by Chapman (1907) from the

?latest Pleistocene of the Buffalo River, East London, almost certainly is

referable here too. Siesser & Salmon (1979) reported ‘Pararotalia inermis’,

supposedly from the Late Eocene Langental Beds of southern Namibia, but

samples from pits dug in these outcrops yielded no trace of Eocene Pararotalia

species (unpubl. studies). The illustrations given (Siesser & Salmon 1979,

fig. 1Sc-d) are certainly of Pararotalia nipponica. It is evident that these

Pararotalia tests, in keeping with those of Elphidium and Ammonia, were

introduced into the Langental region with windblown sand derived from the

littoral. The presence of Elphidium cf. E. crispum (Linné) at Langental (Siesser

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 171

Fig. 17. A-—C. Pararotalia nipponica (Asano). A. Dorsal view, CTC 4. x 98. B. Ventral view,

CTC 4. x 122. C. Side view, same specimen as B. X 122. D. Globigerina (Globigerina)

bulloides d’Orbigny. Ventral view, CTC 3. x 172. E-F. Globorotalia (Globoconella) inflata

(d’Orbigny). E. Ventral view, CTC 3. x 181. F. Side view, CTC 3. x 235. G-H. Neoglobo-

quadrina cf. N. pachyderma (Ehrenberg). G. Ventral view, CTC 3. x 200. H. Side view,

Crea ex Zip:

iV ANNALS OF THE SOUTH AFRICAN MUSEUM

& Salmon 1979) perhaps suggests a latest Eemian—earliest Weichselian age for the

windblown contaminants.

Pararotalia nipponica occurs in the Varswater Formation (McMillan 1987b,

Appendix A) of the Langebaan area. It has also been reported as ‘Rofalia-like

foraminifers’ and ‘Rotalia’ in the Saldanha area (Visser & Schoch 1973). The

species is widespread throughout the ‘Bredasdorp Formation’ (unpubl. studies),

and was noted by Riimke (in Spies et al. 1963) under the name Rotalia audouini

d’Orbigny, though it is distinctly different from D’Orbigny’s European Eocene

and Oligocene species. In the “Alexandria Formation’ the species is widespread

and abundant in the Early Pleistocene Unit II and latest Pleistocene Unit IIJ—

again previously recognized by Riimke (in Engelbrecht et al. 1962) as Rotalia

audouini.

Specimens of ‘Rotalia sp.’ from the Bluff Sandstone Formation at Durban,

described by Parr (1958), are certainly referable to Pararotalia nipponica. Some of

this material was stored on grid slides at the Bernard Price Institute for

Palaeontological Research, University of the Witwatersrand, Johannesburg, and,

through the courtesy of Professor M. Raath, were examined by the present author.

Pararotalia nipponica also occurs, presumably in situ, in the latest Pleistocene

(Unit III) Pecten bed in the lower half of the Uloa exposure, Zululand. As Streblus

taiwanica (Nakamura), this species was encountered in Quaternary sands along the

Inharrime to Vilanculos coast of Mocgambique (Rocha 1965).

The Pararotalia nipponica-group exhibits wide morphological variation, parti-

cularly in the height of the trochospire, size of the umbilical boss, and in the spinose,

lobate or circular test outline. Attempts to divide the latest Pleistocene tests into

discrete morphological formae proved to be impossible for the Orange Shelf material

(McMillan 1987b). However, there does seem to be one tendency toward tests with

a high trochospire, circular or weakly lobate periphery, and large umbilical boss

(Pararotalia nipponica form), and one tendency to low trochospire, spinose

periphery, and small or divided umbilical boss (Pararotalia ozawai form). These two

trends resemble the distinctions made by Ujiié (1966) within the group. Bhalla (1972)

considered sexual dimorphism to be partly responsible for the wide range of

morphologies included within Pararotalia nipponica.

Pararotalia nipponica is closely comparable to Pararotalia serrata, described

by Ten Dam & Reinhold (1941) from the Pliocene of the Netherlands, and also

known from similarly aged Crag deposits of eastern England (Funnell 1981). The

precise differences between the two species remain to be determined.

Family Globigerinidae Carpenter, Parker & Jones, 1862

Genus Globigerina d’Orbigny, 1826

Globigerina (Globigerina) bulloides d’Orbigny, 1826

Fig. 17D

Globigerina bulloides d’Orbigny, 1826: 277, modéles no. 17 (1 ére livraison), no. 76 (4 éme

livraison).

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 173

Remarks

Only brief notes are given for this and the following planktonic species. One

test, with the last-formed chamber broken away. Compared with examples

encountered in the Holocene of the Orange Shelf (McMillan 1987b), this test

displays a rather low-arched aperture but is otherwise typical. The species is

characteristic of subpolar and transitional waters in the world oceans (Bé &

Tolderlund 1971). Kennett & Srinivasan (1983) regarded Globigerina bulloides to

range from the Middle Miocene to the present day.

Family Globorotaliidae Cushman, 1927

Genus Globorotalia Cushman, 1927

Globorotalia (Globoconella) inflata (d’Orbigny, 1839)

Fig. 17E-F

Globigerina inflata d’Orbigny, 1839c: 134, pl. 2 (figs 7-9).

Remarks

Two specimens, one rather more lobate than is typical. Both are sinistrally

coiled, as are almost all examples encountered by McMillan (19876) through the

Holocene of the Orange Shelf. The species is most typical of temperate water

masses (Bé & Tolderlund 1971; Stainforth et al. 1975). Its stratigraphic range is

Late Pliocene to present day (Kennett & Srinivasan 1983).

Genus Neogloboquadrina Bandy, Frerichs & Vincent, 1967

Neogloboquadrina cf. N. pachyderma (Ehrenberg, 1861)

Fig. 17G—H

see Aristerosira pachyderma Ehrenberg, 1861: 303; 1873: 386, pl. 1 (fig. 4).

Remarks

Two specimens of the shelf forma (non pachyderm), both sinistrally coiled

and both exhibiting a kummerform final chamber. These tests show similarities to

the shelf forma tests illustrated by McMillan (19875, pl. 22, figs 14-16) from a

water depth of 46,5 m, close to the Namaqualand coast. However, the presence

of the kummerform final chambers result in distinctly unusual apertures, and the

unique apertural arrangement of Neogloboquadrina pachyderma is not seen. For

this reason, the two tests are placed in the status of confer N. pachyderma.

Sinistrally coiled Neogloboquadrina pachyderma appears to be associated

with upwelling waters of the Benguela Current close to the west coast of southern

Africa (Bé & Tolderlund 1971). The species ranges from the Late Miocene to the

present day (Kennett & Srinivasan 1983).

ACKNOWLEDGEMENTS

I am most grateful to Mr V. H. Valicenti of SOEKOR (Pty) Ltd., Cape

Town, for sending portions of samples collected by him from the municipal

174 ANNALS OF THE SOUTH AFRICAN MUSEUM

trenching, for stratigraphy data on the samples, for much help in collecting the

samples detailed in the three Appendices, as well as for valued discussions

over many years on southern African Cainozoic and Mesozoic geology.

Professor J. R. Haynes of University College of Wales, Aberystwyth, kindly

made available facilities for sample processing, literature searching (including

the Ellis & Messina ‘Catalogue of Foraminifera’), photographic darkroom

facilities, and allowed access to the U.C.W. scanning electron microscope,

during completion of a Ph.D. thesis late in 1987. Comparative material from

the Varswater Formation was supplied through the courtesy of Mr V. H.

Valicenti and Dr A. J. Tankard (formerly South African Museum), from the

‘Bredasdorp Formation’ by Mr J. Malan (Geological Survey, Bellville), and

from the ‘Alexandria Formation’ by Messrs D. Stear and W. Smuts (both

formerly University of Port Elizabeth). Material from the Bluff Sandstone

Formation was kindly provided by Mr J. A. G. Cooper (University of Natal),

and some Uloa Formation material generously supplied by Mr G. Kruger

(Industrial Development Corporation).

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Reuss, A. E. 1848. Die fossilen Polyparien des Wiener Tertiarbeckens. Naturwissenschaftliche

Abhandlungen, Wien 2: 1-109.

Rocua, A. T. 1965. Contribuigéo para o estudo dos foraminiferos do Quaterndrio do sul da

provincia portuguesa de Mocambique. Revista ‘Garcia de Orta’, Junta de investigacoes do

ultramar 13 (3): 407-424.

ROGL, F. & Hansen, H. J. 1984. Foraminifera described by Fichtel & Moll in 1798—a revision

of Testacea Microscopica. Neue Denkschriften des naturhistorischen Museum, Wien

3: 1-143.

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 179

RossET-MOouLINIER, M. 1972. Etude des foraminiféres des Cétes nord et ouest de Bretagne.

Travaux. Laboratoire de Géologie, Ecole Normale Supérieure 6: 1-225.

RovuviLLois, A. 1974. Un foraminifére méconnu du plateau continental du Golfe de Gascogne:

Pseudoeponides falsobeccarii n. sp. Cahiers de Micropaléontologie 3: 1-9.

SaLmon, D. A. 1979. Quaternary foraminifers in piston cores from the S.W. Indian Ocean.

Technical Report. Joint Geological Survey South Africa/University of Cape Town Marine

Geoscience Unit 11: 72-79.

SALMON, D. A. 1981. Revised data on benthic Foraminiferida used as age-dating evidence in

some South African Tertiary sediments. Technical Report. Joint Geological Survey South

Africa/University of Cape Town Marine Geoscience Unit 12: 11-17.

ScHROTER, J. S. 1783. Einleitung in die Conchylienkenntniss nach Linné 1. Halle: J. J.

Gebauer.

SEGUENZA, G. 1862. Prime ricerche intorno ai rizopodi fossili delle argille Pleistoceniche dei

dintorni di Catania. Atti dell’Accademia gioenia di scienze naturali (2) 18: 84-126.

SIESSER, W. G. & SALMon, D. A. 1979. Eocene marine sediments in the Sperrgebiet, South

West Africa. Annals of the South African Museum 79 (2): 9-34.

Spies, J. J., ENGELBRECHT, L. N. J.. MALHERBE, S. J. & VILJOEN, J. J. 1963. Die geologie van

die gebied tussen Bredasdorp en Gansbaai: toeligting van blaaie 3419C en 3419D (Gansbaai)

en 3420C (Bredasdorp). Pretoria: Geological Survey of South Africa.

STAINFORTH, R. M., Lams, J. L., LUTERBACHER, H., BEARD, J. H. & JEFFoRDS, R. M. 1975.

Cenozoic planktonic foraminiferal zonation and characteristics of index forms. Paleonto-

logical Contributions. University of Kansas 62: 1-425 (2 volumes).

TANKARD, A. J. 1975. The marine Neogene Saldanha Formation. Transactions of the

Geological Society of South Africa 78 (2): 257-264.

TEN Dam, A. & REINHOLD, T. 1941. Die stratigraphische Gliederung des niederlandischen

Plio-Pleistozans nach Foraminiferen. Mededelingen van de Geologische Stichting

(C-S) 1: 1-66.

TERQUEM, O. 1876. Essai sur le Classement des Animaux qui Vivent sur la Plage et dans les

Environs de Dunkerque: Deuxiéme Fascicule: 55-100. Paris: Terquem. (Also 1877—

Mémoires de la Société dunkerquoise pour encouragement des sciences, des lettres et des

arts 20: 146-191.)

THompson, L. B. 1978. Distribution of living benthic Foraminifera, Isla de los Estados, Tierra

del Fuego, Argentina. Journal of Foraminiferal Research 8 (3): 241-257.

Ume, H. 1966. Shell structure of Japanese smaller Foraminifera. Part 2: Pararotalia nipponica

(Asano, 1936). Transactions and Proceedings of the Palaeontological Society of Japan

(n.s.) 61: 191-200.

VissER, H. N. & ScHocu, A. E. 1973. The geology and mineral resources of the Saldanha Bay

area. Memoirs. Geological Survey of South Africa 63: 1-150.

WALKER, G. & Boys, G. 1784. Testacea minuta rariora nuperrime detecta in arena littoris

sandvicensis. (A collection of the minute and rare shells lately discovered in the sand of the

sea-shore near Sandwich.) London: Walker, G.

WILLIAMSON, W. C. 1848. On the Recent British species of the genus Lagena. Annals and

Magazine of Natural History (2) 1: 1-20.

WILLIAMSON, W. C. 1858. On the Recent Foraminifera of Great Britain. London: Ray Society

WRIGHT, J. 1886. Foraminifera of the Belfast Naturalists’ Field Club’s cruise off Belfast Lough,

in the steam-tug Protector, June 1885; also Foraminifera found by Dr Malcomson, at

Rockport, Belfast Lough. Proceedings. Belfast Naturalists’ Field Club 1885-6 (Appendix):

317-326.

APPENDIX A

THE LLANDUDNO-SANDY BAY COAST, CAPE PENINSULA

Deposits of the same age as those described in the main body of this article

occur intermittently around the Cape Peninsula. Along the Llandudno to Sandy

Bay part of the coastline, ‘raised beach’ deposits occur about 10 m above present

180 ANNALS OF THE SOUTH AFRICAN MUSEUM

IG°20'E

Karbonkelberg

Fig. 18. Locality map of sampling sides along the Llandudno to Sandy Bay coast, Cape

Peninsula.

mean sea-level as unconsolidated, white shelly sands at 34°01'05”S 18°20'00’E,

just north of the Hamersteen (Fig. 18).

Because of their loose nature, they are poorly exposed and tend to wash

down slope, but occasional clear faces may be seen or excavated on either sides

of paths leading inland from, and parallel to the beach. Large shells, particularly

gastropods, are common and often well preserved. Three samples were collected

from the best available exposures late in 1986, and revealed the presence of

abundant, though rather poorly preserved, often leached, and abraded

foraminifera tests (sample SB—LL 3).

In addition, cemented shelly sandstone was noticed occasionally adhering to

the large granite slabs and boulders in the present-day surf zone. One sample of

sandstone revealed a microfauna very similar in composition to that of the ‘raised

beach’ deposits, but severely affected by calcite leaching and precipitation, and

somewhat less diverse.

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 181

The foraminiferal faunas of the four samples are closely comparable, both in

species and composition, to those from Cape Town city centre. It is clear that

these deposits near Llandudno are derived from the same regressional episode

during the latest Eemian—earliest Weichselian. The faunas are distinguished by

the predominance of Elphidium crispum (Linné) s.1., Pararotalia nipponica

(Asano) and Cibicides lobatulus (Walker & Jacob), the first-named confirming the

Unit III, latest Pleistocene age. Table 2 details the foraminifera recovered from

the four samples.

The absence of Ammonia parkinsoniana (d’Orbigny) s.1. in the Llandudno

samples is of note, and probably reflects the minute rainfall catchment area of this

part of the coast, with a corresponding lack of dilution of littoral sea-water. In

contrast, numbers of Cibicides lobatulus are substantially greater than at Cape

Town, probably indicating a more turbulent environment: a rocky shore rather

than a beach. Again, a littoral to sublittoral environment is indicated by the

foraminifera species present, though in such a wave-influenced shallow marine

environment, considerable post-mortem transport of tests must have occurred.

Macrofaunal debris is composed mainly of mollusc shell, some of which retains its

life colours. In addition, barnacle plates, echinoid spines and sponge spicules

occur.

TABLE 2

Distribution of foraminifera in studied samples from the latest Eemian—earliest Weichselian

deposits of the Llandudno to Sandy Bay coast, Cape Peninsula.

Sample number Sp. Species

SB-LL3 SB-LL2 SB-LL1 SB-1 O.

1 1 Spiroplectammina cf. S. wrightit

1 2 Quinqueloculina contorta

8) 2 2 5) 3 Quinqueloculina dunkerquiana

1 1 4 Quinqueloculina seminulum

3} 4 5 7 5 Quinqueloculina spp. (broken)

72 6 Siphonaperta sp.

1 7 Lagena perlucida

1 3 8 Lenticulina spp.

1 9 Bulimina gibba

1 2 10 Cassidulina laevigata

1 11 Rosalina bradyi

1 1 12. Rosalina cf. R. williamsoni

iL 13 Rosalina sp.

3 1 8) 14 Glabratella australensis

17 3D) 27 71 15 Cibicides lobatulus

1 2 4 4 16 Cibicides spp. (broken)

1 1 3 2 17 Cibicidoides sp.

1 18 HAyalinea balthica

1 19 Elphidium advenum

1 2 20 Elphidium sp. A

38) sy) 36 96 21 Elphidium crispum s.1.

1 1 22 Elphidium macellum s.l.

1 23 Elphidium sp. (broken)

ON 56 24 149 24 Pararotalia nipponica

80 173 105 349 Total number of tests

182 ANNALS OF THE SOUTH AFRICAN MUSEUM

Many of the species present are confined to the west-coast littoral at the

present day, and indicate that, as with the Cape Town fauna, water temperatures

were closely comparable to those experienced today along this coast. It seems

likely that the very close similarity of the foraminifera species in the latest

Eemian-—earliest Weichselian of the south-western Cape and the Orange Shelf

(McMillan 1987b) can be taken to indicate little difference in coastal sea-water

temperatures between the two areas in the latest Pleistocene.

APPENDIX B

LATEST PLEISTOCENE FORAMINIFERA FROM BERG RIVER MOUTH

Further north along the west Cape coast, near the mouth of the Berg River,

deposits of the same age (latest Eemian—earliest Weichselian, Unit III) as those

of Cape Town city centre are preserved beneath the Holocene estuarine

sediments. Recent dredging and channelling operations for a new marina at Port

Owen, just inside the estuary mouth, have excavated muddy and shelly sands that

have been used as fill over the adjacent area.

Two samples were collected in late 1986: one from a brown, clayey and

lignitic sand, apparently in situ, near the base of a telephone pole next to the

dredged marina channel (BR-2), and the other of muddy shelly sand dredged

from the bed of the channel (BR-1). The localities of the samples are shown in

Figure 19, and a list of the recovered foraminifera in Table 3. The two samples

were from closely adjacent sites at 32°47'25"S 18°09’05"E.

TABLE 3

Distribution of foraminifera in two studied samples from the latest Eemian—earliest Weichselian

deposits of the Berg River mouth, west Cape coast.

Sample number Sp. Species

BR-2 BR-1 no.

2 1 Bulimina elongata

3 2 Bulimina gibba

1 3. Trifarina angulosa

1 4 Cassidulina laevigata

1 5 Rosalina cf. R. globularis

1 6 Glabratella australensis

8 6 7 Cibicides lobatulus

1 8 Cibicides sp. (broken)

11 49 9 Elphidium advenum

1 21 10 Elphidium sp. A

18 3 11. Elphidium cf. E. alvarezianum

62 87 12. Elphidium crispum s.1.

10 13. Elphidium macellum s.1.

58 68 14. Ammonia japonica

1 1 15 Ammonia parkinsoniana s.l.

113) 48 16 Pararotalia nipponica

182 293 Total number of tests

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 183

LAAIPLEK

VELDDRIF

Fig. 19. Locality map of sampling sites at Berg River mouth, west Cape coast.

Both samples are rather gypsiferous, though this would seem to be a

secondary feature derived from the overlying Holocene estuary. Present-day

environments in the estuary are markedly hypersaline, and some areas are utilized

for salt production. However, the foraminifera of the latest Pleistocene Unit II

reveal a much more normal marine environment. Cibicides lobatulus (Walker &

Jacob) occurs in fairly small numbers—as at Cape Town but unlike Llandudno-—

and is suggestive of a sandy beach environment rather than a rocky shore.

Ammonia japonica (Hada) in such numbers clearly illustrates a good connection

to the open sea, whereas the lack of Ammonia parkinsoniana (d’Orbigny) s.1.

perhaps implies little fresh water entering the Berg estuary during the latest

Eemian-—earliest Weichselian. Very rare fragments of mollusc shell retain their

184 ANNALS OF THE SOUTH AFRICAN MUSEUM

life colours, whereas all of the foraminifera tests are bleached. Occasional tests of

Ammonia parkinsoniana s.\. with their natural brown colour are substantially

better preserved than the rest of the assemblage, and are considered to be

contamination from the Holocene estuarine deposits.

The brown sand contains much lignite and clay, and it yielded rather more

corroded foraminifera tests than the channel sample. It seems likely that the

clayey sand accumulated in a non-marine or estuarine mud-flat environment, as

plant fragments, seeds and non-marine gastropods also occur. The marine

foraminifera shells were probably carried off the adjacent beach and into the

estuary by onshore winds. However, the absence of Holocene foraminifera in the

clay clearly identifies it as having accumulated before the Holocene transgression.

The abundance of Elphidium crispum (Fichtel & Moll) s.1. in both samples clearly

shows these deposits to have been laid down at the time of the latest

Eemian-—earliest Weichselian regression.

APPENDIX C

LATEST PLEISTOCENE FORAMINIFERA OF HOEDJIESPUNT PENINSULA, SALDANHA

Although the main body of the white, shelly and sandy limestones that cap

the Palaeozoic and older rocks of the Hoedjiespunt Peninsula are the most

obvious latest Cainozoic deposits of this region, there nevertheless are small

residues of an even later unit. Foraminifera from shelly parts of the main

limestones (the Varswater Formation, and including the ‘Saldanha Formation’)

TABLE 4

Foraminifera species of one studied sample from the latest Eemian—earliest Weichselian sands of

Hoedjiespunt Peninsula, Saldanha, west Cape coast.

Sample Sp. Species

no. no

H-1

B) 1 Quinqueloculina dunkerquiana

2 2 Quinqueloculina seminulum

11 3. Quinqueloculina spp. (broken)

1 4 Siphonaperta sp.

1 5 Cassidulina laevigata

24 6 Glabratella australensis

9 7 Glabratella australensis (reworked)

5 8 Cibicides lobatulus

12 9 Cibicidoides sp.

A 10 Elphidium advenum

3 11. Elphidium crispum s.1.

6 12. Elphidium sp. (reworked)

1 13. ?Haynesina sp.

3 14. Ammonia spp. (reworked)

112 15 Pararotalia nipponica

21 16 Pararotalia nipponica (reworked)

238 Total number of tests

PLEISTOCENE FORAMINIFERA FROM CAPE TOWN CITY CENTRE 185

indicate a later Early Pleistocene age (see p. 132), regarded as equivalent in age

to Unit ITA of the ‘Bredasdorp Formation’.

However, bleached white, unconsolidated, shelly sands were recognized by

Mr V. H. Valicenti about 8 m above high tide mark, below the Control Tower,

on the north side of the Hoedjiespunt Peninsula, unconformably overlying the

Varswater Formation limestones. Though severely affected by rainwash down-

slope, and rather obscured by small plants and humus, as well as being

contaminated with weathered debris from the adjacent limestones, one relatively

clean sample was obtained. The locality is at 33°01'36"S 17°57'44’E, on Figure 20,

and the foraminifera assemblage is listed in Table 4.

Despite the apparent cleanness of the sample, reworked foraminifera from

the Varswater Formation do occur, but they can be easily identified by being

cemented, creamy in colour and well preserved. Jn situ tests, in contrast, are

I7°57 E

Outlier of

Varswater

Formation

limestones

+b ® =| HOEDJIESPUNT

BOMGAT

control

tower

Fig. 20. Locality map of the sampling site H—-1 at Hoedjiespunt, Saldanha, west Cape coast.

186 ANNALS OF THE SOUTH AFRICAN MUSEUM

bleached white, rather damaged, often displaying borings, poorly preserved, and

often appear rather ‘leached’.

Again, the age of the shelly sands is regarded as being latest Eemian—earliest

Weichselian, based on the presence of Elphidium crispum (Linné) s.l. The

absence of Ammonia parkinsoniana (d’Orbigny) s.l. would seem to duplicate the

situation along the Llandudno—Sandy Bay coast, with little coastal dilution of

seawater. As with the previous deposits studied, the mollusc shell often retains its

life colours, but unlike elsewhere, tests of foraminifera are relatively rare.

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.,

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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

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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

Sip: . 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 ea 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.

I. K. McMILLAN

FORAMINIFERA FROM THE LATE

PLEISTOCENE (LATEST EEMIAN TO

EARLIEST WEICHSELIAN)

SHELLY SANDS OF CAPE TOWN CITY

CENTRE, SOUTH AFRICA

LANAI

fF Pus OUTH AFRICAN

. : MUSEUM |

KCAPE "TOWN

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BuLtLoucu, 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)

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AN INVESTIGATION OF THE ‘COLDSTREAM STONE’

M. L. WILSON,

W. J.J. VAN RIJSSEN

Archaeology Department, South African Museum, Cape Town

D. A. GERNEKE

Electron Microscope Unit, University of Cape Town

(With 10 figures)

[MS accepted 23 July 1989}

ABSTRACT

The South African Museum has in its collections an unusually well-preserved, polychrome

painted stone recovered from archaeological deposits in the southern Cape Province.

Doubts have been expressed as to the authenticity of this artefact. Documentary research

and micro-analysis of the paints indicate that, while there are a number of points that cannot be

resolved, there is no evidence to refute the authenticity of the painting on the stone.

CONTENTS

PAGE

MMR OGMCHO Mean mire tern ce tent taec ral, we itn ra At in os eh avtyoisyeiel dime deal 187

the documentaryevidence: 2 it osc compa ecs soos Le nea ey hor ee nae 189

| DPONAY'S CEICENE TUONO eho Slgctlaned ara che aha nny coctio te mat Cite en tar te nee ere 192

STMOSSCIOGINE INVISION gr aeeita Ste le utr ecto y eecuera ol ae ect coats NG) Cues sees area 194

Phy sicalianalySes- Othe plementS awn em onde ea. sce aa meas sae 195

IRONS TON Ts Sasa taug kpseseesmer cy et etree a ca as We idler en Oe err 195

INCI palmtsianGipicmiem(Sy nea ee ee es key ee leony 197

NVihivespaintsrandiplenmentSan= criss sacle, acess cel ented dee eee e § 201

Blackspaintsand pilemenmtS aes owes es aes a oe oe ee eta oy8 201

DISCUSSION Maw re eee ao en eet te acai eien ais dyh Wage aiayer ea 4 206

ithe stylerolsthe;pamlines s.r ark eee eas er epee he cee ee Seka 206

DISCUSSION anGd’CONnClUSIONS =. ace re ye een es aus a ek os hers havent aneia le 3 207

PNCKMOW IEC CSTE TICS Tpit ney tee eh na acer Mae 2, sia es 8 2,

J RETTOT RETIN SOS a eo aastates oa been eelive, Sri ana ko. Repth aa Tater ata UAC Ds ee nea 212

INTRODUCTION

In 1911 a remarkable painted stone (Fig. 1F) was reported to have been

found in association with a human burial during excavations in a rock-shelter

near the mouth of the Lottering River on the southern coast of the Cape

Province of South Africa, in what is now the Tsitsikama Coastal National Park

(Fig. 2). To this day this artefact remains unique because of the unusual state of

187

Ann. S. Afr. Mus. 99 (6), 1990: 187-213, 10 figs.

188 ANNALS OF THE SOUTH AFRICAN MUSEUM

BOTSWANA MOZAMBIQUE

SOUTH WEST

AFRICA / NAMIBIA

SWAZILAND

LESOTHO

SOUTH AFRICA

ATLANTIC INDIAN

OCEAN OCEAN

TSITSIKAMA MOUNTAINS

COLDSTREAM

VILLAGE

OAKHURST

LOTTERINGS R.

WITELS R.

ELANDSBOS R.

COLDSTREAM

SHELTER

GEELHOUTBOS R.

INDIAN OCEAN

Fig. 2. Maps showing the location of the Coldstream shelter and Whitcher’s Cave.

AN INVESTIGATION OF THE ‘COLDSTREAM STONE’ 189

preservation of the painting and because it is the only known polychrome

painted stone to have come from archaeological deposits in South Africa.

This stone, together with the associated skeleton, was on display in the

South African Museum for a number of years. When the display was changed

the stone was put for safe-keeping in the Museum’s strong-room, stored in the

wooden box made for its transport from Coldstream, and was seen by very few

people, although a copy was later put on display in the Museum. The skeleton

was integrated into the Museum’s physical anthropology collections with the

others excavated from the shelter but, unfortunately, can now not be identified.

In 1980, pending the removal of most of the Museum’s staff and collections

to temporary premises during building operations, the ‘Coldstream Stone’, as it

has become known, was returned to the Archaeology Department’s artefact col-

lections, where it has since been seen more frequently by researchers. Sight of

the stone itself and comparison of it with other painted stones in the Museum’s

collections, such as those shown in Figure 1, have raised doubts that anything so

fragile as a painting could have survived in so remarkable a state of preser-

vation, given that it was said to have been found in the damp deposits of a

coastal rock-shelter.

Since it is clearly in the Museum’s interest, as well as that of other

researchers, to know whether or not this unique artefact is genuine, various lines

of enquiry were followed. These included documentary research, micro-analysis

of the paints and comparison with paints and pigments from other sources, and

the opinion was sought of those who knew the excavator. These points are dis-

cussed below.

THE DOCUMENTARY EVIDENCE

The first record of the finding in South Africa of painted stones associated

with human burials appears to have been a report by J. S. Henkel, Conservator

of Forests at Knysna, to the then Director of the South African Museum,

L. Péringuey. Henkel’s correspondence cannot now be traced, but Péringuey

wrote to him on 12 April 1909 ‘I never heard of painted stones or slabs placed

above the head. I’ll go further and say that nothing of this kind is known and

your announcement took my breath away’ (S.A. Museum Letter Book

1908-1910: 184).

Péringuey was obviously intrigued by Henkel’s report and possibly stimu-

lated by another letter from him, for he wrote to Henkel again on 7 July ‘I have

a group of, perhaps dancers, perhaps dead bodies judging from their attitude,

painted on a small stone, found in one of the Knysna grottoes or caverns’ (SAM

Letter Book 1908-1910: 376). Again, on 18 December, he wrote to Henkel

‘Hitherto painting or carving on stone could not be traced to these primitive

people—I mean the Strand Looper—except however a small painting which is

alleged, and I am sure rightly so, to have been found in a cave’ (SAM Letter

190 ANNALS OF THE SOUTH AFRICAN MUSEUM

Book 1908-1910: 835). The stone referred to is one found in 1872 by the son of

a Mr Chevalier in a cave in the Eastern Head at Knysna, about two feet (0,6 m)

below the surface (J. Rudner 1971: 54; see also Péringuey 1911: 162-163) and

which is in the Museum’s collections (SAM—AA8386— Fig. 1A).

Péringuey twice again referred to this stone in letters to Henkel the follow-

ing year. On 12 May 1910 he wrote ‘I have had a good deal of information

regarding the contents of some of these caves in the Knysna—Plettenberg Bay

coastal region and I thought we had the only painting, a monochrome, found

there, until you told me of Witcher’s [sic] discovery’ (SAM Letter Book

1910-1911: 136). On 11 October he wrote ‘The painting has been carefully pre-

served—it is a small fragment and had the painted stones of Coldstream not

been found, I would have been left in doubt as to it having been detached from

the sides or roof of the cave’ (SAM Letter Book 1910-1911: 527).

C. J. Whitcher was the owner of a sawmill near the little village of Cold-

stream (Fig. 2) and of extensive tracts of indigenous forest in the surrounding

area. He excavated in the shelter at the mouth of the Lottering River, one of

several shelters on his property, possibly in 1908 but certainly early in 1909,

making the discoveries referred to in Henkel’s correspondence with Péringuey.

Péringuey (1911: 154-158) published excerpts from Henkel’s correspon-

dence in his seminal monograph on the Stone Ages of South Africa, in which

Whitcher’s excavations are described. Initially, Henkel emphasized ‘You will

observe from my account that there was no direct evidence to connect the stones

decorated with drawings with skeletons, although in the first instance I was led

to believe that this was so’ (Péringuey 1911: 158). However, later in the same

extract, Henkel confirmed from personal observation on 14 July 1909 that from

one to three stones, none of them painted, were found overlying skeletons

excavated during his visit.

Péringuey was extremely anxious to obtain for the Museum the artefacts

and skeletons excavated by Whitcher, as well as to ensure that any future exca-

vations were carried out systematically. Henkel negotiated with Whitcher on

Péringuey’s behalf for the purchase of the material and on 7 July 1910 Péringuey

wrote to Henkel, acknowledging receipt of it and commenting “The ostrich egg-

shell beads are interesting because found on the child, and the cockle necklace is

next to the painted stone—the larger one—the interesting part of the collec-

tion’ (SAM Letter Book 1910-1911: 271).

Péringuey’s reference to ‘the larger one’ of the painted stones, implying as

it does that two were received, calls for comment. In a paper on painted ‘burial

stones’ from the Cape, J. Rudner (1971: 56) described four painted stones from

the Coldstream shelter. Rudner’s C1-—3 are said to have been found by Whitcher

and to have the accession number SAM-[AA]8387, whereas his C4, accession

number SAM-[AA]6008, is the painted stone that is the subject of this paper.

According to Rudner, ‘C1 is a lower grinding stone with four human figures

painted on a red background . . . C2 consists of a rough rock fragment with a

black painting on the upper part carrying a long, probably cylindrical quiver and

AN INVESTIGATION OF THE ‘COLDSTREAM STONE’ 191

a bow . . . C3 originally contained four black figures of animals, but these have

now disappeared’.

The Archaeology Department’s accession register lists under SAM-

AA8387 only two painted stones that, from Rudner’s descriptions, are his Cl

and C2. According to Péringuey (1911: 163), ‘Mr Henkel reports that the third

one became obliterated as soon as exposed to light’. It is thus possible that

Péringuey actually received three stones, but omitted to mention the third,

Rudner’s C3, from which the painting had vanished and which was accordingly

not accessioned as a painted stone.

The matter of the acquisition of Rudner’s Cl (Fig. 1D) also calls for

comment. On 12 April 1912 Péringuey wrote to Whitcher ‘I should very much

like you to loan me the tracing taken by Fourcade of the dancing women, on the

stone found in the cave. Our copy has greatly faded and I want to re-touch it’

(SAM file M—18—d). This seems a strange request if the Museum had acquired

the stone itself in 1910. Moreover, Péringuey (1911, fig. 199) had published an

artist’s impression of the stone, coloured lithographs of which are still in the

Museum’s possession. However, Péringuey (1911: 163) observed of the painted

stones from Coldstream “They are fading so rapidly that we had a great difficulty

in reproducing them’. He may therefore have considered that Fourcade’s copy

would provide a more accurate representation than could be obtained from the

stone itself.

Whatever the case, it is evident from the foregoing that, up to the beginning

of 1911, the only painted stones known to exist in South Africa were considered

to be either monochrome: the Knysna Heads stone (Fig. 1A), or bichrome: the

‘dancing women’ stone (Fig. 1D) and the smallest of the three stones from

the Coldstream shelter (Fig. 1C). It should be pointed out that, although—as

mentioned above—Peéringuey considered the Knysna Heads stone to be a

monochrome, the black figures are painted on a red ochre background, as are

the ‘dancing women’.

Where Rudner’s C2, the smallest of the Coldstream stones, is concerned,

microscopic examination of what Rudner (1971: 56) described as ‘a white mark

... that ...may be part of the picture’ has shown that it is in fact white

pigment, evidently depicting a bag, or possibly two, slung over the figure’s

shoulder. Samples of the pigments on this stone were included in the analyses

discussed below.

Péringuey (1911: 163) did not at first consider that the ‘hook-heads’ of the

figures in the ‘dancing women’ painting resulted from the loss of the white

pigment used to depict the faces, a phenomenon now well known to occur in

paintings on rock-shelter walls (Yates et al. 1985: 70; W.J.J.v.R. pers. obs.). He

suggested that it might be a totem, but thought there was no doubt that it was ‘a

distinctive badge’. However, in the final chapter of his monograph, which deals

with the Coldstream shelter and appears to have been written after the main

body of his text, Péringuey (1911: 209), in describing the ‘Coldstream Stone’,

commented ‘The face is painted white and the crescent-shaped process...

192 ANNALS OF THE SOUTH AFRICAN MUSEUM

which I took at first to be possibly the mark of a clan—stands for the hair and is

black’. The conclusion is, therefore, that on the basis of the information then

available, there was no reason to suppose that anything other than monochrome

or bichrome painted stones would be found.

DRURY’S EXCAVATION

On 16 February 1911, on Péringuey’s instructions, James Drury, the

Museum’s taxidermist (Fig. 3), started excavations in the shelter in which

Whitcher had made his finds. The following are excerpts from Drury’s field

notes (SAM file M-—18—d). In the interests of authenticity, the only alterations

that have been made to Drury’s text are the omission of certain passages that

are not directly relevant and the separation of the daily information into para-

graphs. The days of the week, written in the margin of Drury’s notes, have been

incorporated into the text.

‘Mr Whitcher had dug out a part of the cave marked in sketch, he had got

to a depth of 10 feet, so I dug another 2 feet found remains of Baby skeleton

very badly preserved, and as I don’t think skeletons or paintings can be of any

use at that depth I gave up working and made a fresh start, at the back centre of

cave as in sketch. I found the top layers sandy for a few inches then shells stones

and refuse to a depth of just on 3 feet, then a layer of limy ground varying from

an inch to 3 inches thick and so on in alternating layers of ashes earth lime and

shells to a depth of about 4 feet 6 in. then the ground is brownish earthy and

rather damp... .

Thursday. I started a section next to where Mr Whitcher had worked and

am now working down to the same level as first section about 11 feet

deep... . At a depth of 4 feet I found my first skeleton, no. 2. . . their was no

flat stones on top or side of him . . . this skeleton is very rotten (as I find all are)

and I don’t think it will be any good to go too deep. Next skeleton was a baby

but as before too far gone, this was close to the above skeleton, then I found

another just a little deeper . . . lying on right side with a stone on top this time

but their was no trace of Paintings. All through we found lots of colouring sub-

stances but no trace of painting. I got charcoal, Red Ochre also Yellow Ochre

but so far no white.

Friday. I started another section down and when nearly the end of the layer

I found the second full skeleton and the most interesting find so far. I was

having layers carefully removed when I saw back part of skull protruding out of

ground so started to carefully work earth away all round with trowels and hands

and in so doing found a large flat stone lying on shoulder, so with great care

removed the earth from on top, and thus exposed was the finest Painting I have

so far seen anywhere, and with great care got it cleaned of all dirt adhering and

so removed to Mr Whitcher’s house where I finished cleaning, their is three

figures in colours and realy marvellous it is. The position of stone was flat on

shoulder with feet end of figures next to the jaw of skeleton on painting side

AN INVESTIGATION OF THE ‘COLDSTREAM STONE? 193

Fig. 3. James Drury excavating in the Coldstream shelter, February 1911.

(Photograph: S.A. Museum archives.)

194 ANNALS OF THE SOUTH AFRICAN MUSEUM

up... there was a flat stone lying on hips of skeleton also but there were no

tracings of Paintings . . . close by him I| found for the first time two pieces of

white clay which now gives all the colours found on painting, this clay was in a

hole in rock just at back of where the painting was found... .’

Drury continued excavating until 28 February, removing the bulk of the

deposits and finding a total of 28 skeletons. Some of the burials had stones

associated with them, but none of them was painted.

According to J. Rudner (1971: 56) the skeleton with the painted stone was

found at a depth of 6 feet (approx. 1,8 m). This is not stated in Drury’s notes

but the three sketches of burials that accompany his notes are, according to the

captions, of the first three skeletons, that noted as having been found at a depth

of 6 feet being identified as ‘2nd skeleton’ and ‘no. 10’. It will be seen above that

on the Friday, Drury mentioned finding ‘the second full skeleton’ and Rudner’s

identification may thus be correct. None of the three sketches contains infor-

mation about the placement of stones apart from a note that the first skeleton

had ‘no big stones on top’ (SAM file M—18-d).

SUBSEQUENT HISTORY

As already mentioned, Péringuey (1911: 209, pl. 27) described the ‘Cold-

stream Stone’ in his monograph but illustrated only the other two painted stones

from the site and the Knysna Heads stone. This tends to confirm that his chapter

on the Coldstream shelter was a late addition to his manuscript and that he did

not have time to have a plate prepared from an artist’s impression, as was the

case with the ‘dancing women’ stone. One was prepared at some stage and sent

to Germany for the preparation of coloured lithographs, but due to the exigen-

cies of the First World War these did not reach South Africa until after 1920

(R. Summers pers. comm. 1985). It was left to Haughton (1926) to publish the

first illustration of the ‘Coldstream Stone’, after Péringuey’s death in 1924.

Burkitt (1928, fig. 20) illustrated only the ‘dancing women’. Goodwin (1929:

209, pl. 42), in discussing painted stones, merely remarked that ‘Coldstream

shows three female figures and a child’ and illustrated a monochrome from the

collections of the Port Elizabeth Museum (PEM 603). In a later monograph

Goodwin (1946: 115-16) mentioned the coastal caves of the southern Cape and

the ‘amazing efflorescence of culture’ of the area, which included painted ‘grave-

stones’; and elsewhere in the same work (Goodwin 1946: 127) he discussed

Péringuey’s (1911) monograph. As mentioned above, the latter publication con-

tains a description of the ‘Coldstream Stone’ though not an illustration of it.

Goodwin must therefore have known of the stone’s existence; and, as he was at

that time honorary curator of the Museum’s archaeological collections, he could

accordingly be expected to have known of the stone’s presence in the Museum’s

collections. Moreover, he sent a copy of the coloured lithograph to J. D. Clark

(Clark pers. comm. to J. Deacon 1986). It seems, however, that Goodwin never

published a reference to this unique artefact in the years remaining before his

AN INVESTIGATION OF THE ‘COLDSTREAM STONE’ 195

death in 1959. Goodwin’s apparent failure to refer to the ‘Coldstream Stone’ in

any of his prolific writings is a puzzling aspect of this case for which there is now

no ready explanation. On the other hand, E. M. Shaw, until recently Head of

the Ethnography Department, who started at the Museum in 1933, and in whose

charge the archaeological collections were until the appointment of an archae-

ologist to the Museum’s staff in 1967, has said (pers. comm. 1986) that it was

many years before she became aware of the existence of the ‘Coldstream Stone’;

and it may be that it lay forgotten in the Museum’s strong-room. (It was only

accessioned in 1956.)

The ‘Coldstream Stone’, remarkable though it is, appears to have been

ignored for more than thirty years after Haughton’s publication, until Clark

(1959: 214, pl. 15b) mentioned ‘the magnificent painted stones of Coldstream’

and included an illustration of the stone under discussion. Since then it has been

described and/or illustrated by Woodhouse (1968, 1969, 1987, 1988 in Coertze

ed. 1988 (unpaginated)), Lee & Woodhouse (1970), Rudner & Rudner (1970),

J. Rudner (1971), Summers (1975), I. Rudner (1982), Lewis-Williams (1984)

and Willcox (1984). None of these authors commented on the fact that this is the

only known polychrome in the category of painted stones, and only J. Rudner

(1971: 56) and Lewis-Williams (1984: 241) commented on the remarkable state

of preservation of the painting.

PHYSICAL ANALYSES OF THE PIGMENTS

INTRODUCTION

The rarity and consequent importance of the painted stones impose severe

restrictions on the size of the sample available for analysis. The sampling tech-

nique must be as little destructive as possible, which means that only small

quantities of the paint may be removed. The paint is nowhere thick, with an

estimated maximum of 100 u; and this further affects the available size of the

sample. Energy-dispersive X-ray (EDX) micro-analysis is thus the only tech-

nique available for this purpose.

Samples were obtained by placing a 5x 5 mm piece of silver mylar tape

(Scotch 3M 850 PAU333310) with the adhesive face down on to the painted area

and rubbing the back of the tape with a small, blunt object. The effect of this

sampling technique was minimal and the places from which the samples were

removed cannot be detected with the naked eye. The pieces of tape were then

glued to Cambridge 100 stubs and carbon coated, to an interference colour blue,

in a Balzer’s high-vacuum evaporator.

The paint and raw pigment samples were analysed using a KEVEX 7000

EDX system, interfaced with a Cambridge $180 scanning electron microscope.

Standardized operating conditions were used for all the analyses. Conditions

included a calibrated 15 keV probe, 38° take-off angle and a 20 per cent

amplifier dead time. The electron optic parameters gave a probe current of

1,2 x 10-° amps. All the analyses were carried out using the same electron optic

196 ANNALS OF THE SOUTH AFRICAN MUSEUM

parameters so that spectra would be comparable, even though absolute values

were not obtained.

EDX micro-analysis is capable of detecting any element having an atomic

number greater than 11 (Na) and in practice the detection limit is about 0,1

weight per cent (wt %) in the analysed volume. The volume analysed depends

on sample density, but is in the order of 0,08 mm? for a sample of density

8 g/mm? and using a 15 keV probe and a scanning electron microscope magnifi-

cation of 1 000 X on the visual screen.

By measuring the intensity of X-ray emission for each element and

comparing the experimental with the theoretical X-ray yield, the concentration

of constituent elements can be determined. Unfortunately, the relative errors

in this method of analysis can be high, particularly if the sample contains unde-

tectable elements such as oxygen and carbon, and is not homogeneous and

flat. The more satisfactory strategy involves direct comparisons of X-ray yield

with similar accurate standards. The difficulties in this method are obtaining

suitable standards, polishing the samples, and particle selection based on

morphology.

For each sample the secondary electron image was observed and analyses

carried out, initially using area mode (the probe scanned in a raster over an area

of about 30 x 30 u) on visually selected differences. Nine additional spot or area

acquisitions (the probe stationary in one position) were undertaken on selected,

morphologically similar particles or areas in order to establish the nature of the

sample and to determine the average concentration of detectable elements. In

selecting the area or spot for acquisition, the probe was carefully positioned to

assist In minimizing topographical, refluorescence and interaction volume errors.

In this connection, it must be pointed out that, although the samples analysed

were small, because they were almost without exception not homogeneous,

probes that, for example, included a grain of sand would yield results very

different from that of pure ochre. The raw pigments may have been ground on a

quartzite grindstone, using a muller of the same material, which would introduce

foreign material. The pigment was mixed with some sort of binder and probably

liquefied with additional fluid, both of which would introduce more foreign

elements.

Incontrovertible factual information regarding the nature of the binders

used in rock paintings is not available, although much has been written about

these, and a number of experiments carried out by various researchers.

I. Rudner carried out extensive documentary research and fieldwork in regard to

Khoisan paints and pigments and their relationship to rock paintings, and con-

cluded (I. Rudner 1982: 264) that there is virtually no primary information

regarding the nature of the binders used, and that very little of what is available

can be considered as reliable. She was particularly sceptical of the claim by Den-

ninger (1971) that a number of binders had been identified by means of paper

chromatographic analysis of the constituent amino-acids (I. Rudner 1982:

33-34).

AN INVESTIGATION OF THE ‘COLDSTREAM STONE’ 197

The results of the five analyses of each sample that were most similar were

averaged and are presented in graphic form in Figures 4-10 as weight per cent

(wt %) of the elements detected in the analysed volume, normalized to 100 per

cent. Since the samples are not homogeneous and averages were used, the results

characterize the samples rather than provide precise, absolute determinations of

their constituents.

When studying the results of the EDX analyses given below, it must be

borne in mind that none of the samples can be considered as chemically pure.

They have all been subjected to various taphonomic processes: adsorption,

absorption, and leaching of material, in addition to the other forms of contami-

nation mentioned above. It should also be remembered that silicon (Si) and

aluminium (Al) are the two most common elements in the earth’s upper crust

(hence its name, sial). For the purposes of these analyses, the presence of these

elements should be considered as ‘noise’ and, for the most part, irrelevant.

Included in the small collection of non-skeletal material Drury retained

from his excavation (SAM-—AA1378, AA1379) are a few pieces of ochre as well

as one of the pieces of ‘white clay’ that he recorded (see above) as having found

in the deposits. It was therefore decided to analyse samples of these as well as of

the paint on the ‘Coldstream Stone’ and, where necessary, to compare the

results with those of other paints and pigments. Because of a lack of concord-

ance between the results of the analyses of the various white and black

pigments in the earlier series of analyses, samples of marine shell and burnt

animal bone were also analysed.

There is too little black paint left on the ‘dancing women’ stone (Fig. 1D) to

allow for analysis, and the red ochre background is also a complicating factor.

Samples of the black and white paints from the smallest of the three stones from

the site (Fig. 1C) were, however, included in the analyses.

RED PAINTS AND PIGMENTS

Four samples of red paint from the ‘Coldstream Stone’ were analysed, and

two of unprocessed ochre from the excavation that visually matched the light red

and dark red paints most closely. The results are shown in histogram form in

Figures 4—6.

The light red paint from the front (left) figure (Fig. 4A) has less iron (Fe)

and lower frequencies of the minor elements (S—Ti) than the dark red paint

from the central figure (Fig. 4B); otherwise these two samples are broadly

similar.

Both the light red ochre from the excavation (Fig. 5A) and the dark red

(Fig. 5B) have very little iron by comparison with the frequencies in the paints

(Figs 4A—-B) and are generally dissimilar to the paints in most respects. The

light red ochre lacks five of the minor elements present in the dark red. These

minor elements in the latter may be the cause of its darker colour.

Figure 6A is the analysis of the light red paint from the front leg of the last

(right) figure and Figure 6B that from the back leg. It is immediately evident

198 ANNALS OF THE SOUTH AFRICAN MUSEUM

Oo.

a -

Na Mg Al Si P S Cl K Ca Ti Mn Fe Co Ni Cu Ba

ELEMENT

Oo.

| cd

Na Mg Al Si P S CI K Ca Ti Mn Fe Co Ni Cu Ba

ELEMENT

Fig. 4. Histograms of amounts (wt%, normalized to 100%) of elements detected in paint

samples. A. Light red paint from the ‘Coldstream Stone’. B. Dark red paint from the

‘Coldstream Stone’.

AN INVESTIGATION OF THE ‘COLDSTREAM STONE’ 199

WwW

oO.

Na Mg Al Si P S Cl K Ca Ti Mn Fe Co Ni Cu Ba

ELEMENT

O 40

WwW

Oo.

- 30

Ww GST,

= 20

10 mt

WE, ot

aaa an le a —-——— 7A ZI

Na Mg Al Si PS Cl K Ca Ti Mn Fe Co Ni Cu Ba

ELEMENT

Fig. 5. Histograms of amounts (wt %, normalized to 100%) of elements detected in pigment

samples. A. Dark red ochre from the excavation. B. Light red ochre from the excavation.

200 ANNALS OF THE SOUTH AFRICAN MUSEUM

oO.

Na Mg Al Si P S CI K Ca Ti Mn Fe Co Ni Cu Ba

ELEMENT

Oo.

pis

Na Mgr All Si PS) Cl 1K Ca Ti Mn Sees Co Ni MCusEa

ELEMENT

Fig. 6. Histograms of amounts (wt%, normalized to 100%) of elements detected in paint

samples. A. Light red paint from the front leg of the last (right) figure on the ‘Coldstream

Stone’. B. Light red paint from the back leg of the same figure.

AN INVESTIGATION OF THE ‘COLDSTREAM STONE’ 201

that the two samples differ considerably from each other and from those of the

other two figures. The paint from the front leg is similar to that of the light red

ochre from the excavation (Fig. 5A), although this contains only about half as

much iron. The presence of a relatively high frequency of calcium (Ca) in the

sample from the back leg is not easy to explain. Unlike the front leg, there is no

white paint on the back leg, which is, however, outlined in black on its front

(Fig. 1). A possible source of the black paint used on the ‘Coldstream Stone’,

and thus of the calcium in this sample, is given below.

WHITE PAINTS AND PIGMENTS

Samples of white paint from the face of the central figure on the ‘Cold-

stream Stone’ and the white bag(?) carried by the figure on the smallest painted

stone from the site were analysed, as was a sample of the white ‘clay’ Drury

reported as having found in the excavation. The presence of a high frequency of

calcium in the samples could not be explained, particularly as one of the samples

analysed for comparative purposes (but not included here), from a mural paint-

ing in the Transkei, only contained a minimal amount of calcium (approx. 6%).

A sample of calcined marine shell from an excavation in the southern Cape was

therefore also analysed. The results of these analyses are shown in Figures 7-8.

The paint from the ‘Coldstream Stone’ (Fig. 7A) and that from the smallest

stone (Fig. 7B) differ only in the amount of calcium present and in the presence

or absence of various minor elements. The white ‘clay’ (Fig. 8A) has consider-

ably less calcium than the paint samples, partly because of the increased

presence of other elements. Examination of this material under a standard

binocular microscope indicated that it is not clay but an agglomerate of calcined

material that includes marine shell, sand and other unidentified material.

Because of uncertainty as to the precise nature of the white paint, a sample

of calcined marine shell, brown mussel Perna perna, without its periostracum,

was also analysed (Fig. 8B). Despite differences caused by the presence or

absence of small amounts of minor elements in the two paint samples, the result

of the shell analysis strongly suggests that shell was the pigment used for the

white paint on both stones. It also tends to confirm that the white ‘clay’ is prob-

ably a lump of calcined shell with other admixtures. Another possibility, egg-

shell, which is also rich in calcium, was considered later but not tested.

BLACK PAINTS AND PIGMENT

Samples of black paint from the quiver or bag held by the central figure on

the ‘Coldstream Stone’ and from the figure on the smallest of the stones from

the site were analysed. The results of these were compared with that of black

paint from a mural rock painting in the Cedarberg Mountains of the western

Cape. As will be seen below, this bore no resemblance to the paint on the two

stones. On the basis of the discovery that the white paint on these stones had

probably been made from calcined shell, it was decided to analyse a fragment of

202 ANNALS OF THE SOUTH AFRICAN MUSEUM

oO.

Na. Mg Al Si P S* Cl K Ca Ti Mn Fe Co NiCusBa

ELEMENT

oO.

Na a tes Al K Ca Ti Mn Ti Ma Fe. ce) Noe Co Ni Cu Ba

ELEMENT

Fig. 7. A. Histograms of amounts (wt %, normalized to 100%) of elements detected in paint

samples. White paint from the ‘Coldstream Stone’. B. White paint from the smallest of the

painted stones from the Coldstream shelter.

AN INVESTIGATION OF THE ‘COLDSTREAM STONP’ 203

WEIGHT PER CENT

LZ,

0 82 Vea ETE, A SEC EAN TERY

Nae Mg Alsi PS (Cl K Ga Ti Mn Fe Co Ni Cu Ba

ELEMENT

WEIGHT PER CENT

NaiViqurAle Site Penton Clenheu Can til Ninpte Co mNi- Cus Ba

ELEMENT

Fig. 8. Histograms of amounts (wt %, normalized to 100%) of elements detected in pigment

samples. A. White ‘clay’ from the excavation. B. Calcined marine shell.

204 ANNALS OF THE SOUTH AFRICAN MUSEUM

O 40

Oo.

FE 30

Na’ Mg) VAll "Si PS) Cl 3K Cas Ti Mn Fem Com NimCumea

ELEMENT

Na Mg “Al” Si PS) Cl 4K Car it Mn kesiCom NiniceusBa

ELEMENT

Fig. 9. Histograms of amounts (wt%, normalized to 100%) of elements detected in paint

samples. A. Black paint from the ‘quiver’ of the central figure on the “Coldstream Stone’.

B. Black paint from the smallest of the painted stones from the Coldstream shelter.

AN INVESTIGATION OF THE ‘COLDSTREAM STONE’ 205

Oo.

Na Mg Al Si P S Cl K Ca Ti Mn Fe Co Ni Cu Ba

ELEMENT

| ome

O 40

= 30

Na Mg Al Si “P= S Cl K Ca Ti Mn Fe Co Ni Cu Ba

ELEMENT

Fig. 10. Histograms of amounts (wt %, normalized to 100%) of elements detected in paint and

pigment samples. A. Black paint from a rock painting in the Cedarberg. B. Burnt mammal

bone.

206 ANNALS OF THE SOUTH AFRICAN MUSEUM

burnt mammal bone, also from an excavated site in the southern Cape. The

results of these analyses are given in Figures 9-10.

The large amount of calcium in the paint from the ‘Coldstream Stone’

(Fig. 9A) and from the smallest of the painted stones (Fig. 9B) was surprising,

especially when these were compared with the paint from the Cedarberg rock

painting (Fig. 10A), which consists mostly of manganese (Mn), a substance

often found in nodular form in archaeological deposits, and common in the

sandstones of the Table Mountain Group. Since it was known that carbon, an

organic element that is used to prepare samples for electron microscopy, would

not register in the EDX analyses another source of the black pigment was

sought: burnt bone. Although the result of this analysis (Fig. 10D) shows differ-

ences from the paint on the two stones from the Coldstream shelter in the

presence or absence of a number of minor elements, and differences in the fre-

quency of others, the burnt bone sample is more like the two Coldstream paint

samples than any of the three are like the Cedarberg paint sample. Although

these analyses have not demonstrated conclusively that the black paint was

made from burnt bone, they do show that the pigment used was not derived

from a manganese-based compound.

DISCUSSION

As mentioned in the introduction to this section, EDX analysis is at present

the only feasible method of analysing very small samples of paint from rock

paintings. The method has its limitations in that it cannot identify all the con-

stituents of the paints and, because of their lack of homogeneity, cannot identify

them precisely.

However, given these limitations, the analyses have shown that it is possible

to characterize paints and raw pigments to a degree that enables it to be deter-

mined whether any two samples are alike or not. Moreover, as was shown in the

cases of the white and black paints, it is possible to identify possible, even prob-

able, sources of the pigments from which the paints were made. The analyses

have therefore proved useful in the investigation of the ‘Coldstream Stone’.

THE STYLE OF THE PAINTINGS

The figures on the ‘dancing women’ stone (Fig. 1D) are now only barely

visible, but examination of the coloured lithograph in the Museum’s possession

(see Péringuey 1911, fig. 199; Burkitt 1928, fig. 20, etc.) shows that, in general,

they differ from the figure on the smallest stone only in having ‘hook-heads’ and

in being more robust. This latter aspect may be a consequence of the different

sizes of the paintings, though this is not true of the larger ‘Coldstream Stone’. In

general, however, figures of women in rock paintings are more amply propor-

tioned than those of men (W.J.J.v.R. pers. obs.).

The ‘hook-heads’ of the ‘dancing women’ provide a stylistic link with the

figures on the ‘Coldstream Stone’: were the men’s white faces to disappear, they

AN INVESTIGATION OF THE ‘COLDSTREAM STONE’ 207

would be left with ‘hook-heads’. That apart, the stylistic differences between the

paintings on the three stones from the Coldstream shelter are marked. The

‘Coldstream Stone’ appears to have the most elaborate style of the three, but it

should be borne in mind that the painting of the ‘dancing women’ is now only a

faint vestige of what it was originally.

The style of the painting on the two smaller stones is closer to that of the

figures in the mural paintings in Whitcher’s Cave and the small shelter above it

(Woodhouse 1966) than to that of the ‘Coldstream Stone’. However, the

Whitcher’s Cave paintings are in red only and do not have ‘hook-heads’ indica-

tive of the loss of the white paint used for the faces on the ‘Coldstream Stone’

and, probably, the ‘dancing women’ stone. Whitcher’s Cave is in the southern

foothills of the Tsitsikamma Mountains, some 10 km north-east of the Cold-

stream shelter (Fig. 2), and is the closest known site with mural rock paintings.

Relatively little has been published about the rock art of the southern Cape,

but L. Abel, who has studied the art of the area and has seen an illustration of

the ‘Coldstream Stone’, is of the opinion (pers. comm. 1985) that the style of the

painting on the stone is similar to that of mural rock paintings in the Langkloof

and in the Willowmore district, which are to the north of the Tsitsikamma

range. Rudner & Rudner (1970: 126) found the figures on the stone reminiscent

of some of those in the mural paintings on Kriedouwkrantz Hill, which is in the

Clanwilliam district of the western Cape Province.

Since there is no information as to the stratigraphic provenance of the

painted stones from the Coldstream shelter, even their relative chronology

cannot be established. The most that can be said, then, is that their styles fall

within the range of painting styles of the rock art of the area.

The last figure on the ‘Coldstream Stone’ differs from the other two figures

in some respects. In these, the body and limbs were sketched in white before the

red paint was applied, and the lower part of the bodies as well as the front of the

legs were outlined in black. In the last figure, only the front leg and the lower

part of the body were first sketched in white and the black outlining was

extended to the back of the trunk and to the back of the rear leg. Parts of this

figure, and particularly the rear leg, have the appearance of having been

retouched, a suggestion that is reinforced by the differences indicated above in

the element composition of the paint on the two legs.

DISCUSSION AND CONCLUSIONS

Discussing the ‘Coldstream Stone’, Lewis-Williams (1984: 241) commented

‘The extraordinarily good preservation of the painting on this stone, coming as it

does from a rather damp cave deposit, has led some workers to question its

authenticity. Two possibilities exist: Either the stone is a complete forgery, or it

has been considerably touched up. The second of these possibilities may be

correct because it seems unlikely a forger in 1911, when the stone was found,

208 ANNALS OF THE SOUTH AFRICAN MUSEUM

would have known about some of the details in the painting and because, if it

had been touched up, the workers of that time may not have considered it

necessary to record their restoration’. The details Lewis-Williams referred to are

those such as the red lines across the faces, which he interpreted as stylized

depictions of nasal haemorrhage induced during trance performance, rather than

the purely technical details such as the style and manner in which the painting

was executed.

The possibility that the painting was touched up subsequent to its discovery

seems unlikely. The present condition of the paint is not uniform, most of the

black, in particular, having all but disappeared. If any touching up was done, it

seems most likely to have been of the rear leg and lower part of the body of the

last figure; but if this was so, it was a clumsy attempt at restoration to have out-

lined the back of the trunk and leg in black when the back of the front leg is

outlined in white, as are the legs of the other figures. Moreover, had it been the

practice to restore such paintings, this would surely have been done to the

‘dancing women’ stone which, as mentioned above, was in a very poor condition

as early as 1912. At some time, attempts were made to preserve some of the

Museum’s other painted stones, notably those from the caves on the Robberg

Peninsula at Plettenberg Bay (SAM-—AA2822 to AA2828), by covering them

with clear varnish. This has now discoloured to the extent that some of the

paintings are almost obliterated, and it is fortunate that the painted stones from

Coldstream, whatever their present condition, were not subjected to this

treatment.

Discussing the painted ‘gravestones’ of the southern Cape, Battiss (1948:

93) commented ‘It is difficult to examine the paintings on these gravestones for

SO many are in a very poor State of preservation. In a letter to me Mr A. J. H.

Goodwin writes: ‘““The many gravestones from Robberg have almost been

destroyed. This was owing to the way in which the paint was forced out of the

stone by the efflorescence of salt crystals and so formed a fine impalpable

powder— just like a pastel”’’. The Robberg Peninsula is some 35 km west of the

Coldstream shelter and in the same general environment, although the Cold-

stream shelter is higher above sea-level than some of the Robberg caves from

which the painted stones were recovered, and better protected by vegetation

from wind-blown salt spray. The shielding vegetation does, however, enhance

the dampness of the shelter’s deposits (M.L.W. pers. obs.).

There is no indication on any of the three painted stones from Coldstream

of any crystalline efflorescence, nor any indication of the paint having been

forced out of the stone by such action. On the contrary, most of the paint is

retained in the pores and other surface irregularities on the stones. The same is

true of the painted stones from Klasies River Mouth Cave 5 (Fig. 1B, E; Singer

& Wymer 1982, figs 48—49) as well as the other painted stones in the Museum’s

collections. The painted stone from a rock-shelter in the Caledon River Kloof,

on the farm Danielskraal in the Calitzdorp district (Fig. 1G), has a calcareous

accretion on its sides and base, but it is the only one of the painted stones in the

AN INVESTIGATION OF THE ‘COLDSTREAM STONE’ 209

Museum’s collections that shows any such damage, perhaps excluding those

from Robberg already mentioned, which were subjected to preservative treat-

ment. The Danielskraal painted stone was one of two found on the floor of the

shelter, the deposits having been washed away (J. Rudner 1971: 57).

The differential preservation of the painting on the three painted stones

from the Coldstream shelter may be explained partly by differences in the

nature of the pigments and/or binders used, partly by the nature of the rock

surface on to which they were painted, and partly by differences in the way in

which the artefacts were handled during and after excavation.

The nature of the binders clearly has some bearing on the way in which the

paint adheres to its stone base. In the absence of any clear information in this

regard, however, and particularly with regard to the painted stones from the

Coldstream shelter, it is not possible to enter into any discussion on this subject.

It will be seen that, in the examples illustrated in Figure 1, red is the colour

that is generally best preserved, followed by white, and with black least well pre-

served, although in the case of the ‘dancing women’ stone the white has

completely disappeared, leaving only ‘hook-heads’. In the case of mural rock

paintings, the order in which paints are best preserved is generally red, yellow,

black and white (W.J.J.v.R. pers. obs.; see also Yates et al. 1985: 70). This

makes it the more surprising that white is better preserved than black on the

‘Coldstream Stone’. This may have to do with the use of different types of pig-

ments in different areas.

The ‘dancing women’ are painted on a lower grindstone, the surface of

which is accordingly smoother than that of the ‘Coldstream Stone’, although this

is a small water-worn boulder. The smallest of the three stones is an irregular

piece of cave rock, quartzite like the others, but with a more uneven surface.

The different surface textures almost certainly play a part in the retention of the

paint on its base.

From the information Henkel gave Péringuey, the ‘dancing women’ stone

may have been the first found by Whitcher who, being unaware of the existence

of such artefacts, may not have handled it as carefully as he did the other two

painted stones, although the third was left exposed to the weather. Drury began

his excavation in the knowledge that painted stones might be found, and he

would accordingly have treated any stones he found very carefully, as his field

notes in fact reveal he did when the ‘Coldstream Stone’ was found. Figure 3

shows that the stones and other material from his excavation were laid on

sacking.

One of the aspects of Drury’s field notes that seemed suspicious was his

mention, the day before making his great discovery, of finding red, yellow and

black pigments in the deposits ‘but so far no white’. In this, he seems almost to

have anticipated the discovery of a polychrome painted stone. It must be stated

again, however, that the smallest of the three painted stones obtained from

Whitcher is a black and white bichrome, and was already in the Museum’s pos-

session. As discussed earlier, Drury had no reason to suspect the existence of

210 ANNALS OF THE SOUTH AFRICAN MUSEUM

polychrome painted stones, the only three painted stones known before his dis-

covery being monochromes or bichromes. Moreover, yellow ochre, one of the

pigments he mentioned, appears not to have been used in the painting. He

would, on the other hand, have known that polychrome mural paintings existed,

as indicated by the comment in his field notes given above, that this was ‘the

finest Painting I have so far seen anywhere’. The Museum’s earliest other acqui-

sition of polychrome paintings, two pieces of a large slab from a mural painting

in a cave on the farm Zamenkomst in the Maclear district of the eastern Cape

(SAM-AA1617), was only obtained in 1912 (SAM Antiquities Register: 93).

If Drury were the perpetrator of a forgery, the logistics of this act, though

not insuperable, would have been considerable. The stone on which the painting

was done measures 300 x 240 x 90 mm and weighs 8,5 kg. Had Drury prepared

the stone in Cape Town, he would have had to take it with him on the train

journey to Assegaibosch, and thence by cart on the close on 70 km journey to

Coldstream. He stayed with Whitcher at the latter’s home, ‘Oakhurst’, which is

about 5 km from the shelter (Fig. 2), and even though Whitcher gave him the

use of a cart, its closest access to the shelter was about a mile (1,6 km) from the

site (SAM file M—18—D). Drury would thus have had to transport the stone to

Coldstream, get it into and out of Whitcher’s house and then to the site, all

unnoticed by Whitcher, who probably accompanied him to the shelter on at

least the first day.

An alternative possibility is that Drury painted the stone at the site. He

could have obtained the stone from the site: as mentioned above, he found a

number of unpainted stones in the deposit (see Fig. 3, behind Drury); or he

could have brought it up from the beach, some 20 m below. He could have used

some of the pigments he recorded as having found in the deposit, although this

is somewhat negated by his record that it was only after he had found the

painted stone that he found two pieces of ‘white clay’ in a niche in the shelter

wall behind the burial. In this case, Drury would almost certainly have had to

bring a drawing, or drawings, with him, that he could use to copy on to the

stone. He employed four labourers, who would have been witnesses to this

action, as well as to his ‘salting’ of the deposit, or to his pretence that he had

found the stone on the shoulder of the skeleton, as he reported.

While neither of these scenarios is beyond the bounds of possibility, both

tend to seem far-fetched, and to credit Drury with remarkable deviousness. The

strongest argument that can be advanced against Drury’s having done the paint-

ings is the evidence provided by the analyses of the paints and pigments. These

indicate, with a high degree of probability, that the white paint was made from

ground shell and the black from burnt bone. Drury thought that the white

material he found at the back of the cave after he had found the painting was

clay; and he said in his field notes that charcoal was the black colouring

material. It seems highly unlikely that he would have considered that shell or

bone could or would have been used as pigments. That there is a close match

between the light red ochre from the site and the paint on the front leg of the

AN INVESTIGATION OF THE ‘COLDSTREAM STONE’ PAS

last figure but not on the back leg, which is the one that might appear to have

been retouched, seems to provide further support for the argument that Drury

was not involved in either the painting or the retouching. Since the red colour of

the first and last figures is visually similar, and different from that of the central

figure, it seems more logical to have used the same pigment for both. The

retouching could well have been done before the stone was buried.

Apart from the fact that Drury made numerous plaster casts of rock

engravings, some from as far afield as Namibia, nothing is known of his interest

in, or knowledge of, rock art. He joined the Museum in 1902 but, although

Péringuey is known to have had a high regard for Drury’s craftsmanship in the

fields of taxidermy and life-casting (Summers 1975: 103), the touching-up of the

photographic negatives Péringuey used in his monograph was done by

A. R. Walker, one of the Museum assistants (Péringuey 1911: 163); and he

commissioned a local artist, G. M. Winkles, to prepare the copies of the ‘Cold-

stream Stone’ and the ‘dancing women’ stone from which the coloured

lithographs were later prepared (Summers 1975: 107). It may be that Péringuey

did not consider Drury’s artistic abilities, even as a copyist, to be adequate for

such a task. There is, too, no evidence that Drury spent any time in the southern

Cape, although it is possible that he may have done so during the Anglo-Boer

War, which brought him to South Africa from Scotland (Summers 1975: 103).

Although Drury had a reputation for eccentricity and secretiveness

(Summers 1975: 128), E. M. Shaw (pers. comm. 1985) and R. H. N. Smithers

(1985 in litt.), both of whom knew him for many years, stressed that he was a

man of the highest integrity. Smithers also mentioned that Drury was a meticu-

lous worker who would probably have taken greater care in the execution of the

painting had he done it; and here the anomalous treatment of part of the last

figure may be mentioned again.

Presumably, the reason for anyone wanting to perpetrate such a forgery

would be to acquire, or share in, the prestige attaching to such a spectacular

‘discovery’. In the event, the ‘Coldstream Stone’ did not arouse international

interest, and appears to have been ignored, even locally, for nearly 50 years,

apart from Haughton’s (1926) publication. Although Péringuey was an entomol-

ogist by profession, rather than an archaeologist, there can be no doubt that he

would not have allowed the stone to be exhibited if he had had even the slightest

suspicion about its authenticity. There is, moreover, no indication that, until

recently, the genuineness of this artefact has been doubted by any of those who

have seen it, and then only on the grounds of the remarkable state of preser-

vation of the painting.

The conclusion to be drawn is that, although this investigation has shown

some lacunae, if not enigmas, in the documentary evidence, the pigment

analyses strongly suggest that Drury was not the author of the painting, nor

responsible for the retouching. The remarkable state of preservation of the paint

on the stone must be considered a taphonomic problem for which there is at

present no ready answer. Ultimately, however, the evidence yielded by this

22 ANNALS OF THE SOUTH AFRICAN MUSEUM

investigation suggests that there are no grounds for doubting the authenticity of

the painting on the ‘Coldstream Stone’.

ACKNOWLEDGEMENTS

Thanks for helpful comment on earlier drafts of this paper are due to

Dr M. J. Wicomb, Director of the Electron Microscope Unit, University of the

Witwatersrand, Johannesburg; Dr D. Crawford, Core Consulting, London;

Dr J. Deacon, National Monuments Council, Cape Town; and Dr A. J. B.

Humphreys, Department of Anthropology, University of the Western Cape,

Bellville. Louis Lawrence, Archaeology Department, South African Museum,

produced the maps. The two painted stones from Klasies River Mouth Cave 5

(Figs 1B, E) were first published by the excavators, Ronald Singer and John

Wymer (1982, pls 48—49), and thanks are due to Professor Singer for agreeing to

allow them to be photographed for use in this paper. Thanks are also due to

Dr G. A. Robinson, formerly Warden of the Tsitsikama Coastal National

Park, now Deputy Director, National Parks Board, who first took M. L. W. to

the Coldstream shelter and other sites, and made available his considerable

knowledge of the archaeological sites of the area.

REPERENGES

Battiss, W. 1948. The artists of the rocks. Pretoria: The Red Fawn Press.

Burkitt, M. 1928. South Africa’s past in stone and paint. Cambridge: The University Press.

CLARK, J. D. 1959. The prehistory of southern Africa. London: Pelican Books.

CoERTZE, R. D. ed. 1988. San rock art. University of Pretoria. Woodhouse exhibition. Pretoria:

University of Pretoria.

DENNINGER, E. 1971. The use of paper chromatography to determine the age of albuminous

binders and its application to rock paintings. Special Publications. South African Associa-

tion for the Advancement of Science 2: 80-84.

Goopwin, A. J. H. 1929. The Wilton Industry. Jn: Goopwin, A. J. H. & VAN Ret Lowe, C.

The Stone Age cultures of South Africa: 251-270. Annals of the South African Museum 27:

vii—298.

Goopwin, A. J. H. 1946. The loom of prehistory. South African Archaeological Society Hand-

book Series 2: 1-151.

HaucGuton, S. H. 1926. Note on a burial-stone. Transactions of the Royal Society of South

Africa 13 (2): 105-106.

Lee, D. N. & WoopuouseE, H. C. 1970. Art on the rocks of southern Africa. Cape Town:

Purnell.

Lewis-WILuiAMs, J. D. 1984. Ideological continuities in prehistoric southern Africa: the evi-

dence of rock art. Jn: SCHRIRE, C. ed. Past and present in hunter gatherer studies: 225-252.

Orlando, Fla.: Academic Press.

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): 1-128.

Rupne_r, I. 1982. Khoisan pigments and paints and their relation to rock paintings. Annals of

the South African Museum 87: 1-281.

RupDNeER, J. 1971. Painted burial stones from the Cape. Special Publications. South African

Association for the Advancement of Science 2: 54-61.

RUDNER, J. & RuDNER, I. 1970. The hunter and his art. A survey of rock art in southern Africa.

Cape Town: Struik.

SINGER, R. & Wyner, J. 1982. The Middle Stone Age at Klasies River Mouth in South Africa.

Chicago & London: Chicago University Press.

AN INVESTIGATION OF THE ‘COLDSTREAM STONE’ MANS)

SumMERS, R. F. H. 1975. A history of the South African Museum 1825-1975. Cape Town:

Balkema.

WiLicox, A. R. 1984. The rock art of Africa. London: Croom Helm.

WoopHousE, H. C. 1966. Whitcher’s Cave: the paintings. South African Archaeological Bulletin

21 (2): 92-93.

WoopHousE, H. C. 1968. The Coldstream burial stone. South African Journal of Science 64 (9):

341-344.

WoopHousE, H. C. 1969. The Coldstream burial stone. South African Journal of Science 65 (4):

WoopHousE, H. C. 1979. The Bushman art of southern Africa. Cape Town: Purnell.

WoopHousE, H. C. 1987. The Coldstream burial stone: a review. South African Journal of

Science 83 (9): 538-541.

YaTES, R., Gouson, J. & HALL, M. 1985. Trance performance: the rock art of Boontjieskloof

and Sevilla. South African Archaeological Bulletin 40 (142): 70-80.

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.,

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°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

eo... . the Pisure depictins 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.

M. L. WILSON, W. J. J. van RIJSSEN

| &

‘D. A. GERNEKE

AN INVESTIGATION OF THE

‘COLDSTREAM STONE’

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