ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 97 Band
August 1986 Augustus
Part 6 Deel
LATE TERTIARY AND EARLY
QUATERNARY FOSSIL MOLLUSCA
OF THE HONDEKLIP AREA,
CAPE PROVINCE, SOUTH AFRICA
By
BRIAN KENSLEY
&
JOHN PETHER
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LATE TERTIARY AND EARLY QUATERNARY FOSSIL MOLLUSCA
OF THE HONDEKLIP AREA, CAPE PROVINCE, SOUTH AFRICA
By
BRIAN KENSLEY
National Museum of Natural History,
Smithsonian Institution, Washington, D. C.
&
JOHN PETHER
Department of Cenozoic Palaeontology,
South African Museum, Cape Town
(With 42 figures and 1 table)
[MS accepted 10 February 1986]
ABSTRACf
A brief review of the Quaternary mollusc research of the west coast of South Africa is
provided. Molluscan fossils were obtained from two coastal regressive sedimentary complexes.
The older complex records a regression from c. +50 m, and is called the 50 m Complex.
Stratigraphic context in terms of global sea-level curves and the presence of Equus in this
complex suggests a Late Pliocene age. The subsequent complex records a regression from
c. +30 m, is called the 30 m Complex, and is considered Early Pleistocene in age.
The following 21 species and one subspecies are described as new: Patella hendeyi, Patella
hoffmani, Clanculus lutosus, Gibbula zonata patula, Bolma anoropha, Crepidula deprima,
Argobuccinum casus, Epitonium lycocephalum, Thais arenae, Ocenebra petrocyon, Trophon
carringtoni, Burnupena rogersi, Burnupena aestus, Fasciolaria dinglei, Melapium hawthornei,
Pseudoliva lutulenta, Drillia tempestae, Terebra canisaxi, Glycymeris fulleri, Isognomon
gariesensis, Cardita unica, Dosinia sicarisinus. A number of previously described mollusc species
are discussed or redescribed. The faunal composition and biogeography of the Hondeklip
assemblage are discussed, and the assemblage is compared with other west-coast fossil
assemblages. Temperature tolerances of some extant forms. present indicate that a warmer water
regime relative to modern conditions existed along the Namaqualand coast during the Late
Pliocene and Early Pleistocene. A temperature contrast across the Plio-Pleistocene boundary is
not clear-cut due to the faunal similarity of the 50 m and the 30 m Complexes. Significantly,
about 50 per cent of the fauna is extinct. Decreased diversity and the appearance of Choromytilus
meridionalis in the 30 m Complex may indicate cooling in the Early Pleistocene. Subsequently,
the restriction of warmer waters to the north led to the establishment of the modern west-coast
fauna. Lack of a broader regional database precludes more precise conclusions at this stage.
PAGE
Introduction. . . . . . . . . . . . . . . . . . . . . 142
Geological setting . . . . . . . . . . . . . . . . . . . . . . . . . 142
The Hondeklip and Avontuur A deposits. . 144
The Koingnaas and Swartlintjiesrivier localities. 148
Systematic descriptions . . . . . . . . . . . 149
Discussion . . . . . . . . . . . . . . . . . . . . 210
Faunal composition . . . . . . . . . 210
Zoogeographic affinities 217
Faunal comparisons . . 219
Concluding remarks. . 221
Acknowledgements 223
References. . . . . . . . . . . . . . . . . . . . . . 223
The fossil mollusc occurrences on the west coast of South Africa have
received scattered attention during the past 50 years, with reports becoming more
numerous in the last decade. Haughton (1932) presented an overview of the
west-coast deposits and described several of the fossil molluscs. Barnard (1962)
listed all the known Late Tertiary and Pleistocene molluscs from South Africa.
New species and records for the west coast have since been added by Carrington
& Kensley (1969), Kilburn & Tankard (1975), and Kensley (1972, 1977).
Diamond-mining activities on the Namaqualand coast have exposed coastal
marine deposits in several areas. Fossil molluscs from three localities, Koingnaas,
Swartlintjiesrivier, and Strandfontein, were the subject of the paper by
Carrington & Kensley (1969). Material recorded in the present paper also comes
from Koingnaas, plus two additional properties, Avontuur A, and Hondeklip
(Fig. lA, B). Exposures of several fossil-bearing beds were bulk-sampled, while
selective sampling was carried out when appropriate.
The bulk of the material recorded in the accompanying list (Table 1) is
housed in the South African Museum, as are holotypes and paratypes, which bear
SAM-PQ catalogue numbers. Where possible, paratypic material has also been
deposited in the National Museum of Natural History, Smithsonian Institution,
Washington, D.C. and bears USNM catalogue numbers.
GEOLOGICAL SETTING
This paper is a sequel to that of Carrington & Kensley (1969) in which several
new species obtained from similar deposits were described and a summarized
general account of the Cenozoic coastal Namaqualand stratigraphy was presen-
ted. Important observations recorded in that work were the recognition of two
distinct littoral marine complexes, the 45-50 m and the 17-21 m transgression
complexes, and their zone fossils Donax haughtoni Carrington & Kensley, and
Donax rogersi Haughton, respectively. The complex names indicate perceived
transgressive altimetric maxima. The older 45-50 m complex was considered
typified by fine, green quartzose sands and the associated thin-shelled D. haugh-
toni, while the younger 17-21 m complex was characterized by brown-stained,
coarse, quartzo-feldspathic sands, high-energy bedforms, and the robust
D. rogersi. An evolutionary relationship was inferred between the two Donax
species, and linked to the changed environment implicit in the contrasted
lithologies of the two complexes. Additionally, species obtained from the
45-50 m complex suggested a fauna of warm-water affinity.
Subsequent fieldwork on the properties of Hondeklip and Avontuur A has
furnished more information on the depositional environments, ages, and sea-level
history of the 45-50 m and 17-21 m complexes. This work will be presented in
detail in another article (Pether in prep.); thus only a condensed account is given
here. Modifications of the succession presented by Carrington & Kensley (1969)
are shown in the following summarized stratigraphy.
ATLANTIC
OCEAN
SOUTH
AFRICA
C
<10J30 Marine
~ pacKages
> 30 masl.
20-30 masl.
10- 20 masl.
Fig. 1. A. Locality map. B. Hondeklip and surrounding properties with main sampling areas
indicated. C. Bedrock contour map of Hondeklip and a portion of Avontuur A in metres above
sea-level (masl). Large numerals (90, 50, 30) indicate the overlying deposits of the marine
complexes; < 10 refers to the low-elevation, Mid-Pleistocene to Holocene raised beaches.
Palaeoshorelines at transgressive maxima of the 30 m and 50 m Complexes are shown.
Succession Age
*2 metre Beach Holocene
*5 metre Beach Late Pleistocene
8 metre Beach Middle Pleistocene
30 metre Complex Early Pleistocene
Regressive complex (incl. 29-34 m Beach and 17-21 m Complex)
50 metre Complex Late Pliocene
Regressive complex (= 45-50 m Complex)
90 metre Complex Early Pliocene
Regressive complex? (= 75-90 m Complex)
(Includes lower shoreface to off~hore deposits preserved in bedrock
depressions underlying the 50 m Complex at lower elevations.)
Kaolinized fluviatile deposits Latest Oligocene
to Early Miocene
Bedrock Archaean
Bedrock
The bedrock consists of Archaean quartzo-feldspathic gneiss, and bedrock
topography (Fig. Ie) has profoundly influenced the local character of the
overlying deposits. A major feature in the study area is a broad, coast-parallel,
channel-like structure in the bedrock, which has its western flank defined by a
bedrock ridge. Emergence of the bedrock ridge during regression resulted in a
barrier along the seaward flank of the channel, protecting the channel area from
open-coast conditions.
Kaolinized fluviatile deposits
This, the oldest sedimentary entity encountered, is locally preserved in a
bedrock depression. It is the remains of a fluvial arkosic infill deposited in the
bedrock channel. The underlying bedrock and this fluvial infill are kaolinized and
associated with it are silcrete slabs and boulders that are indurated portions of the
fluvial deposits that have been exhumed by extensive erosion.
The bedrock is considered to have been incised during the major Oligocene
regression that has been identified in the offshore record (Siesser & Dingle 1981).
Subsequently, infilling Late Oligocene to Earliest Miocene fluvial deposits were
kaolinized during the Early Miocene under tropical conditions; within the
weathering profile titanium-rich silcrete developed. Sea-level fluctuations and
concomitant marine and fluvial erosion during Middle Miocene to Late Pliocene
times exhumed and modified the bedrock channel. No Miocene marine deposits
are preserved.
The 90 m Complex deposits
This high elevation complex, the 75-90 m Complex of Carrington & Kensley
(1969), is present between 50-90 m above sea-level (masl), but no exposures at
these elevations exist in the study area. However, underlying the younger 50 m
Complex in a local bedrock depression, is a remnant deposit that may have been
deposited, wholly or in part, during the same sea-level cycle. This is a partly
indurated, mouldic, coquinoid, muddy conglomerate, which exhibits extensive
(though not laterally continuous), authigenic phosphorite (microsphorite) deposi-
tion. A ubiquitous, similar, phosphorite gravel content in overlying 50 m
Complex gravels indicates the erosion of this previously more extensive lithology.
Fragmentary, worn, phosphate-mineralized, marine and terrestrial verte-
brate fossils are associated with this bed. These have a Mio-Pliocene aspect
(Q. B. Hendey pers. comm.). The mouldic coquinoid fauna has both intertidal
and offshore aspects. The bed is considered to have been storm-deposited in a
lower shoreface to offshore environment during regression, in an embayed
situation. Extensive microsphorite precipitation driven by upwelling took place
during calmer periods. This bed has previously been referred to as 'lower E stage'
(e.g. Tankard 1975b, 1975c).
The 50 m Complex deposits
This complex is present mainly to the east of the bedrock ridge (Fig. 1C).
Exposures in open-coast situations reveal a regressive vertical facies sequence of
lower-shoreface, upper-shoreface, foreshore, and aeolian environments. The
basal gravel is seen to be a transgressive veneer, which has been redeposited and
supplemented during regression in a lower-shoreface context. The bedrock ridge
promoted the development of back-barrier environments in its landward lee
during regressive emergence.
Evidence of a minor transgressive sea-level fluctuation is preserved in this
complex and consists of a laterally persistent surface produced in back-barrier
deposits, an anomalously thick foreshore deposit, a vertical aspect to normally
lateral open-coast barrier facies deposition, and a tidal inlet-dominated barrier
overlying back-barrier deposits.
The 30 m Complex deposits
The 50 m Complex was eroded by a subsequent transgression and overlying
the transgressive disconformity is the 30 m Complex. This is a seaward-thickening
wedge, which, beneath foreshore deposits extending from the transgressive
maximum near +30 m, progressively incorporates upper-shoreface, and then
lower-shoreface deposits. This complex is preserved mainly to the west (seaward)
of the bedrock ridge (Fig. 1C). Sea-level altitude relative to bedrock topography
suggests that generally during 30 m Complex times the effect of bedrock
topography was not as marked as during the previous sea-level cycle. In the study
area there is no evidence of back-barrier environments during 30 m Complex
times.
The 30 m Complex subsumes the 17-21 m Complex and the 29-34 m Beach
of Carrington & Kensley (1969). No evidence of a stillstand at c. 20 masl is
present in Hondeklip exposures. Instead, 30 m Complex lower-shoreface gravels
extend seawards from that elevation. However, the possibility that stillstands
occurred subsequently during the regression cannot be dismissed at this stage.
The complex is present almost up to the modern coastline, where it is overlapped
by younger palaeoshorelines at elevations of less than 10 mas!'
Age of the marine complexes
Onshore and offshore evidence of Tertiary sea-levels around southern Africa
has been synthesized in a curve by Siesser & Dingle (1981) (Fig. 2A); this curve
exhibits a general correspondence with the trends of global sea-level produced by
Vail & Hardenbol (1979). Sea-level history and age inferred for the deposits at
Langebaanweg (Fig. 2B) is consistent with the global sea-level record (Hendey
1981a, 1981b) and, pointing out the feasibility of altimetric correlation along the
southern-African west coast, Hendey proposed a preliminary correlation scheme
between Langebaanweg and west-coast localities involving the high-elevation
(> 10 masl) deposits.
"The correlation scheme was related to the sea-level curve of Vail &
Hardenbol (1979). Beard et al. (1982) provided greater resolution for the
Quaternary portion of the 'Vail curve' (sea-level cycles Ql to Q8 (Fig. 2C)).
Although methodological aspects and the chronology of the curve are controver-
sial, it is accepted here as presented and considered to be of provisional utility.
For example, Vail & Hardenbol (1979) and Beard et al. (1982) placed the Plio-
Pleistocene boundary at 2,8 Ma; in contrast, in terms of the most recent ICS
decision (Aguirre & Pasini 1985), the boundary at the Vrica section is ~ 1,64 Ma.
Regardless of this controversy, it is the sea-level curves, as gross indicators, that
are deemed relevant, and interestingly they do show a general correspondence
with our present (though scant) knowledge of west-coast deposits. In this paper
the Plio-Pleistocene boundary is taken at ~1,6 Ma; thus sea-level cycle Ql
(Fig. 2C) is considered Late Pliocene.
An important feature of Hendey's (1981a, 1981b) correlation scheme was the
correlation of the 90 m Complex in Namaqualand with the Early Pliocene
Varswater Formation at Langebaanweg; the latter, in turn, related to the
transgressive sea-level period TPI of Vail & Hardenbol (1979) (Fig. 2B). An
Early Pliocene age for the 90 m Complex is consistent with observations at
Hondeklip. The likelihood of the complex being a regressive package in
Namaqualand suggests correlation with sea-level cycle TP2.
The presence of a species of Equus in the 50 m Complex suggests that it
cannot be older than the 1,9 Ma mammalian dispersal event in Africa (Lindsay
et al. 1980). Similar to the age inferred by Hendey (1981a) for the Baards Quarry
Fluviatile Deposits in the south-western Cape which contain Equus, the 50 m
60 50 .0 30 20 10 my
CR. PA L. EOCENE OLIG MIOCENE PE L E M L E L E M L L
- 4
-3
- 2
- 1 +
0 m(x 102)
- 2
- 3
-4
-5
4 3 my
m
90?II TIAM :~ Surface 60, PPM I deposit, ?_GM_? QSM \ 1,, 30,. ., , BQF
\ : "'---.., ""'.. . , \ +\ I \ ,\ , I , 0\ . ,
30
Bedrock
75 50 25
Years B.P x 103
I
3 -2.8 2
PLIOCENE
1
QUATERNARY
Fig. 2. A. Gross Tertiary sea-level trends after Siesser & Dingle (1981). B. Langebaanweg
deposits (south-western Cape) related to global sea-level trends of Vail & Hardenbol (1979)
(after Hendey 1981b). GM = Gravel Member, QSM = Quartzose Sand Member, PPM = Pelletal
Phosphorite Member, AM = Anyskop Marine Deposits-all of the Varswater Formation.
AT = Anyskop terrestrial deposits, BQF = Baard's Quarry fluviatile deposits. C. Gross
Quaternary sea-levels of Beard et ai. (1982).
Complex is considered Late Pliocene in age and is correlated with the regressive
portion of sea-level cycle Q1 of Vail & Hardenbol (1979) and Beard et al. (1982).
The 30 m Complex must reflect a subsequent high sea-level, and it is thus
correlated with cycle Q2 in the Early Pleistocene. A Late Pliocene and Early
Pleistocene age for the 50 m and 30 m Complexes, respectively, is probably
consistent with the warmer-water faunal elements present, relative to the colder-
water assemblages of the younger, low-elevation complexes and of the modern
coast.
The age of these low-elevation ? 10 masl) palaeo-shorelines has recently
been assessed (Hendey & Volman 1986). Fossil evidence associated with the 8 m
Beach at Saldanha suggests a late Early Pleistocene age to Hendey & Cooke
(1985) and it was correlated with sea-level cycle Q2. Since this report prefers to
relate the 30 m Complex to cycle Q2, the 8 m Beach is here considered to reflect
cycle Q3. This younger (Middle Pleistocene) age is inconsistent with the age
constraints inferred from the !fossil evidence at Saldanha, an issue yet to be
resolved. The 5 m and 2 m shorelines are considered to be Late Pleistocene
(Eemian) and Holocene respectively (Hendey & Volman 1986). Flemming (1977)
has submitted evidence that the Flandrian transgression reached 3 masl and
profoundly influenced the development of the Langebaan lagoon. Recently a
radiocarbon date of 3800 BP has been obtained at 2,5 masl at Verlorevlei
(1. Parkington pers. comm.).
THE KOINGNAAS AND SWARTLINTJIESRIVIER LOCALITIES
Some basal exposures of the deposits on De Beers properties neighbouring
Hondeklip (Fig. 1B) were examined and seen to contain molluscs not well
represented at Hondeklip. In the case of the 50 m Complex outcrops (Koingnaas
KN and Swartlintjies SL), this is due to the occurrence of thicker, muddy, distal
lower shoreface deposits in bedrock depressions at these localities. At Hondeklip
the deposits of this environment are thinner and more cryptic, due to the
em bayed palaeocoastal geomorphology formed by the bedrock topography, in
contrast to the more open-coast, palaeocoastal situation considered applicable to
the De Beers properties. This facies overlies bedrock or local 90 m Complex
remnants, and is overlain by gravelly, proximal lower shoreface deposits of the
50 m Complex. It is characterized by a molluscan fauna with both intertidal and
sublittoral components, and in these respects the similarity with the 90 m
Complex remnants is noteworthy.
An outcrop of the 30 m Complex on Koingnaas also yielded species not
encountered at Hondeklip (Koingnaas KL). At this exposure a massive sand,
containing thin gravel beds and a mud lens -20 cm thick, overlies bedrock at
10 masl. It attains its greatest thickness (-2 m) in a local bedrock depression.
This unit is overlain by the 30 m Complex upper shoreface facies. It is thus
interpreted as 30 m Complex proximal lower shoreface deposits. The open-coast
forms Isognomon sp., Mactra sp., and Turritella carini/era dominate the
assemblage.
In the following section, in addition to the previously undescribed material,
discussion is given to some previously recorded taxa, where relevant.
As is frequently the case in fossil molluscan systematics, generic allocation
cannot always be done with certainty, due to the lack of essential diagnostic
information (e.g. radular and opercular structure). This uncertainty in generic
placement was constantly in the forefront of the authors' thoughts when writing
descriptions; nevertheless it was decided not to indicate this uncertainty with
quotation marks or any other such device.
Unless otherwise stated, all material mentioned in this work was collected by
the authors.
Class GASTROPODA
Family Haliotidae
H aliotis saldanhae Kensley, 1972
SAM-PQ-A V229, several fragments, Avontuur A, Trench 3.
SAM-PQ-KN370, numerous fragments, including seven triangular frag-
ments, each the upper columella portion of an individual shell, Koingnaas KN-l.
Previous records
Langebaanweg, Pliocene.
Remarks
Several large abalone specimens have been seen in situ at Avontuur, but the
shell material is usually so fragile and friable as to make recovery almost
impossible. Several fragments of shell, however, have been recovered, bearing
the distinctive rounded spiral ridges and broadly rounded oblique axial ridges
characteristic of H. saldanhae.
A single specimen (SAM-PQ-AV506), c. 60 x 140 mm, was recovered,
which retains its overall shape and proportions. Unfortunately none of the
exterior shell structure remains, making positive identification impossible. The
size of this specimen places it well beyond the maximum for H. spadicea
Donovan, 1808, but within the size range of living H. midae Linnaeus, 1758, of
the west coast.
Family Fissurellidae
Amblychilepas scutellum (Gmelin, 1791)
Amblychilepas scutella: Barnard, 1962: 191; 1963: 286, figs 21b, 22d-f.
Amblychilepas scutellum: Kilburn & Rippey, 1982: 35, pI. 2 (fig. 11), pI. 6 (fig. la, b).
Material
SAM-PQ-HB319, 4 specimens; TL 22,5 mm, 23,3 mm, 29,0 mm, 1 dam-
aged, Hondeklip Zone 12. SAM-PQ-HB74, 2 specimens, Hondeklip, 50 m
Complex.
Previous records
Living: Angola to Natal.
Fossil: Saldanha, Little Brak River, Sedgefield, Durban, Inhambane.
Remarks
The living subspecies, A. s. scutellum of the west coast, is characterized by
the possession of 'moderately to extremely raised ends' (Kilburn & Rippey 1982:
35). The present material consists of six specimens not at all saddle-shaped, i.e.
sitting flat on a horizontal surface, and one specimen with very slightly raised
ends.
Fissurellidea aperta (Sowerby, 1825)
Pupillaea aperla Sowerby, Barnard, 1962: 191.
Fissurellidea aperla (Sowerby) Barnard, 1963: 288, fig. 21e. Kensley, 1973: 32, fig. 46. Kilburn &
Rippey, 1982: 36, pI. 3 (fig. 3), pI. 6 (fig. 13).
Material
SAM-PQ-KN507, 1 specimen, 22,8 x 12,8 mm, 4 fragments, Koingnaas
KN-l.
Previous records
Living: Namibia to Transkei, shallow infratidal.
Fossil: Pleistocene, Algoa Bay.
Tugali barnardi (Tomlin, 1932)
Tugalia barnardi: Barnard, 1963: 300, figs 21d, 22a-c.
Material
SAM-PQ-KN392, 2 specimens, 4,0 x 2,8 mm, 8,1 x 4,6 mm, Koingnaas
KN-l.
Previous records
Living: off Cape Point, 360 m; False Bay, 18 m; St. Francis Bay, 12 m.
Dead: Cape Point to Cape Morgan, intertidal to 360 m.
Remarks
The fossil specimens have been compared with material in the South African
Museum, and agree very closely with specimens from St. Francis Bay
(SAM-A9299). The external ridge sculpture is as figured by Barnard (1963: 291,
fig. 22b).
Family Patellidae
Patella hendeyi sp. novo
Figs 3, 4
Material
Holotype. SAM-PQ-A V454, 126 x 107 mm, Avontuur A, 50 m Complex.
Paratypes. SAM-PQ-AV455-457, 7 specimens, 68 x 57 mm, 74 x 61 mm,
90 x 76 mm, 94 x 77 mm, ? x 109 mm, 142 x 133 mm, 146 x 137 mm, Avon-
tuur A, 50 m Complex. SAM-PQ-HB458, 148 x 130 mm, Hondeklip Z4A,
50 m Complex (with attached barnacle shells). USNM 400979, 2 specimens,
122 x 112 mm, 138 x 122 mm, Avontuur A, 50 m Complex.
Fig. 3. Patella hendeyi. Holotype in dorsal view.
Scale = 30 mm.
Fig. 4. Patella hendeyi. Holotype in upper right corner, remainder of specimens paratypes.
Scale = 30 mm.
Non-type material. SAM-PQ-AV459, 4 damaged and incomplete speci-
mens, Avontuur A. SAM-9921, 2 specimens, 106 X 91 mm, 98 x 85 mm,
Graauweduinen, Vanrhynsdorp District coast, Cape Province, coil. H. Harger.
SAM-9932, 136 x 124 mm, Graauweduinen, Vanrhynsdorp District coast, coli.
H. S. Harger.
Description
Shell heavy, relatively low, with subcircular outline, only slightly longer than
wide, with apex anterior to midline, apex in all specimens eroded. Sculpture
consisting of fine radial lines of subequal strength (about 300 on circumference of
holotype, 200-300 in paratypes), becoming divided and/or distorted close to
circumference. No juvenile scars on any specimens.
Remarks
The specimens numbered SAM-9921 and 9932 are those referred to by
Haughton (1932: 30) as 'Patella sp. (large)'.
Of present-day patellids living on the southern African coast, in terms of
size, only P. tabularis Krauss, 1848, reaches the size of the largest specimen
considered here. This species, however, characteristically has very large acute
radiating ribs alternating with weaker ribs, and an irregular shell margin.
Patella safiana Lamarck, 1819, of West Africa has more regularly spaced but
stronger ribs.
Patella argenvillei Krauss, 1848, has a narrower and higher shell.
Only very large shells of P. granularis Linnaeus, 1758, approach the present
material in shape and sculpture. Prof. G. Branch (U.c.T.) has donated a
specimen of P. granularis (91,0 x 74,9 mm), that in shape closely resembles some
of the smaller of the present fossils, and has faint radial sculpture as in the largest
fossil. The possibility exists that P. hendeyi represents either a precursor of
P. granularis or an earlier and much larger form of the same species.
Patella (Ancistromesus) Juenzalidai Herm, 1969, from the Pliocene of central
Chile, bears a striking resemblance to the present species. The holotype and
paratype of this species, at 209 mm and 188 mm length, are even larger than
specimens of P. hendeyi, but have a very similar outline and general proportions
and also the very fine radiating ribs seen in the Namaqualand species.
Etymology
The species is named for Dr Q. B. Hendey of the South African Museum.
Patella hoffmani sp. novo
Figs 5, 6
Material
Holotype. SAM-PQ-AV508, 56,7 x 51,0 x 31,0 mm
height), Avontuur A.
Paratypes. SAM-PQ-AV402, 9 specimens,
32,3 x 27,8 x 10,6 mm; 34,8 x 31,4 x 13,8 mm;
23,3 x 18,9 x 8,3 mm;
42,9 x 38,5 x 15,8 mm;
44,5 x 41,0 x 20,0 mm, 47,7 x 43,6 x 22,4 mm; 47,3 x 42,9 x 20,0 mm;
46,7 x 41,8 x 25,4 mm; 54,2 x 48,2 x 30,8 mm; Avontuur A. SAM-PQ-HBI88,
48,4 x 41,2 x 27,1 mm, Hondeklip, 50 m Complex. SAM -PQ-HBI91, 2 speci-
mens, 48,5 x 41,9 x 23,0 mm; 51,6 x 45,8 x 26,7 mm; Hondeklip, 50 m Com-
plex. SAM-PQ-HBI42, 3 specimens, 39,7 x 32,0 x 18,7 mm; 46,9 x 38,8 x
21,7 mm; 47,7 x 40,2 x 25,6 mm; Hondeklip, 50 m Complex. SAM-K4755,
49,5 x 46,0 x 24,1 mm; Swartlintjies SL2, colI. A. J. Tankard. SAM-
PQ-HB509, 3 specimens, 43,8 x 40,4 x 32,3 mm; 49,4 x 45,0 x 29,8 mm;
51,2 x 44,1 x 34,5 mm; Hondeklip Zone 4C, 50 m Complex. USNM 400980,
6 specimens, ? x 37,2 x 18,4 mm; 42,2 x 38,5 x 17,0 mm; 44,4 x 39,4 x
18,6 mm; 45,8 x 40,6 x 27,5 mm; 52,3 x 47,5 x 31,1 mm; 54,3 x 49,0 x 25,3 mm;
Avontuur A, 50 m Complex.
Fig. 5. Patella hoffmani. A-C. Lateral, dorsal and ventral views of holotype.
Scale = 20 mm. D. Specimen showing juvenile scars. Scale = 10 mm.
Non-type material. SAM-PQ-HB249, 4 specimens, Hondeklip, B Block,
50 m Complex. SAM-PQ-A V402, 12 specimens, Avontuur A, 50 m Complex.
SAM-PQ-AV51O, 4 specimens, Avontuur A, 50 m Complex. SAM-K4754,
3 specimens, Koingnaas, coli. A. J. Carrington. SAM-K4756, 2 specimens,
Somnaas SM4, 50 m, coli. A. J. Tankard. SAM-PQ-HB461, 27 specimens,
Hondeklip, Zone 4C, 50 m Complex.
Description
Shell generally high-conical; length/width ratio range 80-93 per cent; sides
very faintly convex; apex at or close to midpoint of antero-posterior line.
Sculpture consisting of radiating flattened ribs of equal strength; under
magnification, ribs seen to be built up of numerous concave growth lines,
Fig. 6. Patella hoffmani. Range of specimens in dorsal view, some showing colour-rays.
Scale = 30 mm.
somewhat flattened, separated by shallow sharp groove. Number of ribs on
circumference of large specimens ranging from 105 to 124. Shell margin finely and
evenly serrate. Large shells often with several oval juvenile scars, up to nine per
shell; scars usually 4-10 mm greatest diameter. Some shells bearing dark-brown
radiating bands separated by narrow pale bands; dark rays more distinct closer to
shell apex; near margin, dark bands coalescing to form continuous dark-brown
band.
Remarks
The almost circular circumference and tall conical shape of Patella hoffmani
bears little resemblance either to any of the living or fossil southern or West
African patellids. The sculpture of equal flattened radiating ribs, however, is
almost identical to that of Patella argenvillei.
The subcircular and high-conical shape of P. hoffmani is reminiscent of
Nacella clypeater (Lesson, 1831) from Chile. This species, however, possesses
fewer radiating ribs which, unlike the flattened ribs of P. hoffmani, are low and
rounded.
Etymology
The species is named for Mr F. Hoffman, of Transhex Pty Ltd diamond
company.
Family Trochidae
Clanculus lutosus sp. novo
Fig. 7
Material
Holotype. SAM-PQ-KN511, 16,3 x 18,3 mm, outer lip damaged, Koing-
naas KN-l.
Paratypes. SAM-PQ-KN512, 5 specimens, Koingnaas KN-l.
Non-type material. SAM-PQ-KN513, 12 damaged specimens, Koingnaas
KN-l.
Description
Shell top-shaped, of at least 4 postnatal whorls. Whorl profile straight to
barely convex. Sculpture of spiral lines bearing bead-like, close-set tubercles;
second whorl with 4-6 spirallirae; third whorl with 10-12 lirae; body whorl with
10 lirae above shoulder, 12-14 on base. Shoulder narrowly rounded. Umbilicus
broadly open, smooth, with single low spiral ridge. Columella with bipartite
tubercle on lower portion. Outer lip slightly thickened, but lacking internal
plicae.
Remarks
Clanculus murrayi has been described from the Quaternary of the Honde-
klipbaai area (Carrington & Kensley 1969). This species, while having beaded
spirallirae as in C. lutosus, is characterized by a stepped profile, with two strong
lirae on the whorl periphery.
Clanculus atricatena Tomlin, 1921, known from Transkei to Zululand,
possesses a grooved lower tubercle on the columella, giving it a bipartite
appearance. The spiral lirae of this species, however, are more finely beaded,
while the outer lip possesses a strong posterior tubercle on the inner surface.
Clanculus kraussi Philippi, 1846, from West Africa, possesses finely beaded
spiral lirae, but lacks the bipartite columella tubercle of C. lutosus. Two other
West African species, however, show a stronger resemblance to the present
material. Clanculus santamariae Gofas, 1984, from Benguela, Angola, has spiral
lirae with fewer but larger beads, a crenulate umbilical opening, and a non-bifid
columella tooth. Clanculus pseudocorallinus Gofas, 1984, also from Bf:nguela,
Angola, possesses sculpture very similar to that of C. lutosus, but also has a
crenulate umbilical opening, and a non-bifid columella tooth. Clanculus
corallinus (Gmelin, 1790) of the Mediterranean has fewer spirallirae with larger
beads, but does have a bifid columella tooth, and a non-crenulate umbilical
opening as in C. lutosus.
Etymology
The specific name is derived from the Latin 'lutosus', muddy, and refers to
the sediments from which the material was collected.
Gibbula zonata patula subsp. novo
Fig. 8
Material
Holotype. SAM-PQ-HB514, 5,0 x 6,9 mm, body whorl damaged, Honde-
klip Zone 3.
Paratypes. SAM-PQ-HB117, 4 specimens, 4,9 x 5,6 mm, 4,3 x 5,5 mm,
6,1 x 7,2 mm, 6,3 x? mm, Hondeklip Zone 3. SAM-PQ-HB345, 6,8 x
6,5 mm, Hondeklip Zone 12.
Description
Shell broader than high, somewhat globular, of 4 whorls. Profile of whorls
evenly rounded. Sculpture of rounded spiral bands; penultimate whorl with
6 bands; body whorl sculpture variable, with 8-10 bands on upper whorl; some
specimens with intermediate finer lirae; ventral whorl with 8-12 lirae. Umbilicus
broad, open in all specimens. Raised rounded bands in some specimens having
darker colour than rest of shell.
Fig. 8. Gibbula zonata patula. Holotype in upper left corner, remainder of specimens
paratypes. Scale= 3 mm.
Remarks
The present material is very similar to Gibbula zonata (Wood, 1828), known
from the intertidal and shallow infratidal of South West Africa-Namibia to False
Bay, Cape. The variable spiral sculpture of the present material agrees well with
the living species. The umbilicus is usually closed in adults, but occasionally
remains open in the living species, while all the fossil specimens have open
umbilici. As the fossils come from a distinctly warm-water assemblage, and
G. zonata is characteristically a cold-water species, it seems essential to regard
the fossil as a chrono-subspecies, or as a precursor species of G. zonata.
Etymology
The specific name derives from the Latin 'patulus', standing open, and refers
to the open umbilicus of the species.
Family Turbinidae
Holma anoropha sp. novo
Fig. 9
Material
Holotype. SAM-PQ-KN516, operculum, length 23,2 mm, Koingnaas
KN-l.
Paratypes. SAM-PQ-KN517, 6 opercula, lengths 17,6, 20,0, 25,0, 26,0,
26,1 mm, one damaged, Koingnaas KN-l. SAM-PQ-SL518, operculum, length
12,9 mm, Swartlintjies SL-20.
Description
Operculum narrowly elongate-oval, narrower at apical end; internal surface
gently convex, with numerous growth-lines. Apex on outer ventral margin.
External surface smooth, with rounded ridge in dorsal half. Operculum thickest
posterodorsally.
Remarks
None of the living southern African turbinids possesses an operculum
resembling the present material. Bolma andersoni (Smith, 1902), known from
Transkei to Zululand, possesses a more broadly oval operculum, but has similar
features, i.e. a smooth external surface, posterodorsally thickset, and with the
apex on the outer ventral margin.
Unfortunately, no turbinid shell that could be associated with these opercula
has been found in the deposits.
Etymology
The specific name, from the Greek 'anorophos', without a roof, alludes to
the fact that no shell has been found associated with the present opercula.

Family Turritellidae
Turritella declivis Adams & Reeve, 1850
Turritella declivis: Barnard, 1963: 167, fig. 33c. Kensley, 1973: 74, fig. 243.
Material
SAM-PQ-KN350, numerous fragments, up to 47 mm in length, maximum
diameter 13 mm, Koingnaas KN-l.
Previous records
Living: False Bay, 51 m; off Cape St. Francis, 12-112 m.
Dead: False Bay to Agulhas Bank, East London, 12-248 m.
No fossil record.
Remarks
Most of the present material is unworn, and shows the straight, even profile
of typical T. declivis. A few fragments show slight development of a cingulum
above the suture, but none show the hollowing of the whorls seen in the
T. excavata form (see Barnard 1963: 168).
Family Crepidulidae
Calyptraea kilburni nom. novo
Fig. 10
Calyptraea aurita striata Carrington & Kensley, 1969: 201, fig. 2c, pI. 22. Kilburn, 1980: 194.
Material
SAM-K1433 (holotype of C. aurita striata), 8,1 x 19,6 mm, Swartlintjies,
45-50 m transgression. SAM-PQ-HB148, 2 specimens, 9,8 x 27,2 mm, one
damaged, Hondeklip HB-4, 50 m Complex. USNM 400981, 2 specimens,
Fig. 10. Calyptraea kilburni. Left, dorsal view; centre and right, ventral view.
Scale = 10 mm.
9,0 x 23,3 mm, 11,4 x 30,4 mm, numerous fragments, Hondeklip, 50 m Com-
plex.
Remarks
Kilburn (1980) correctly pointed out that the name 'striata', used as a
subspecific epithet by Carrington & Kensley (1969), was thrice preoccupied.
Further, C. aurita (Reeve) from Chile differed markedly from the living South
African species now named C. barnardi Kilburn, 1980. The west-coast fossil
species, while having fine radiating striae similar to the Chilean C. aurita, differs
in the structure of the septum. A new name is thus required for the Namaqualand
fossil species.
Etymology
The species is named for Dr R. N. Kilburn of the Natal Museum, foremost
southern African malacologist.
Crepidula deprima sp. novo
Fig. 11
Material
Holotype. SAM-PQ-HB519, 22,6 x 9,2 mm, Hondeklip Zone 4A.
Paratypes. SAM-PQ-HB520, 2 specimens, 12,3 x 5,5 mm, 16,9 x 7,2 mm,
2 fragments, Hondeklip Zone 12. SAM-PQ-HBI25, 1 specimen, 14,2 x
7,4 mm, Hondeklip HB-4. SAM-PQ-HB521, 3 specimens, 16,6 x 8,1 mm,
16,0 x 6,3 mm, 15,1 x 7,9 mm, 2 fragments, Hondeklip Zone 4A.
USNM 400982, 1 specimen, 16,1 x 8,0 mm, 2 fragments, Avontuur A.
Description
Shell elongate-oval, strongly dorso-ventrally depressed, sides roughly paral-
lel, anteriorly evenly rounded, dorsum convex in long axis, dorsal surface with
irregular growth lines. Apex terminal on rounded posterior end. Ventral 'shelf'
between posterior margin and start of septum. Free edge of septum evenly
concave.
Remarks
Crepidula deprima bears little resemblance in its general shape and
proportions to any of the southern or West African crepidulids. Crepidula plana
Say, 1822, of the east coast of the U.S.A. is a similarly depressed species with a
concave septum, but is much larger (up to 40 mm in length), with a narrowly
rounded posterior end and a truncate anterior end.
Etymology
The specific name is derived from the Latin 'deprimo', depressed, and refers
to the dorso-ventrally depressed condition of the present species.
Fig. 11. Crepidula deprima. A. Holotype in dorsal and ventral view. B. Paratypes.
Scales = 10 mm.
Family Naticidae
Natica d. andansoni (Blainville, 1824)
Fig. 12
Natica adansoni (Blainville), Nickles, 1950: 80, fig. 117.
Material
SAM-PQ-KN522, 8 specimens, ? x 5,7 mm, 6,3 x 6,4 mm, 15,8 x
14,3 mm, ? x 15,2 mm, 18,2 x? mm, 20,9 x 17 ,8 mm, 20,4 x 19,0 mm,
21,5 x ? mm, Koingnaas KL south face.
Previous records
Living: Madeira; Cape Verde Islands; Morocco to Angola.

Description
Shell slightly wider than long; spire low. Umbilicus with slight ridge at outer
margin; columella callus forming convex lobe or bulge in umbilicus. Single
specimen with broad band of brown pigment stretching from above umbilicus on
to base and anterior end of outer lip.
Remarks
As all the present specimens are to some degree abraded, a definite
identification is difficult. The form of the umbilicus, however, most closely
resembles that of N. adansoni. The presence of a band of pigment around the
umbilicus and base, typical of the living species, further suggests this identifica-
tion.
Sinum concavum (Lamarck, 1822)
Fig. 13
Material
SAM-PQ-HB515, longest diameter 16,8 mm, widest diameter 12,9 mm,
Hondeklip B Block.
Previous records
Senegal to Mo<;amedes, Angola.
Description
Shell thin, fragile. Protoconch indistinct; 4 postnatal whorls. Spire low,
barely protruding above surface of body whorl. Sculpture consisting of low
irregular flattened spiral lirae of varying widths, becoming obscure in columella
area. Lirae +50 on body whorl, difficult to count as wider lirae tend to divide
obscurely into several finer bands. Twenty-five lirae on penultimate whorl;
obscure on earlier whorls.
Family Cymatiidae
Argobuccinum casus sp. novo
Figs 14, 15
Material
Holotype. SAM-PQ-A V523, 89,9 x 50,9 mm, apex and base damaged;
Avontuur A, 50 m Complex.
Paratypes. SAM-PQ-AV405, 3 specimens, 66,2 x 37,8 mm, 50,7 x
28,6 mm, 43,8 x 27,8 mm, Avontuur A, 50 m Complex. SAM-PQ-HB339,
4 specimens, 62,7 x 37,4 mm, 40,3 x 19,4 mm, 32,5 x 18,2 mm, 31,2 x 19,4 mm,
Hondeklip Zone 12. SAM-PQ-AV524, 75,5 x 48,0 mm, 8 smaller specimens
48,8 x 21,9 mm to 29,0 x 15,8 mm, Avontuur A, 50 m Complex. USNM 400983,
2 specimens, 48,7 x 29,0 mm, 33,3 x 17,9 mm, Hondeklip Zone 12.
USNM 400984, 58,9 x 32,9 mm, Avontuur A, 50 m Complex. USNM 400985,
4 specimens, 27,8 x 15,1 mm to 42,0 x 22,8 mm, Avontuur A, 50 m Complex.
Non-type material. SAM-PQ-A V525, 22 specimens, Avontuur A, 50 m
Complex. SAM-PQ-HB526, 15 specimens, Hondeklip Zone 4A.
Description
Shell high-spired, very slightly dorso-ventrally compressed, with at least
5 postnatal whorls bearing a total of about 6 varices, latter not continuous from
whorl to whorl. Profile of whorls evenly convex. Shell somewhat variable with
regard to degree of elongation; few squatter specimens resembling Argobuccinum
pustulosum (Lightfoot, 1786) in general proportions. Siphonal canal less than half
length of aperture, slightly flexed to left. Outer lip with 11 or 12 ridges, becoming
paired in older specimens. Columella smooth. Postnatal sculpture consisting of
strong flattened spiral ridges or ribbons with narrow intervening grooves, 7 or 8
on earlier whorls, 13 to 15 on body whorl with finer alternating ridges appearing.
Fig. 14. Argobuccinum casus. A. Range of paratypes. Scale = 20 mm. B. Holotype.
Scale = 10 mm.
Spiral ribbons becoming worn on body whorl, and seen to consist of 2 high ridges
connected and covered by outer shell layer. Spiral ridges running on to dorsum of
siphonal canal. Very faint rounded axial ridges seen in two specimens, 7 ridges
between varices, becoming obsolete on body whorl.
Remarks
Of the cymatiids currently living on the west coast, A. casus most closely
resembles Argobuccinum pustulosum (Lightfoot). This latter is said to occur in
two forms or subspecies (see Kilburn & Rippey 1982: 75), with A. pustulosum
proditor (Frauenfeld, 1865) being the west-coast form. Argobuccinum casus
generally has a narrower shell even than this west-coast form, lacks the rounded
tubercles on the spiral bands, has rounded tubercles rather than elongate ridges
on the inner surface of the outer lip, has a more clearly defined siphonal canal,
and lacks the short marginal digitiform projections of the lower outer lip. Even on
the few specimens showing faint axial ribs, where these intersect the spiral bands,
no rounded tubercles are formed as in the living species.
The present species was also compared with Pleistocene material of
A. pustulosum (Lightfoot) from the Saldanha Bay area, and again, all the
differences listed above are apparent.
Cymatium parthenopeum (von Salis, 1793), living on the coasts of West and
south-east Africa, and known as a fossil from the Pliocene of Italy, possesses
fewer spiral bands, but stronger axial sculpture than A. casus.
Etymology
The specific name, the Latin for 'adventure', is derived from the farm-name
Avontuur, a locality for the present species as well as several others described in
this work.
Family Epitoniidae
Epitonium (Gyroscala) lycocephalum sp. novo
Fig. 16
Material
Holotype. SAM-PQ-HB527, 14,5 x 6,6 mm, Protoconch damaged, Honde-
klip Zone 4A, 50 m Complex.
Paratypes. SAM-PQ-HB528, 2 specimens, 13,0 x 5,0 mm (apex missing),
12,9 x 4,6 mm, Hondeklip Zone 12, 50 m Complex.
Non-type material. SAM-PQ-HB529, 15 fragments, Hondeklip A Block,
30 m Complex.
Fig. 16. Epitonium lycocephalum. A. Paratype. Scale = 3 mm.
B. Holotype. Scale = 5 mm.
Description
Protoconch unknown; at least 6 postnatal whorls. First whorl with 12-13
well-defined rounded lamellae; second to fifth whorls each with 15 lamellae; body
whorl with 14 lamellae. At least 20 very fine spiral lines visible between lamellae,
becoming obscure near upper and lower sutures, and difficult to count. Strong
basal lira on body whorl. Suture crenulate. Outer lip varicoid.
Remarks
The present material resembles the living east-coast species E. coronatum
(Lamarck, 1816) in the possession of a basal lira, but is more slender. The
common Epitonium of the west coast, E. kraussi (Nyst), lacks both spiral lirae
and a basal lira. Epitonium lamellosum (Lamarck, 1822) of West and South
Africa is a somewhat broader shell, with fewer lamellae (9-10 on body whorl),
and lacks distinct spiral sculpture.
Etymology
The specific name, from the Latinized Greek meaning 'wolf head', alludes to
a low hill or koppie, Wolfkop, in the vicinity of Hondeklip Bay.
Family Thaididae
Nucella praecingulata (Haughton, 1932)
Figs 17, 18
Thais praecingulata Haughton, 1932: 48, pI. 5 (figs 6-10). Barnard, 1958: 219; 1962: 182.
Material
Syntypes. SAM-9730, 3 specimens, Reuning's Claim, Alexander Bay, Cape
Province, coli. J. B. Cilliers. SAM-10598, 1 internal mould, The Point,
Namaqualand coast, Cape Province, oyster horizon, coli. E. Reuning.
SAM-9731, 4 specimens, Alexander Bay; SAM-9742, 1 specimen; SAM-9743,
1 specimen; SAM-9745, 1 specimen; SAM-9746, 1 specimen; SAM-9749,
1 specimen; SAM-9750, 1 specimen; SAM-9752, 1 specimen; SAM-9753,
1 specimen; SAM-9754, 1 specimen; SAM-9934, 7 specimens; all previous
specimens from Graauweduinen, Vanrhynsdorp District coast, Cape Province,
unless otherwise stated, coIl. S. Haughton.
Non-type material. SAM-PQ-AV398, 18 specimens, 18-58 mm,
Avontuur A, 50 m Complex. SAM-PQ-HB344, 28 specimens, 16-57 mm,
Hondeklip Zone 12. SAM-PQ-HB334, 5 specimens, 81-110 mm, Hondeklip
Zone 12. SAM-PQ-KN460, 2 specimens, 103 mm, 110 mm, Koingnaas, 50 m
Complex.
Description
Shell thaidid-like, variable, with aperture longer than spire. Protoconch of
I-H whorls, apparently smooth (but all specimens apically somewhat eroded);
4+ postnatal whorls, generally with well-marked shoulder. Spire in relation to
body whorl variable, squat in some specimens and half length of aperture, to
more slender and three-fourths length of aperture. Sculpture variable, squatter
specimens generally with stronger sculpture; early whorls with 4 strong spiral
bands and numerous axial lamellae between bands and upper and lower suture
lines, giving cancellate appearance. Spiral bands increasing on to body whorl,
with intermediaries appearing in some specimens; up to 20 major spiral bands,
running on to base. Cancellation usually lost on body whorl. Specimens having
weaker spiral sculpture usually with shoulder marked by single somewhat
stronger band; sculpture in these specimens often becoming obsolete on body
whorl. Columella smooth, concave in lower half. Base rounded, fasclole strong.
Inner surface of outer lip generally plicate, folds becoming obsolete in larger
specimens.
Remarks
Barnard (1962: 182) noted that Nucella praecingulata resembles both Nucella
cingulata (Linnaeus, 1758) and N. squamosa (Lamarck, 1816), but reaches a
much greater size than either of these living species.
Haughton (1932) quotes a note from Tomlin, in which the latter suggests the
name 'praecingulata', and also observes the similarity to N. cingulata and
N. squamosa. Haughton, while not presenting a formal description, noted the
variability of the species, did not designate a holotype, but sawall the material
listed above under 'Syntypes' (Fig. 17).
In spite of being highly variable, large specimens of N. praecingulata
(approx. 100 mm) are easily identified, being the only whelk of such dimensions
in the west-coast deposits. Smaller specimens, in the size range of N. cingulata
and N. squamosa, however, are more difficult to assign.
In the majority of smaller specimens of N. praecingulata the body whorl and
at least the two earlier whorls have a faint but distinct shoulder on the upper part
of the whorl. The strength of the spiral bands varies. In those specimens in which
the spiral bands are few and strong, the resemblance to N. cingulata is
unmistakable, even to the possession of five axial lines between the spiral bands,
leading to the squamose appearance sometimes seen in N. cingulata. In these
specimens, the whorls tend to have an evenly rounded profile. In those specimens
in which the spiral bands are finer and more numerous, the axial lines lead to a
cancellate appearance very similar to N. squamosa (see Fig. 19). In these, the
slightly angular whorl profile is more apparent, but even here, evenly rounded
whorls are encountered.
It is suggested that with the change to a colder regime on the west coast
during the Pleistocene, Nucella praecingulata gave rise to two forms, Nucella
cingulata and N. squamosa, both being intertidal to shallow infratidal forms,
neither attaining the dimensions of the ancestor, and with N. cingulata being
confined to the colder waters of the west coast to False Bay, while N. squamosa,
perhaps with greater temperature tolerance, extends to the Transkei coast.
Nucella praecingulata has not been recorded concurrently with N. cingulata
or N. squamosa, a further indication of the form's possible ancestral role.



Fig. 18. Nucella praecingulata. A. AvontuiJr specimens. Scale = 30 mm. B. Hondeklip specimens. Note oyster shells on lower left specimen.
Scale = 30 mm.

Thais arenae sp. novo
Fig. 20
Material
Holotype. SAM-PQ-KN530, almost complete specimen, length 52,5 mm,
greatest width 27,3 mm, Koingnaas KL south face.
Non-type material. SAM-PQ-KN307, 3 fragments (body whorls, somewhat
worn), Koingnaas KL south face.
Description
Shell with at least 4 postnatal whorls; aperture longer than spire. (The
holotype, an almost complete specimen, lacks the tip of the spire.) Whorls with
distinct shoulder in upper half. Columella smooth; short anterior canal flexed
slightly to left. Second extant whorl with 14 axial ribs, spiral sculpture worn away.
Third whorl with 11-12 axial ribs, 2 strong spiral keels, upper forming shoulder;
axial ribs rounded at intersections with strong keels; fine spiral sculpture over
entire whorl, 11-12Iirae between suture and shoulder, 9-10 below shoulder.
Body whorl with 10 axial ribs, becoming obsolete on outer lip; 5 strong keel-like
spiral bands; 12-13 fine lirae between suture and shoulder; 10-12 fine lirae
between each strong spiral band; 4 fine lirae between lowest band and base.
Remarks
The present species bears some resemblance to Thais capensis (Petit, 1852),
especially in the fine spiral lirae, but the latter is a relatively squatter species
having somewhat stronger tubercles, especially on the body whorl. Thais
haemastoma (Linnaeus, 1767), especially in the form occurring at present at the
southern end of its West African range, has a much squatter shell bearing variable
rounded tubercles on the shoulder, rather than the rounded ribs of T. arenae.
Lathyrus armatus A. Adams (see Nickles 1950: 106) from the Azores, the
Canary Islands, and Senegal, has a similar overall shape and sculpture. Although
the holotype of T. arenae shows some wear on the axial ridges and tubercles,
these are still more rounded than in the West African species. Also, the anterior
canal is more defined in the latter species.
Etymology
The specific name, from the Latin, of a sandy place, refers to the coarse
sands of the type-locality at Koingnaas KL south face.
Family Muricidae
Ocenebra bonaccorsii (Carrington & Kensley, 1969)
Tritonalia bonaccorsii Carrington & Kensley, 1969: 196, pI. 19.
Material
SAM-PQ-HB531, 39,1 x 22,0 mm, Hondeklip A Block, 30 m Complex.
SAM-PQ-HB532, 2 specimens, 8,4 x 4,8 mm, 8,1 x 4,0 mm, Hondeklip
Zone 12, 50 m Complex. SAM-PQ-HB152, 22,1 x 10,8 mm (protoconch miss-
ing), Hondeklip Zone 3.
Remarks
The PQ-HB531 specimen above is both larger and better preserved than the
holotype. The description of the species can thus be supplemented:
Protoconch of 2-2! whorls; 4! postnatal whorls. First two postnatal whorls
bicarinate; third whorl with upper carina stronger than lower; postnatal whorl 2
with 4 fine spiral lirae between suture and upper carina, 2-3 fine lirae between
upper and lower carina; 2-3 lirae between lower carina and suture. Postnatal
whorls 3 and 4 with 8-9 lirae between upper suture and upper carina; 4-5 lirae
between carinae; 4 lirae between lower carina and lower suture. Body whorl with
13-15Iirae between upper suture and strongest spiral ridge; 8 strong spiral
ridges, 9 fine lirae between uppermost strong ridge and next, number of fine lirae
decreasing anteriorly to two or three between stronger ridges. Inner surface of
outer lip with 8 rounded ridge-teeth.
Ocenebra petrocyon sp. novo
Fig. 21
Material
Holotype. SAM-PQ-HB533, 9,9 x 4,8 mm, Hondeklip, 30 m Complex.
Paratypes. SAM-PQ-HB534, 10,6 x 5,7 mm, 8,6 x 4,6 mm; Hondeklip,
30 m Complex. SAM-PQ-HB535, 5 damaged specimens, 20 juvenile speci-
mens; Hondeklip A Block, 30 m Complex.
Fig. 21. Ocenebra petrocyon. Holotype at left, remainder of specimens paratypes.
Scale = 3 mm.
Description
Protoconch of H smooth whorls; 4 postnatal whorls. Aperture subequal to,
or slightly longer than, spire. Sculpture consisting of rounded axial ribs and spiral
lirae. Axial ribs: 11-12 on first whorl; 11-12 on second whorl; 11 on third whorl;
11 on body whorl. Spiral lirae: 4 on first whorl; 5 on second whorl; 5 on third
whorl; 5 (with 1 or 2 very fine intermediates) on fourth whorl; 18-20 on body
whorl (outer lip), including several finer intermediates; lirae running on to
anterior canal. Lirae somewhat broadened at intersection with axial ribs, but
never becoming bead-like. Narrow, smooth columella callus present. Inner
surface of outer lip with 8-10 faint spiral ribs.
Remarks
Ocenebra petrocyon most closely resembles O. purpuroides (Reeve, 1845)
(with which it was collected), especially in general shape and body-whorl
sculpture. The new species differs from O. purpuroides in that the upper two or
three whorls are not cancellate; in having more spirallirae per whorl; in having
spiral lirae evenly spaced between the upper and lower suture lines (0. pur-
puroides has a distinct shoulder and no lirae between shoulder and the upper
suture line); and in having the axial ribs more rounded (almost carinate in
O. purpuroides).
Etymology
The specific name is the Latinized form of the Greek for 'dog stone', and
alludes to the farm Hondeklip.
Trophon carringtoni sp. novo
Fig. 22
Latiaxis sp. Carrington & Kensley, 1969: 195, pI. 19.
Material
Holotype. SAM-PQ-A V536, 42,5 x 22,5 mm, Avontuur A, 50 m Complex.
Paratypes. SAM-PQ-AV537, 2 specimens, 37,lx22,4mm, 40,8x
Fig. 22. Trophon carringtoni. A. Holotype. Scale = 10 mm. B. Range of paratypes.
Scale = 30 mm.
24,5 mm, Avontuur A, 50 m Complex. SAM-KI447, 62,5 X 37,0 mm, outer lip
missing, Strandfontein, 45-50 m Complex, coIl. A. J. Carrington.
SAM-PQ-AV538, 8 specimens (all with eroded apices), 58,0 x 32,0 mm,
53,0 x 28,0 mm, 42,5 x 23,5 mm, 40,2 x 24,0 mm, 40,0 x 23,5 mm, 38,2 x
22,0 mm, 34,8 x 20,0 mm, 20,0 x 13,0 mm, Avontuur A, 50 m Complex.
USNM 400986, 3 specimens (all with eroded apices), 42,0 x 24,8 mm,
43,0 x 23,3 mm, 38,0 x 25,9 mm, Avontuur A, 50 m Complex.
Description
Shell of 4-5 postnatal whorls; aperture longer than spire. Whorls with single
prominent keeled shoulder. Early whorls with irregular axial ridges, giving
appearance of cancellate sculpture below keel, forming more or less well defined
squamae where keel intersected. In later whorls, 6-8 axial ridges only faintly
indicated above keel, stronger below keel, especially on body whorl, forming
faint swelling at intersection with keel. Columella smooth, with callus developed.
Inner surface of outer lip with 5 tubercles below carinal notch. Short anterior
canal flexed to left.
Remarks
In the original mention of this species (Carrington & Kensley 1969: 195) the
resemblance to Latiaxis tortilis (= L. nakamigawai io Kilburn, 1974) was
mentioned. This comparison no longer stands, as the present species is more
slender, does not have a strongly flexed anterior canal, and appears to lack spiral
sculpture, other than the carina.
The general shape and sculpture are reminiscent of members of the
Trophoninae, but resemble nothing recorded in the southern African fauna.
Etymology
The species is named for Mr A. J. Carrington of De Beers Consolidated
Diamond Mines.
Family Buccinidae
Burnupena rogersi sp. novo
Fig. 23
Material
Holotype. SAM-PQ-HB539, 58,1 x 34,5 mm (body whorl plus 2! whorls,
remainder of spire missing), Hondeklip Zone 3, 50 m Complex.
Paratypes. SAM-PQ-HB540, 14 specimens, 48,3 x 30,5 mm to 14,7 x
8,7 mm (larger specimens with outer lip and/or body whorl damaged and spire
missing; smaller specimens complete). Hondeklip Zone 3, 50 m Complex.
USNM 400987, 5 specimens, 38,4 x 24,3 mm to 15,7 x 8,9 mm, Hondeklip
Zone 3,50 m Complex.
Fig. 23. Burnupena rogersi. Holotype at upper left corner, remainder of specimens
paratypes. Scale = 20 mm.
Description
Shell about 1,5 times longer than wide. Protoconch unknown; at least
5 postnatal whorls. Profile of earlier whorls evenly convex; of last 3 whorls with
distinct, rounded shoulder. Sculpture of fine spiral bands separated by narrow
incised grooves; 7-8 on third from last whorl, 14-15 on penultimate whorl, 35-42
on body whorl, extending on to outer surface of anterior canal, flattened bands
becoming obscure, divided by shallow incised lines on lower body whorl. No axial
sculpture apart from faint growth lines, and several aperture margin ridges in
larger specimens. Narrow gutter formed at suture line on last whorl, becoming
closed on earlier whorls. Inner surface of outer lip with about 10 faint ridges.
Base narrowly umbilicate; columella lacking pleats; with weak callus; anterior
canal short, flexed slightly to left.
Remarks
The present material, showing a strongly sunken suture, bears some
resemblance to the genus Babylonia. Burnupena rogersi, however, is plumper
than the two South African species of Babylonia, neither of which possesses fine
spiral sculpture.
While the generic position of the present material cannot be established in
the absence of radula, soft parts, and operculum, the species does bear a general
resemblance to species of Burnupena, and especially to B. papyracea (Bruguiere,
1789), which sometimes does show a fairly sunken suture.
Etymology
The species is named for Dr John Rogers of the Department of Geology,
University of Cape Town.
Burnupena aestus sp. novo
Figs 24,25
Material
Holotype. SAM-PQ-AV541, 69,9 x 31,0 mm, Avontuur A, 50 m Complex.
Paratypes. SAM-PQ-A V542, 3 specimens, 62,4 x 30,3 mm (apex and outer
lip damaged), 67,1 x 29,9 mm, 68,4 x 32,0 mm, Avontuur A, 50 m Complex.
SAM-PQ-HB543, ?66,6 x 29,7 mm (apex damaged), Hondeklip Zone 12,50 m
Complex. SAM-KI445. 61,3 x 27,0 mm (body whorl damaged), Swartlintjies,
colI. A. J. Carrington, 1967. SAM-KI446, 67,1 x 34,6 mm (body whorl dam-
aged), Swartlintjies, call. A. J. Carrington, 1967.
Description
Shell elongate, relatively slender, spire longer than aperture. At least
5 postnatal whorls. Profile of whorls convex, with slight hollowing just below
suture. Columella smooth, basally flexed slightly to left, with rounded ridge just
below suture; base rounded. Inner surface of outer lip with 20-22 rounded ridges.
Sculpture consisting of fine spiral lirae, occasionally doubled on lower whorls;
spiral lirae narrowly rounded to carinate. Second whorl with 10-12 lirae; third
with 12-15; fourth with about 20; body whorl with up to 56 lirae.
Remarks
In general proportions, the present species most resembles Afrocominella
capensis (Dunker, 1844), but lacks the obscure axial sculpture of the early whorls
typical of this species. The spiral lines are finer and more numerous than in
A. capensis, while the aperture is generally subequal in length to the spire. The
present species attains a much larger size than either A. capensis or Burnupena
papyracea, the other species to which it bears some similarity. This latter species
is much squatter, with fewer spiral lines.
There is a possibility of confusing small specimens of this species with large
specimens of Triumphis dilemma (see below). The two species may, however, be
easily separated on the following features: columella smooth in B. aestus, with
2-4 tubercles in T. dilemma; aperture longer than spire in T. dilemma, shorter
than spire in B. aestus; outer lip frequently thickened in T. dilemma, unthickened
in B. aestus; spiral cords on body whorl and anterior canal often doubled,
normally rounded to carinate in B. aestus, spiral cords single, rounded to tabulate
in T. dilemma.
Etymology
The specific name is derived from the Latin 'aestus', of the surf, and refers to
the probable habitat of the species.
Triumphis dilemma Kilburn & Tankard, 1975
Fig. 26
Triumphis dilemma Kilburn & Tankard, 1975: 200, fig. 10.
Material
Holotype. SAM-K4565, 16,3 x 10,4 mm, Langebaan, Early Pleistocene,
coli. A. J. Tankard.
Non-type material. SAM-PQ-HB544, 10 specimens, 27,2 x 17,8 mm to
39,1 x 22,0 mm, Hondeklip Zone 4A, 50 m Complex. SAM-PQ-AV545,
6 specimens, 25,3 x 15,0 mm to 33,0 x 19,9 mm, Avontuur A, 50 m Complex.
SAM-K4862, 4 specimens (2 damaged), 24,9 x 13,2 mm, 30,9 x 16,3 mm, Som-
naas SN4, Hondeklip Bay, colI. A. J. Tankard.
Previous records
Early Pleistocene, Langebaan, 9,5 m beach.
Remarks
With a wider range of material available, the original description can be
supplemented: parietal area with single tooth in most cases; 2 teeth (as in
holotype) only very occasionally seen. Outer lip only occasionally not thickened;
with 10-17 internal ridges. Columella with 2-5 basal tubercles, 2 most common.
Spiral cords rounded to tabulate. Aperture longer than spire, generally not as
constricted as in holotype. Kilburn & Tankard (1975) mentioned two damaged
specimens considerably larger than the holotype, about which they had
reservations. With good material in this size range, it can be seen that the two
earlier specimens are in fact this species, and that the higher counts of spiral cords
are typical of the species.
Family Nassariidae
Bullia annulata (Lamarck, 1816)
Bullia annulata: Barnard, 1959: 127, figs 25f, 27e; 1962: 189.
Bullia magna Haughton, 1932: 46, pI. 5 (figs 1, 4, 5). Barnard, 1959: 129; 1962: 182, 189.
Material
SAM-9897 (syntypes of B. magna), 2 incomplete specimens, Graauwe-
duinen, Vanrhynsdorp District coast, Cape Province. SAM-7998, (syntypes of
B. magna), 16 incomplete specimens, Doornbaai, south of Olifants River mouth,
Fig. 26. Triumphis dilemma. Range of material from Hondeklip, 50 m
Complex. Scale = 30 mm.
Cape Province. SAM-K4863, 1 specimen, 67,6 x 32,2 mm, Swartlintjies SL2,
Hondeklip Bay, colI. A. J. Tankard. SAM-PQ-HB210, 3 specimens,
63,7 x 30,1 mm, 68,7 x 32,2 mm, 55,6 x? mm (body whorl incomplete), Honde-
klip Zone 12, 50 m Complex. SAM-PQ-AV233, 7 specimens (3 incomplete),
64,3 x 30,7 mm, 60,8 x 30,8 mm, 59,9 x 30,3 mm, 51,2 x 26,9 mm, Avontuur,
50 m Complex. SAM-PQ-AV588, 12 specimens, Avontuur, 50 m Complex.
SAM-PQ-HB613, 1 specimen, Hondeklip Zone 4A, 50 m Complex.
SAM-PQ-HB889, 2 specimens, Hondeklip Zone 12, 50 m Complex.
Remarks
Barnard (1962) expressed the view that Bullia magna was merely a worn and
fossilized B. annulata. All the syntypic material is extremely worn, only one
specimen showing faint spiral sculpture.
The present material, while friable, shows good preservation of detail, and
possesses the distinctive spiral lirae characteristic of B. annulata.
While only one of the specimens from the 50 m Complex shows the
development of a shoulder on the upper whorl, approaching the condition in
typical B. annulata, Barnard (1962) does note that some recent specimens show
only a weak development of the shoulder. The present material removes any
doubt that B. magna really is B. annulata.
The Hondeklip specimens occurred in a very fine sand, in a layer dominated
by specimens of the bivalve Dosinia sicarisinus sp. nov., often with both valves in
life position.
Distribution
Living B. annulata occurs from Saldanha Bay to Mozambique, from the low-
tide mark, but more usually infratidally, to 100 m (Barnard 1958; Kilburn &
Rippey 1982).
Fossil B. annulata has been recorded from Langebaan, Milnerton, Swart-
kops, Redhouse and Coega (Barnard 1959, 1962).
Nassarius kochianus (Dunker), Kilburn & Rippey, 1982: 100,213, pI. 23 (fig. 8).
Nassa kochiana: Barnard, 1959: 104, figs 22b, 23b; 1962: 189.
Material
SAM-PQ-KN546, 3 specimens, 10,9 x 6,1 mm, 9,1 x 5,1 mm, 7,6 x
4,1 mm, Koingnaas KL south face.
Previous records
Living: False Bay to Transkei, intertidal to shallow infratidal.
Fossil: Algoa Bay.
Remarks
With slightly fewer axial ribs (13-14 per whorl) than is usual for
N. kochianus (15-18), the present material, however, does have only a single
columella nodule, and in profile more closely resembles this species than it does
N. signatus (Dunker, 1847). More specimens are needed to decide the range of
variation in the number of axial ribs, before a more definite identification can be
attempted.
Family Fasciolariidae
Fasciolaria dinglei sp. novo
Figs 27,28
Material
Holotype. SAM-PQ-AV547, 101 x 41 mm (apex and upper whorls eroded),
Avontuur A, 50 m Complex.
Paratypes. SAM-PQ-A V548, 3 body whorl fragments, 7 specimens,
39,4 x 19,8 mm to 83,4 x 40,3 mm, Avontuur A, 50 m Complex.
Description
Shell with aperture longer than spire. Four to five postnatal whorls. Profile of
whorls evenly convex. Third whorl with 12-13 fine spiral lirae; fourth with 15-
16 lirae; body whorl with 50-55 lirae running on to anterior canal; lirae on upper
'. ;,
\)1
body whorl near suture having finer intermediates appearing; lirae on base of
body whorl becoming coarser and broader. Columella with 3 pleats, sometimes
obscured in large specimens; low ridge on upper columella just below suture.
Anterior canal about half length of rest of aperture, flexed to left. Inner surface of
outer lip with up to 25 rounded ridges.
Remarks
Of the two specimens of Fascialaria figured by Carrington & Kensley (1969,
pI. 18c), the smaller may well be the present species. The larger, however, is
undoubtedly a large Burnupena, lacking as it does an elongate anterior canal.
Carrington & Kensley (1969) noted the similarity of their specimen to
Fascialaria lugubris (Reeve, 1847). This similarity in overall proportions and
general sculpture is again noted, but several differences easily separate the two
species. The most distinctive feature of the new species is the presence of three
pleats on the columella (F. lugubris is unpleated). The spiral sculpture of
F. dinglei is much finer and the lines more numerous (up to 55 on the body whorl)
than in F. lugubris (up to 30 on body whorl, and more raised).
Fig. 28. Fascialaria dinglei. A. Range of paratypes. Scale = 10 mm. B. Range of paratypes.
Scale = 30 mm.
Etymology
The species is named for Professor Richard V. Dingle, Department of
Geology, University of Cape Town.
Family Olividae
Melapium hawthornei sp. novo
Fig. 29
Material
Holotype. SAM-PQ-KN549, 27,5 x 24,0 mm, Koingnaas KN-l.
Paratypes. SAM-PQ-KN550, 12 specimens, 23,4 x 16,7 mm to 26,1 x
20,1 mm, Koingnaas KN-1, 50 m Complex. USNM 400988, 4 specimens,
23,4 x 17,1 mm to 27,0 x 18,9 mm, Koingnaas KN-1, 50 m Complex.
Non-type material. SAM-PQ-KN551, 9 specimens, all worn and damaged,
Koingnaas KL south face. SAM-PQ-KN366, 16 specimens, all damaged,
Koingnaas KN-1, 50 m Complex.
Description
Shell 1,2 times longer than wide, of 4 postnatal whorls. Spire somewhat
sunken, but still protruding. Body whorl globose, profile evenly convex, with very
faint, shallow, spiral lines, and irregular axial growth lines. Parietal callus
extending from body-whorl suture in evenly convex line to lower columella.
Posterior canal short, but with distinct notch; anterior canal flexed to left; lower
columella narrowed, with fasciole strong but barely visible in apertural view;
outer lip evenly convex to notch at anterior canal.
Remarks
Two living and one fossil species of Melapium have been recorded from
southern Africa, all of which differ from the present species.
Melapium lineatum (Lamarck, 1822), known from Still Bay to Zululand in
depths of 30-160 m, is of comparable size to M. hawthornei, but possesses a more
elevated spire; also, the lower columella is broader, with the fasciole being strong
and visible in apertural view. Melapium lineatum also has a distinct concavity just
above the fasciole origin, and below the bulge of the body whorl; M. hawthornei,
by contrast, is evenly convex in this region.
Melapium elatum (Schubert & Wagner, 1829), from deeper water off Natal
and Mozambique, has a much larger shell, with the lower columella very broad
and with a very strong fasciole, and a distinct shoulder on the upper body whorl.
Melapium patersonae Bullen Newton, 1913, from the Neogene of Bredas-
dorp to Alexandria, Cape Province, is also a large shell, up to 70 mm in length,
has a sunken spire, a well-developed posterior canal with the outer lip reaching
posteriorly above the spire, and a very broad columella with a very strong
carinate fasciole.

Etymology
The species is named for Mr J. B. Hawthorne of De Beers Consolidated
Diamond Mines.
Pseudoliva lutulenta sp. nov.
Fig. 30
Material
Holotype. SAM-PQ-KN552, 31,6 x 22,7 mm, Koingnaas KN-l.
Paratypes. SAM-PQ-KN553, 8 complete specimens, 20,7 x 15,8 mm to
29,2 x 21,2 mm, Koingnaas KN-l. USNM 400989, 3 complete specimens,
23,1 x 18,1 mm to 28,3 x 21,5 mm, Koingnaas KN-l.
Non-type material. SAM-PQ-KN373, 13 specimens (damaged), Koingnaas
KN-l.
Description
Shel1 globose, thick-wal1ed, longer than wide, of 4 whorls. Spire short. Whorl
profiles evenly convex; suture somewhat incised. Columel1a cal1us smooth,
convex, reaching posteriorly to level of suture, outline evenly convex on inner-lip
body whorl; parietal node somewhat elongate, forming shal10w groove-like
posterior canal. Base with strong furrow running from shallow sinus in anterior
outer lip, around lower body whorl, to midpoint of columel1a cal1us; second, less
clearly defined groove anterior to strong groove, neither groove marked on inner
surface of outer lip.
Remarks
Three major differences separate the present material from the apparently
very similar living Pseudoliva crassa (Gmelin), known only from Angola, and
figured and described as P. plumbea (Chemnitz) by Nickles (1950: 107, fig. 189).
The living species possesses two strong well-defined grooves on the anterior body
whorl, the upper of which is marked by a narrow 'excroissance' on the internal
surface of the outer lip. In the present species there is one strong groove and a far
less wel1-defined more anterior groove. Neither groove is reflected by any
structure on the internal surface of the outer lip. The columel1a callus in P. crassa
is concave, and posteriorly narrowed to the posterior canal. In P. lutulenta the
columella callus is convex, and posteriorly broadly rounded.
Bohm (1926) described two species of Pseudoliva from the Cretaceous of
Bogenfels, South West Africa-Namibia, neither of which bears much resem-
blance to the present species. Pseudoliva thielei Bohm shows enormous columella
callus development, and has several spiral ridges on the shell base; P. leutweini
B6hm is a much larger species than P. lutulenta, with a massive upper columella
cal1us.

Etymology
The specific name is derived from the Latin 'lutulentus', covered with mud,
and alludes to the fine sediments in which the species was found.
Family Turridae
Drillia tempestae sp. novo
Fig. 31
Material
Holotype. SAM-PQ-HB554, 12,1 x 5,1 mm, Hondeklip Zone 3, 50 m
Complex.
Paratypes. SAM-PQ-HB555, 10 specimens, 6,3 x 3,4 mm to 19,4 x
7,7 mm, Hondeklip Zone 3, 50 m Complex. USNM 400990, 10 specimens,
6,3 x 2,0 mm to 16,4 x 7,2 mm, Hondeklip Zone 3, 50 m Complex.
Non-type material. SAM-PQ-SL260, 1 specimen, 19,9 x 8,4 mm, Swart-
lintjies River, spoil heap. SAM-PQ-HB556, approx. 50 specimens, Hondeklip
Zone 3, 50 m Complex.
Description
Protoconch of H smooth whorls; 6!- 7 postnatal whorls. Spire 1,3-1,5 times
length of aperture. Sculpture consisting of rounded axial ribs and fine spirallirae;
axial ribs slightly oblique, occasionally perpendicular, rounded, with distinct
shoulder well below upper suture line, reaching lower suture. Postnatal whorl 3
with 9 axial ribs, 6-9 spiral lirae; ribs increasing to 11 on body whorl, lirae to
10-18 on penultimate whorl; outer lip with 20-24 spirallirae running on to outer
surface of siphonal canal. Lirae equal in width on spire, becoming broader on
body whorl and siphonal canal. Anal sinus in outer lip shallow. Columella callus
smooth; anterior siphonal canal short, well marked.
Remarks
The present species resembles Drillia caffra (Smith, 1882) in overall
proportions, but differs in details. The earlier species has more axial ribs per
whorl (11-12 on early whorls, 15-18 on later whorls), more spiral lirae (6 on
early whorls, up to 30 on last whorl), and possesses a deep anal sinus.
As generic placement in the Turridae is so dependent on radula structure,
the present material is placed in Drillia purely because of an overall resemblance
to some species in that genus.
Etymology
The specific name, from the Latin for 'of a storm', alludes to the storm
conditions that probably caused some of the Hondeklip fossil accumulations.
Fig. 31. Drillia tempestae. A. Holotype. Scale = 2 mm. B. Range of paratypes.
Scale = 5 mm.
Material
Holotype. SAM-PQ-HB557, 21,4 x 6,5 mm, Hondeklip Zone 12, 50 m
Complex.
Paratypes. SAM-PQ-HB558, 10 specimens, 15,2 x 4,9 mm to
23,1 x 6,9 mm, Hondeklip Zone 12, 50 m Complex. USNM 400991, 10 speci-
mens, 15,7 x 5,6 mm to 23,1 x 6,4 mm, Hondeklip Zone 12, 50 m Complex.
Fig. 32. Terebra canisaxi. Holotype at upper left corner, remainder of specimens paratypes.
Scale = 10 mm.
Non-type material. SAM-PQ-HB559, 4 specimens, 14,6 x 4,9 mm to
17,7 x 5,7 mm, many fragments, Hondeklip Zone 12, 50 m Complex.
SAM-PQ-HB331, 30 specimens, Hondeklip Zone 12, 50 m Complex.
Description
Shell of protoconch of 2l smooth whorls, plus 7 postnatal whorls. Postnatal
whorl sculpture consisting of strong, rounded, axial ribs running from upper to
lower suture, sometimes alternating from one whorl to next, sometimes
continuous. Postnatal whorl 1 with 9 ribs, following 6 whorls each with 8-9 ribs,
body whorl with 9-10 ribs. Anterior canal short, open, flexed to left. Base
rounded. Columella lacking pleats, smooth; low fasciole present. Spiral groove
absent.
Remarks
This species closely resembles Terebra light/ooti Smith, 1903, recorded alive
from Saldanha Bay to the Agulhas Bank to a depth of 62 m. Barnard (1969: 596)
described this species as having eight postnatal whorls. Terebra light/ooti is a
marginally more slender species than T. canisaxi, with slightly more numerous
axial ribs (11-12 on last three whorls, as against 8-9 in T. canisaxi).
Etymology
The specific name is the Latinized form of 'dog stone', a direct translation of
the name of the type-locality, Hondeklip.
Class BIVALVIA
Family Glycymeridae
Glycymeris Julleri sp. novo
Figs 33,34
Material
Holotype. SAM-PQ-SL561, both valves, 58,0 x 61,5 mm, Swartlintjies
SL-20, 50 m Complex.
Paratypes. SAM-PQ-SL562, both valves, 38,0 x 40,5 mm, Swartlintjies
SL-20, 50 m Complex. SAM-PQ-KN560, right valve, 40,2 x 44,1 mm, Koing-
naas KN-l, 50 m Complex. SAM-PQ-KN563, 3 left valves, 25,0 x 26,8 mm,
36,2 x 40,0 mm, 46,3 x 46,0 mm (distorted), 1 right valve, 37,1 x? mm, Koing-
naas KN-l, 50 m Complex. USNM 400992, 2 left valves, 23,6 x 24,9 mm,
42,9 x? mm, 1 right valve, 34,8 x? mm, Koingnaas KN-l, 50 m Complex.
Description
All material in poor condition, with external surfaces exfoliating, and shape
frequently deformed. Shell slightly inequilateral, posterior margin slightly more
pointed than anterior margin; slightly wider than high. Sculpture of about
40 radiating, apparently flattened ribs, becoming obsolete in anterior and
Fig. 33. Glycymeris fulleri. Holotype, external and internal views of both valves.
Scale = 30 mm.
posterior part of shell. Ventral margin faintly crenulate in external view,
internally more strongly crenulate. Anterior and posterior adductor muscle scars?
situated on faint ledge. Hinge area with teeth discontinuous below umbo, divided
into two groups, each of 7-9 teeth.
Remarks
Glycymeris fulleri resembles the living southern African east-coast species
G. queketti (Sowerby, 1897) in having the muscle scars situated on faint ledges,
but is a smaller and more inequilateral species. Barnard (1962: 183) mentions two
specimens referred to G. queketti from Skulpfontein Point, Hondeklipbaai area,
by Krige (1927), but questioned the identification.
Glycymeris connollyi Tomlin, 1923, known living from Table Bay to Natal, is
smaller than the present species, and more trigonal in outline.
Glycymeris borgesi (Cox, 1946) (= G. africana Cox, 1939), known from the
Neogene of Ysterplaat, Cape Province (Tankard 1975a) as well as from the
Alexandria Formation in the Port Elizabeth area, is a much larger species (up to
100 mm diameter), with the hinge teeth in a continuous band.
Glycymeris ovata (Broderip), recorded from the Pliocene and Pleistocene of
Chile by Herm (1969) also shows the tooth row of the hinge divided into two
parts, but this species is less markedly inequilateral and somewhat more trigonal
than the present species.
Etymology
The species is named for Professor A. O. Fuller, Department of Geology,
University of Cape Town.
Family Ostreidae
Ostrea ct. subradiosa Bbhm
Fig. 35
Ostrea digitilina, non Dubois, B6hm & Weisfermel, 1913: 61, fig. la, pI. 8 (fig. 2).
Ostrea subradiosa B6hm, 1926: 56, text table A (figs 1-3).
Material
SAM-PQ-AV571, one left valve, 93 x 83 mm, Avontuur A, 50 m Complex.
Remarks
The present specimen has been compared with material in the South African
Museum (SAM-K4936) from Bogenfels, as well as with the original descriptions
and figures. Neither of the figures of Bbhm & Weisfermel (1913) shows a row of
pits in the shell margin close to the hinge area. The figure of Bbhm (1926),
however, shows such a row of pits. The present specimen, while having the strong
radiating external ribs seen in O. subradiosa, lacks these marginal pits. A more
definite identification on the basis of a single specimen is not possible.
Family Isognomonidae
Isognomon gariesensis sp. novo
Fig.36A
Material
Holotype. SAM-PQ-HB564, incomplete right valve, upper (dorsal) part of
valve missing, greatest length 158 mm, greatest width (across hinge area)
90,9 mm, Hondeklip A Block, 50 m Complex.
Paratypes. SAM-PQ-HB263, 1 incomplete left valve, Hondeklip A Block,
50 m Complex. SAM-PQ-KN565, 2 incomplete left valves (one 145 mm in
length), Koingnaas KL south face.
Non-type material. SAM-PQ-KN368, numerous friable fragments, Koing-
naas KN-l.
Description
Shell very thick (up to 40 mm on ventral margin), becoming thinner
posterodorsally. Hinge with 11 elongate ligamental grooves; hinge width
36,5 mm.
Remarks
The genus Isognomon is largely a warm-water form. On the west African
coast, living Isognomon occurs as far south as the Congo (Nickles 1950: 172). On
the east African coast, I. anomioides (Reeve, 1858) commonly occurs in rock
pools' as far south as the Transkei coast. Isognomon perna occurs occasionally in
Natal. Both these latter species reach a total length of about 66 mm and are thin-
shelled.
Isognomon ct. gaudichaudi (d'Orbigny, 1842) (from the Miocene of Chile)
was recorded from Needs Camp, Cape Province (Woods 1908; Newton 1913) and
originally was thought to be of Cretaceous age, but later was referred to the
Cenozoic. The species has also been recorded from Redhouse, Koega, and
Bushmans River, Cape Province. The specimens examined in the South African
Museum are from Redhouse and Swartkops (Fig. 36B). Newton (1913) compa(ed
the Cape Province specimens with I. maxillata (Lamarck, 1801) from Europe,
1. conradi (d'Orbigny, 1842) from Virginia, U.S.A., and 1. gaudichaudi
(d'Orbigny) from Chile, and on the basis of the hinge similarities decided that the
South African specimens most closely resembled the South American species.
The present west-coast material is similar to the material from the Eastern
Province in hinge structure, but is much thicker and heavier. The ventral marginal
sinuosity of the Isognomon ct. gaudichaudi is more marked than in the Hondeklip
material. While it is unlikely that either the present samples or the Eastern
Province material is conspecific with the Chilean species, there is precedent for
this distribution pattern. Kensley & Penrith (1970) recorded three species of
mytilid bivalve molluscs, and one brachiopod species from northern South West
Fig. 36. A. Isognomon gariesensis. Holotype. B. Isognomon 'gaudichaudi'. Three specimens
from Redhollse, Cape Province. Scales = 10 mm.
Africa-Namibia and Angola that occur in Pacific South America, while Kensley
(1985b) recorded the South American thaidid Concholepas as a fossil from the
west coast of South Africa. Until more complete material is available, it is wiser
to give the present material new specific status, rather than confuse it with
earlier-described species.
Etymology
The specific name is derived from Garies, the district in which the Hondeklip
and Koingnaas localities are found.
Family Carditidae
Cardita unica sp. novo
Fig. 37
Material
Holotype. SAM-PQ-KN566, 1 right valve, 44,0 x 67,5 mm, Koingnaas KL
south face.
Description
Shell robust, thick, oblong, anterior margin rounded, ventral margin
flattened, posterior margin more pointed than anterior. External sculpture
consisting of 27 radiating ribs, becoming broadly flattened near ventral margin,
separated by narrow grooves; ribs on posterior part of shell narrower than on rest
of shell. Concentric growth lines irregular, wavy. Lower anterior and posterior
margin and ventral margin on shell interior crenulate, crenulations largest at
posteroventral corner. Anterior adductor scar oval; small circular hollow dorsal
to anterior muscle scar, hidden under anterior hinge line. Posterior adductor scar
anteriorly truncate, posteriorly rounded. Hinge line broad, solid; anterior lateral
tooth almost vertical, short, separated from anterior cardinal tooth by triangular
pit; anterior cardinal tooth acutely triangular with posterior margin twice length
of anterior margin, separated from posterior cardinal tooth by narrowly
triangular oblique groove; posterior cardinal tooth elongate, with horizontal
dorsal part and oblique ventral part. Ventral margin of hinge area sinuous.
Remarks
The present specimen bears no resemblance to any carditid, fossil or living,
recorded from southern or west Africa.
Etymology
The specific name, derived from the Latin 'unicus', meaning unique, refers to
the fact that only a single valve of this species has been found, and that it bears no
resemblance to any African carditid.
Fig. 37. Cardita unica. Holotype, external and
internal view of right valve. Scale = 10 mm.
Cuna aquaedulcensis Kensley, 1977
Fig. 38
Cuna aquaedulcensis Kensley, 1977: 203, fig. 15.
Material
SAM-PQ-HB567, about 50 valves, largest specimen 7,5 X 7,1 mm, Honde-
klip, 30 m Complex.
Description
Right valve, hinge with elongate anterior cardinal tooth, median tooth
narrowly triangular, separated from anterior tooth by narrow triangular gutter.
Fig. 38. Cuna aquaedulcensis. Hinge details
of left (upper) and right (lower) valves.
Posterior cardinal tooth reduced to short low rounded ridge. Left valve, hinge
with elongate rounded ridge on anterior margin, running from anterior adductor
muscle scar to umbo; 2 median teeth, anteriormost larger, smaller tooth narrow
and lower; posterior tooth separated from margin by relatively deep groove.
Previous records
Quartzose Sand Member, Langebaanweg, Pliocene.
Remarks
As only two right valves were available when the species was described, it
was thought useful to supplement the description, now that far more material is
available.
Family Veneridae
Dosinia (Dosinia) sicarisinus sp. novo
Figs 39,40
Material
Holotype. SAM-K4877, both valves, 75,7 x 72,4 mm, thickness of two
valves together 27,1 mm, Somnaas SN4, Hondeklip, 50 m Complex, colI. A. J.
Tankard.
Paratypes. SAM-PQ-HB568, 4 left valves, 66,7 x 65,5 mm, 63,1 x 65,0 mm,
57,4 x 57,7 mm, ? x 66,9 mm, 1 right valve, 67,4 x 70,7 mm, Hondeklip
Fig. 39. Dosinia sicarlSlnus. A. Holotype, external and internal
view of both valves. B. Paratype. Scale = 20 mm.
Fig. 40. Dosinia sicarisinus. Hinge details of right (upper) and
left (lower) valves.
Zone 12, 50 m Complex. SAM-PQ-HB569, both valves (hinge of right valve
missing), 62,8 x 68,0 mm, Hondeklip Zone 4A, 50 m Complex. USNM 400993,
both valves, 63,6 x 62,5 mm, 2 left valves, 66,3 x 64,5 mm, 61,8 x 63,2 mm,
Hondeklip, 50 m Complex.
Description
Shell proportions somewhat variable, generally longer than wide but
occasionally wider than long. Sculpture of concentric lines, barely lamellose
anteriorly and posteriorly. Lunule well defined; escutcheon lacking. Pallial sinus
horizontal to slightly ascending, apically narrowed. Left and right hinges with
small anterior lateral tooth set at angle to anterior cardinal tooth; posterior
cardinal tooth in right valve bifid; anterior cardinal tooth in left valve much less
markedly bifid, with very narrow longitudinal slit.
Remarks
The present material bears little resemblance to the extant species of Dosinia
from southern Africa. Apart from attaining a much larger size than any of the
living species, D. sicarisinus also differs in the very distinctive narrowly triangular
pallial sinus that reaches to below the umbo. Dosinia exoleta (Linnaeus) of the
Mediterranean, and also known from Norway to North Africa, has a broader
pallial sinus, and possesses lamellar concentric rings.
In general appearance and size, the present material resembles some of the
Tertiary species from New Zealand, especially D. (Raina) bartrami Laws, 1930.
This latter, however, has a shallow pallial sinus, and the shallow lunule of the
subgenus (see Keen 1969: N679). Dosinia sicarisinus in placed in the subgenus
Dosinia for lacking an escutcheon and lamellose concentric rings.
Etymology
The specific name is derived from the Latin 'sicarius', a murderer, and
'sinus', a bay, and alludes to Moordenaarsbaai, a coastal embayment close to the
Hondeklip type localities.
Family Pholadidae
Barnea truncata (Say, 1822)
Barnea truncata: Nickles, 1950: 232, fig. 454. Barnard, 1964: 565. Kilburn & Rippey, 1982: 203.
Kensley, 1985a: 116.
Material
SAM-PQ-KN356, 2 specimens, left and right valves, approx. 60 mm and
75 mm in length; right valve, approx. 50 mm in length, Koingnaas KN-l. SAM-
PQ-HB26, 3 worn hinges, Hondeklip, 30 m Complex.
Previous records
Living: Senegal to Angola; east coast of U.S.A.; numerous fresh dead shells
from Table Bay in South African Museum collection.
Fossil: Milnerton, Table Bay (Late Pleistocene); borehole in Kuiseb River,
South West Africa-Namibia, 9 miles (14,5 km) from sea (Geological Survey,
determined by K. H. Barnard).
Class SCAPHOPODA
Family Dentaliidae
Dentalium sp.
Fig. 41
Material
SAM-PQ-KN570, fragments, Koingnaas KN-l. SAM-PQ-KN376, several
fragments, Koingnaas KN-l.
Description
Shell gently convex; with 16 rounded ribs at narrower end (1,8 mm
diameter); ribs equal in width to concave furrows between them; at about 3,0 mm
diameter, much narrower intermediate rounded ribs (1 between 2 larger ribs)
becoming apparent; 26 ribs present.
Remarks
The present material bears little resemblance to any living species of
Dentalium described from southern Africa.
FAUNAL COMPOSITION
The following table is a broad-scale analysis of the faunal composition of the
fossil molluscs from the Hondeklip area collected during this study. Material
identified to generic level only is excluded (15 genera).
Total number of species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92
Total number of extinct species 45
Lyellian percentage (percentage extant) .
Total number of species still living on east and west coasts. . . . . . . 27
Total number of species living on east coast only. . . . . . . . . . . . . . . 11
Total number of species with West African-Mediterranean
affinities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
(49% )
51%
Extinct species also present in east-coast deposits .
Total number of species from 50 m Complex .
Total number of extinct species from 50 m Complex .
Lyellian percentage .
50 m Complex species recorded by Carrington & Kensley
(1969) but not found during course of this study .
(of which 3 are extinct)
Total number of species from 30 m Complex .
Total number of extinct species from 30 m Complex .
Lyellian percentage .
Number of species common to 50 m and 30 m Complexes .
Number of extinct species in common .
211
4
78
37 (47%)
53%
4
48
19 (40%)
60%
34
11
From the previous summary, several topics require further comment.
Notably, the 50 m Complex shows a higher diversity than the 30 m Complex
(78 species as against 48). A.glance at the species list (Table 1), which records the
depositional environment of the beds from which the molluscs were obtained,
shows that a greater range of environments was available in the 50 m Complex
(viz. back-barrier, tidal~inlet, and distal lower-shoreface deposits, as against only
near-shore open-coast deposits in the 30 m Complex). It is thus tempting to
attribute the higher diversity of the 50 m Complex to the wider range of habitats.
However, exclusion of specimens obtained only from 50 m Complex tidal-inlet
and back-barrier deposits from the species list removes only nine species (two
extinct), and the comparative diversity becomes 69 species (35 extinct) as against
48 species (19 extinct). This reflects the fact that the shelly beds in the 50 m
Complex back-barrier-related facies are transported assemblages consisting of a
mixture of calm-water and open-coast forms. Additionally, calm-water species
may also inhabit the substrate below average wave-base, and become incorpor-
ated into open-coast facies, provided the open-coast temperature regime is within
species tolerances.
As shown for the Late Pleistocene at Verlorevlei (Tankard 1975), thermally
anomalous back-barrier environments increase species-diversity by the addition
of warm-water taxa to the faunal composition. A modern example of such
localized habitats is Sandvis Lagoon (Sandwich Harbour) near Walvis Bay
(Kensley & Penrith 1977). However, in such cases the contrast between the
back-barrier and open-coast assemblages is clear-cut. This is evidently not the
case with the Hondeklip fauna, but this observation is qualified by the absence of
large life assemblages preserved in the 50 m Complex back-barrier facies.
In the 50 m Complex back-barrier facies, only Dosinia sicarisinus, Donax
haughtoni, Phaxas decipiens, Standella namaquensis, and Tivela ct. compressa
have been found in life positions. In open-coast facies, Lutraria sp. has been
found in life position around the lower-shoreface-upper-shoreface facies bound-
ary in the 50 m Complex; in the 30 m Complex, Donax rogersi and Standella
namaquensis occur in life positions in the upper shoreface. Striostrea margariticea
has been found attached to rocks in open-coast contexts in both complexes.
In Table 1 the species obtained from deposits on the properties of Koingnaas
and Swartlintjiesrivier have been indicated separately owing to the unique addi-
tions to the faunal list those beds have contributed. In the case of Koingnaas (KN)
and Swartlintjiesrivier (SL) samples, this is further warranted since the beds are
distal lower-shoreface deposits of the 50 m Complex. The curious mix of intertidal
and deeper-dwelling forms (e.g. Dentalium, Ringicula, Tugali, and Turritella
declivis) in these samples, together with the muddy nature of the enclosing
sediment, is consistent with a depositional environment transitional to offshore
conditions. As such, the KN sample has contributed most to environmentally-
based diversity in the 50 m Complex.
Removing species unique to these samples (KN, SL, and KL) from the
faunal list results in a 50 m-30 m Complex diversity contrast of 64 species
(30 extinct) as against 39 species (14 extinct), respectively. Excluding these
examples, as well as species from the 50 m Complex tidal-inlet and back-barrier
facies from the faunal list, results in a diversity contrast of 54 species (28 extinct)
as against 39 species (124 extinct) for the 50 m and 30 m Complexes, respect-
ively. Only by not excluding the KL sample from the latter calculation is the
diversity contrast narrowed: 54 species (50 m Complex) as against 48 (30 m
Complex). Thus the higher diversity of the 50 m Complex is either real, or an
unavoidable bias due to the nature of the available exposures. In addition, any
reworked forms possibly present in the 30 ill Complex will reduce the diversity
contrast.
TABLE 1
Species of Mollusca obtained from four properties in the Hondeklip area of the Namaqualand
coast, reflecting the complex and depositional facies from which they were obtained.
Habitat: R = Rocky, S = Sandy, RiS = Rock in sand, M = Muddy. W/E: Species confined to
either west (W) or east (E) coasts. TIL = Tidal-inlet facies. BBR = Back-barrier facies.
USH = Upper-shoreface facies. LSH = Lower-shoreface facies. DLS = Distal lower-shoreface
facies. KN = Koingnaas KN. KL = Koingnaas KL. SL = Swartlintjies SL. t = Extinct.
50 m Complex 30 m Complex
Habi- DLS LSHSpecies tat WIE TIL BBR USH LSH (KNfSL) USH LSH (KL)
GASTROPODA
Afrocominella capensis ...... .... R + + +
(Dunker in Philippi, 1844)
Amblychilepas scutellum ........ R + + + +(Gmelin, 1791)
tArgobuccinum casus sp. noY. 0 ??? + + +
tBolma anoropha sp. noy ..... .... +
Bullia annulata (Lamarck, 1816) .. S + +
Bullia digitalis (Dillwyn, 1817) ... S + + + + +
Bullia laevissima (Gmelin, 1791) S/M +
tBurnupenaaestussp. noy ........ + +
Burnupena papyracea .......... '. R + + +
(Bruguiere, 1789)
tBurnupenarogersisp. noy ....... +
tCaliiostoma depressa ........... +
Carrington & Kensley, 1969
Cantharidus (Jujubinus) striatus .. W +(Linnaeus, 1758)
Calyptraea helicoidea ........... R E +Sowerby, 1883
tCalyptraea kilburni nom. noy ..... ?R + + + + + + +
tCalyptraea viridarena ........... ?R + +
Carrington & Kensley, 1969
tClanculus lutosus sp. noy ........ +
tClanculus murrayi .............. + + + +Carrington & Kensley, 1969
Conus mozambicus ............. R + + + +Hwass in Bruguiere, 1789
Crepidula porcellana ............ R W + +Lamarck, 1801
tCrepidula deprima sp. noy ....... R + + +
Cylichna tubulosa Gould, 1859 ... E +
Diodora elevata (Dunker, 1846) .. R + + + + + + +
tDriliiatempestaesp. noy ......... + + +
tEpitonium lycocephalum s . noy. + +
50 m Complex 30 m Complex
Habi- DLS LSHSpecies tat WfE TIL BBR USH LSH (KN/SL) USH LSH (KL)
GASTROPODA (contd)
tFasciolaria dinglei sp. nov .. . - .... +
tFissurelia glarea ............ .... ?R + +
Carrington & Kensley, 1969
tFissurella robusta .. . ... . .... . ? . R + + + +
Sowerby, 1892
Fissurellidea aperta .. ........... R +
(Sowerby, 1825)
Fusus faurei .... ... . .... . ... . ? . ? ? ? ? ?
Barnard, 1959 (Koingnaas)
Gibbula zonata patula ssp. novo ? . R + +
tHaliotis saldanhae . . . .......... R + +
Kensley, 1972
Helcion sp. .... . . . . . . . . . . . . . . . . R +
tHespererato oppenheimeri ....... ? ? ? ? ? +
Carrington & Kensley, 1969
Littorina sp .... ...... . ... . ..... R + +
Marginella sp. ............ .... . + + +
Melanella sp ...... .... . ........ +
tMelapium hawthornei sp. novo ... + +
tNamamurex odontostoma .. . .... + + +
Carrington & Kensley, 1969
Nassarius kochianus .. .. . ....... E +
(Dunker, 1846)
tNassarius litorafontis ..... . ... ? ? ? ? ? +Carrington & Kensley,"1969
Natica d. adansoni .......... ... S/M W +Blainville, 1824
Nucella dubia (Kr~uss, 1848) .... R + +
tNucella praecingulata ........... + + + + + + +
(Haughton, 1932)
tOcenebra bonaccorsii . .......... + +
Carrington & Kensley, 1969
Ocenebra purpuroides ........ ? . R +
(Reeve, 1845)
tOcenebra petrocyon sp. novo 0 ???? +
Oxystele sinensis (Gmelin, 1791) R E + + + + + +
Patella argenvillei Krauss, 1848 R + + + +
Patella barbara Linnaeus, 1758 ... R + + + +
Patella granatina Linnaeus, 1758 .. R +
tPatelia hendeyi sp. novo 0 ???????? R + +
tPatelia hoffmani sp. novo .. ...... R + + + +
Patella miniata Born, 1778. ...... R + +
Patella sp ................ ...... + +
tPseudoliva lutulenta sp. nov .. .... +
Ringicula turtoni Bartsch, 1915 .. E +
50 m Complex 30 m Complex
Habi- DLS LSHSpecies tat WfE TIL BBR USH LSH (KN/SL) USH LSH (KL)
GASTROPODA (contd)
Sinum concavum (Lamarck, 1822) W +
tTerebra canisaxi sp. novo 0 ??? .... + + +
tThais arenae sp. novo .. ...... . +
Tricolia capensis (Dunker, 1846) R + +
tTriumphis dilemma .. ..... .. . + + +
Kilburn & Tankard, 1975
tTrophon carringtoni sp. novo +
Tugali barnardi (Tomlin, 1932) . .. E +
Turbo cidaris Gmelin, 1791 ...... R + + + + + + + +
Turritella carini/era . ....... . ?? o ? S + + + + +
Lamarck, 1822
Turritella declivis . .............. E +
Adams & Reeve, 1848
t Turris nigrovitta .... ....... . .... ? ? ? ? ?
Carrington & Kensley, 1969
(Koingnaas)
Vermetus sp. ?????? ?? 0 ?????? ... + + + + +
BIVALVIA
Arca avellana Lamarck, 1819 .... ?R E +
tArca halmyrus ............. ... ?R ? ? ? ? ?
Carrington & Kensley, 1969
(Strandfontein)
Arca noae Linnaeus, 1758 ...... ?R W +
Aulacomya ater (Molina, 1782) ... R + + +
Barnea truncata (Say, 1822) . . . ... R W + +
tCardita unica sp. novo . ... . ...... +
tCarditella calipsamma ... .... . + + + + + + +Carrington & Kensley, 1969
Cardium sp. ............ .. . . ... +
tChamelea krigei Haughton, 1926 S + + + +
Chlamys sp .... .......... ..... . +
Choromytilus meridionalis . ..? '0' R +
(Krauss, 1848)
tCorbula palaegialus .... . ....... ? ? ? ? ?
(Carrington & Kensley, 1969)
(Strandfontein)
tCuna aquaedulcensis .. .. - ...... +Kensley, 1977
t Donax haughtoni. ... . ... S + + + +
Carrington & Kensley, 1969
t Donax rogersi Haughton, 1926 ... S + +
tDosinia sicarisinus sp. novo .. .. S/M + + +
t Gastrana fibrosa .... . ..... . .... S +
Kilburn & Tankard, 1975
50 m Complex 30 m Complex
Habi- DLS LSHSpecies tat WfE TIL BBR USH LSH (KN/SL) USH LSH (KL)
BIVALVIA (contd)
tGastrana rostrata .... S
.......... . . +
Carrington & Kensley, 1969
Gastrana sp .................... S + +
tGlycymeris fulleri sp. nov ........ S +
?Hiatellasp ................ .... R + +
Hinnites sp. ................. . . +
tlsognomon gariesensis sp. novo ... + + +
Leporimetis hanleyi .......... ... S/M W + +
(Dunker, 1853)
Lutraria sp. . . . . . . . . . . . . . . . . . . . S/M + + + + + +
tMactrad. dernbergi ............ S/M +
Bbhm & Weisfermel, 1913
Melliteryx capensis .. .......... . RiS E +
(Sowerby, 1889)
tNotocallista schwarzi . . ??????? o ? S +
(Newton, 1913)
Nuculana bicuspidata ...... ..... S/M W + +
(Gould, 1845)
tOstrea d. subradiosa ........... R +
(Bbhm & Weisfermel, 1913
Perna perna (Linnaeus, 1758) .... R + + +
tPetricola prava ........ ..... ... . R + +
Kilburn & Tankard, 1975
Phaxas decipiens (Smith, 1904) ... S/M + + +
Scissodesma spengleri ........ ... S +
(Linnaeus, 1767)
tStandella namaquensis ....... ... S + + + +
Carrington & Kensley, 1969
Striostrea margaritacea ......... R E + + + + + + + +
(Lamarck, 1819)
Tellina ponsonbyi .......... .... S/M E + +
(Sowerby, 1889)
Tellina trilatera Gmelin, 1791 .... S + + + +
Theora sp. . . . . . . . . . . . . . . . . . . . . S +
Tivela cf. compressa ............ S + + +
(Sowerby, 1851)
Venus verrucosa Linnaeus, 1758 .. S W + + + +
SCAPHOPODA
Dentalium sp. ................. S/M +
POLYPLACOPHORA
Chaetopleura pertusa ........... R +
(Reeve, 1847)
d. Chiton sp .......... .???.. '0' R + + + + +
d. Dinoplax sp ................. R +
Table 1 indicates the preferred habitats of most species. In a number of cases
there is uncertainty about habitat, especially for extinct species. While most are
open-coast forms, there are calm-water components present (e.g. Bullia
laevissima, Phaxas deeipiens, Dosinia siearisinus, Lutraria sp., Leporimetis
hanleyi). Among the open-coast forms, rocky-shore species outnumber sand and
mud inhabitants almost two to one. Among the rocky-shore forms are a number
of grazers and algal inhabitants such as Haliotis, Crepidula, Oxystele, Turbo,
Calliostoma, and Patella. Predators such as Argobueeinum, Namamurex, Thais,
Oeenebra, Conus, and Terebra are also abundant. Filter feeders such as
Turritella, scavengers like Bullia and Nassarius, and parasitic forms such as
Epitonium and Melanella are also present, indicating a rich and diverse
environment with numerous habitats. Of the sand and mud dwellers there are
suspension and filter feeders (Donax, Dosinia, Seissodesma, Venus, Lutraria), as
well as deposit feeders (Tellina).
Of the species recorded by Carrington & Kensley (1969), only four have not
been found during the present study. These are Fusus faurei, Turris nigrovitta,
Corbula palaegialus, and Area halmyrus, all recorded from the 50 m Complex.
Nassarius litorafontis and Hespererato oppenheimeri, recorded from the 50 m
Complex by Carrington & Kensley (1969), have been found in the 30 m Complex
in this study.
ZOOGEOGRAPHIC AFFINITIES
There is a small west African-Mediterranean component of nine species in
the faunal list: Area noae, Barnea truneata, Cantharidus (Jujubinus) striatus,
Crepidula poreellana, Leporimetis hanleyi, Nueulana bieuspidata, Natiea cf.
adansoni, Sinum eoneavum, Venus verrueosa. Of these, Area noae, Cantharidus
(Jujubinus) striatus and Natiea cf. adansoni are strictly West Africa-Mediterra-
nean in distribution; the present fossil records are the most southerly for all three.
Nueulana bieuspidata occurs live in West Africa, and is known as a Pleistocene
fossil from Velddrif (Kruispad), Table Bay (Milnerton), and Port Elizabeth
(Redhouse) (See Kilburn & Tankard 1975: 206). Leporimetis hanleyi is known
live from Luanda, while an isolated population occurs at Sandvis, just south of
Walvis Bay. It is recorded as fossil in the Late Pleistocene in the Saldanha area,
and also at Redhouse, Knysna, Sedgefield, and Klein Brak River. Venus
verrueosa and Crepidula proeellana both occur from North Africa, around the
Cape, to Natal. Barnea truneata is known from the Late Pleistocene of Table Bay
(Kensley 1985a) and also from fresh dead shells washed ashore at the same
locality. Sinum eoneavum is an extant West African species, known from Senegal
to Angola (Nickles 1950). This so-called West African component thus represents
a mix of widespread temperature-tolerant species, as well as less temperature-
tolerant forms now restricted to more tropical areas.
The present-day distribution of living species represented in the faunal list
shows a majority of forms (27 species, 29 %) occurring on both east and west
coasts of southern Africa. Only 11 species are at present restricted to the east
coast: Calyptraea helieoidea, Melliteryx eapensis, Nassarius koehianus, Oxystele
sinensis, Ringieula turtoni, Area avellana, Striostrea margaritaeea, Tellina
ponsonbyi, Tugali barnardi, Turritella declivis, Cyliehna tubulosa. Of these, only
the Indo-Pacific Area avellana has a distribution extending north of southern
Mozambique.
The inferring of past sea-temperature regimes based on the known
temperature ranges of living species is a complex issue fraught with pitfalls. Some
guidelines do exist. Tankard (1975b: 33) presented temperature minima for a
range of Pleistocene molluscs of South Africa, and indicated a warm-water
affinity for a number of west-coast fossils. Several of these occur in the present
suite:
Temperature
minima
Nueulana bieuspidata 17 ?C
Seissodesma spengleri . . . . . . . . . . . . . . . . . . . . . 14?C
Tellina ponsonbyi. . . . . . . . . . . . . . . . . . . . . . . . . 14?C
Leporimetis hanleyi . . . . . . . . . . . . . . . . . . . . . . . 17?C
Kilburn & Rippey (1982) mention that Striostrea margaritaeea spawns when
monthly average temperatures exceed 20?C.
Examining the nine 'West African', and the 11 east-coast forms (with their
implied warmer-water requirements), it is seen that 7 species, including Striostrea
margaritaeea and Nueulana bieuspidata, occur in both the 50 m and 30 m
Complexes. Fourteen of the 20 species occur in the 50 m Complex, 12 in the 30 m
Complex. It is thus inadvisable to attribute a colder temperature regime to the
30 m Complex and a warmer regime to the 50 m Complex. Nevertheless, there is
undeniably a component showing a warm-water bias in the present suite of fossils,
deduced from known temperature ranges of living species. The temperature
requirements for the extinct forms, with the greater preponderance of species in
the 50 m Complex, can only be guessed at. The reasons for the extinction of these
forms are probably related in part to sea-temperature changes.
An occulinid coral, found encrusting rocks at Hondeklip in 50 m Complex
exposures, and tentatively identified as Sehizoeulina fissipara (Milne Edwards &
Haime), casts further light on the question of temperature regimes. According to
Laborel (1974), modern S. fissipara is adapted to low-salinity, warm, Guinean
waters, and has a ramose morphology. However, encrusting to subramose forms
(as in the present case) occur at the extremes of its range where periodic
upwelling takes place. This instance probably represents the most southerly
occurrence of S. fissipara in the fossil record and implies the extension of tropical
water southward to Hondeklip latitudes. Lithological evidence of upwelling off
Hondeklip during 50 m Complex times comes from phosphorite rinds inter-
bedded with regressive storm gravels deposited in the bedrock-defined embay-
ment during the earlier stages of bedrock emergence, prior to the establishment
of back-barrier conditions. The thin phosphorite rinds mark intervening
fairweather periods during the deposition of the storm gravel. Hondeklip was
probably situated adjacent to a marine regime characterized by the interaction of
upwelling and south-flowing tropical currents during the period of deposition of
the 50 m Complex.
The warm-water species common to both the 50 m and the 30 m Complexes,
especially the abundant Striostrea margaritacea, undoubtedly point to a sea-
temperature regime considerably higher than that prevailing on the west coast at
present. Over the period of deposition of both complexes, however, a cooling
trend must be inferred from the presence of numerous species living at present on
the west coast. The apparent reduction in diversity reflected by the 30 m Complex
may be indicative of cooling. Significantly, Choromytilus meridionalis evidently
first appears during 30 m Complex times (Early Pleistocene). The cooling trend is
also indicated by the extinction of Striostrea margaritacea on the west coast
subsequent to the Early Pleistocene.
The Hondeklip fossil fauna probably represents a mixture of forms at
different points in their history, reflecting both the influence of the cold Benguela
system and subtropical waters, fluctuating, glacially controlled sea-levels and
temperatures, along with varying temperature requirements. Thus, some
specimens may represent the last members of a population that was dying out
owing to decreasing temperatures, others a population adapting to fluctuating
conditions, others an 'experimental' pioneering stock having tenuous repro-
ductive success, and still other temperature-tolerant and actively reproducing
populations.
FAUNAL COMPARISONS
Evaluation of the fauna from Hondeklip and vicinity must necessarily take
the following into consideration.
The faunal list is undoubtedly incomplete. An unavoidable bias is, of course,
the very nature of the geological record. Only deposits accumulated during
regression from the sea-level maxima are available for examination; deposits
closer to the regressive maxima are not available, nor are deposits of the
transgressions. Thus the sedimentary packages available for sampling were
deposited during the earlier stages of recovery from major polar deglaciations. In
this sense successive complexes are somewhat directly comparable, but this may
also make for the reduction of faunal dissimilarity between complexes. Original
community patchiness, differential transport, preservational bias due to dissolu-
tion, and sampling bias (e.g. the tendency for small forms to be overlooked) all
contribute to an incomplete faunal list.
The latitudinal ranges of species in deposits equivalent to the 50 m and 30 m
Complexes are largely unknown or uncertain. Sampling was restricted to a few
localities in close proximity.
The temporal ranges of species are similarly unknown or uncertain. The Late
Tertiary and Middle Pleistocene are practically unsampled. The best comparative
faunal lists exist for the Late Pleistocene (e.g. Tankard 1975a; Kensley 1985a).
In spite of these limitations, comparisons with other west-coast assemblages
are instructive.
Bogenfels Tertiary Deposits
While the overall fauna of the Tertiary deposits of Bogenfels, South West
Africa-Namibia, is distinctive and very different from any present-day assem-
blage (B6hm & WeisfermeI1913; B6hm 1926) there are a few elements that hint
at a distant relationship with the Hondeklip suite.
The presence of a species of Pseudoliva (with one living species in West
Africa, and two extinct species from Bogenfels), mactrid hinges closely
resembling Mactra (Barymactra) dernburgi B6hm & Weisfermel from Bogenfels
(Fig. 42), and a single oyster valve almost identical with Ostrea subradiosa B6hm
(Fig. 35) suggest that there may well be Tertiary stragglers or their descendants in
the Hondeklip assemblage. This suggestion, however, must be treated as highly
speculative.
Marine Neogene of Ysterplaat
Tankard (1975a) provided a list of 13 molluscs recovered from the Miocene
of Ysterplaat, Cape. Olson (1985), on the basis of fossil penguin material from
the same locality, placed the assemblage in the Early Pliocene. The bivalves
Cardium edgari Newton, 1913, and Glycymeris borgesi (Cox, 1939) support a
Neogene age. Also taken from the Ysterplaat site were Donax serra Dillwyn,
1817, Scissodesma spengleri, Dosinia lupinus and a Pitar, since identified as
Notocallista schwarzi. Of these all but Donax serra and Dosinia lupinus occur at
Hondeklip. While most of the Ysterplaat Dosinia specimens are within the usual
size range of D. lupinus, a few reach the size of D. sicarisinus described above.
Unfortunately, the Ysterplaat material is all mouldic, and a clear impression of
hinge and mantle details is lacking, making a positive identification difficult.
Donax serra is discussed in the next section.
Early Pliocene molluscs from Langebaanweg, Cape
Kensley (1972, 1977) recorded a number of molluscs from Langebaanweg,
several of which also occur at Hondeklip. These are Cuna aquaedulcensis, Bullia
digitalis, B. laevissima, Tricolia capensis, Haliotis saldanhae, and Thais dubia.
Cuna aquaedulcensis is locally abundant in the 30 m Complex at Hondeklip.
Bullia digitalis is abundant in both the 30 m and 50 m Complexes; its living range
from South West Africa-Namibia to Transkei would indicate that this is a
temperature-tolerant species, and its presence in deposits from the Pliocene to
the Holocene is not surprising.
The fossil distribution of Donax serra, however, is difficult to explain. It has
been recorded from the Pliocene of Ysterplaat (Tankard 1975a), and Langebaan-
weg (Kensley 1972, 1977), and the Pleistocene of Liideritz, Orange River mouth,
Velddrif, Sedgefield (Barnard 1962), and Table Bay (Kensley 1985a). It has not
been seen in the Hondeklip deposits under discussion, where D. rogersi (30 m
Complex) and D. haughtoni (50 m Complex) are both abundant. Its present-day
distribution is from South West Africa-Namibia to Transkei, while De Villiers
(1975) reported the species to reproduce in a temperature range of 13-1rc.
Inappropriate temperature thus seems unlikely to be the reason for its absence
from the Hondeklip area. It is possible that local competition from the two extinct
species during the Early Pleistocene precluded the establishment of a population
of D. serra in the Hondeklip area.
Verlorevlei-Saldanha Pleistocene deposits
A number of species recorded from the Pleistocene of the south-western
Cape around Verlorevlei and Saldanha (Kilburn & Tankard 1975) are present in
the Hondeklip assemblage. The extinct species Petricola prava, Nucella prae-
cingulata, Triumphis dilemma and Fissurella robusta were obtained from deposits
at 10 masl at Saldanha Bay. The latter two species occur in the 50 m Complex and
the former two in both the 50 m and the 30 m Complexes at Hondeklip.
Cerithidea bifurcata Kilburn & Tankard, 1975, an extinct species also obtained
from the 10 masl deposit, has not yet been found in Namaqualand.
Tankard (1975c) correlated the Saldanha deposit at 10 masl with the 50 m
Complex in Namaqualand on the basis of the four species mentioned above. The
presence of Fissurella robusta and Triumphis dilemma suggests that this
correlation may be correct but it should be verified by additional criteria.
Accepting the equivalance of the deposits, comparison of the faunal lists shows an
additional 13 extant species in common with the 50 m Complex at Hondeklip, all
of which occur on both west and east coasts. Of the Saldanha fauna, only Patella
tabularis Krauss, 1848, P. concolor Krauss, 1848, and Peristernia nassatula
(Lamarck) are south- and east-coast species not present off Saldanha today.
Tankard (1975c) attributed the absence of Striostrea margaritacea and Donax
haughtoni from the Saldanha deposit to colder water conditions there during the
50 m Complex times.
Gastrana fibrosa, a probably extinct species recorded from the Late
Pleistocene in the south-western Cape, extends back into the Late Pliocene in
central Namaqualand, as shown by its occurrence in the 50 m Complex. The
extinct Late Pleistocene species Crepidula capensis praerugulosa Kilburn &
Tankard, 1975, is not present in the Hondeklip assemblage, though Crepidula
porcellana is.
Significantly, the warm-cold distinction between the 50 m and 30 m
Complexes respectively (or across the Plio-Pleistocene boundary), is not clear-
cut on the basis of existing data, due to the considerable faunal similarity between
the complexes. Notably, there are 34 species (11 extinct) common to both
complexes. Of the species unique to the 30 m Complex, only five are relatively
abundant, viz. Donax rogersi, Fissurella glarea, Choromytilus meridionalis, Cuna
aquaedulcensis, Ocenebra petrocyon. The reduction of diversity and the appear-
ance of Choromytilus meridionalis may suggest a cooler regime during 30 m
Complex times, relative to the 50 m Complex.
The value of Donax haughtoni and Donax rogersi as zone fossils for the 50 m
and 30 m Complexes, respectively, is verified in this study. Donax haughtoni is
present at least as far south as the Olifants River. Donax rogersi, the 30 m
Complex zone fossil, is present at least as far south as Doringbaai and has been
found as far north as Walvis Bay (pers. obs.). However, there are no faunal lists
from 30 m Complex correlates in the south-western Cape for comparison. It
appears that only subsequent to the Early Pleistocene did the Namaqualand
west-coast mollusc fauna more closely resemble the modern fauna. This probably
reflects the increasing dominance of the Benguela system and restriction of
warmer waters to the north, the cooler regime facilitating a species radiation
northwards from the south-western Cape. The ecological niche of Striostrea
margaritacea was usurped, to some extent, by the mytilids and patellids, the latter
to become an important food resource of early man on the west coast. The first
appearance of Patella compressa, the kelp limpet, already present by the Late
Pleistocene, is an important datum yet to be established.
Sedimentological aspects of the complexes suggest that, in the Hondeklip
area during 50 m Complex times, the coast was characterized by a sea-level
interaction with the bedrock topography that resulted in an embayed coast
concomitant with conditions for the development of extensive back-barrier
environments. In contrast, the 30 m Complex is characterized by relative
insulation from antecedent topographic effects, resulting in a more exposed
coastal regime. Superimposed upon these littoral influences are the effects of the
adjacent oceanographic temperature regime. Thus a secure database from which
to deduce Late Cenozoic palaeoclimatic influences will only exist once the
regressive packages are examined sedimentologically and faunistically sampled
on a regional basis in order to resolve the large-scale changes in the boundaries of
marine zoogeographic provinces in time, to recognize the environmental vectors
such as depositional palaeodepth and possibly thermally anomalous lagoons, and
allied to the latter, to evaluate the roles of antecedent topography and sediment
supply.
In conclusion, this study suggests that molluscs have potential for a
significant contribution in the unravelling of Late Cenozoic history. More extinct
molluscs were found than expected and the relatively high level of endemicity
holds promise that more zone fossils may emerge. The extant component of the
fauna permits some extrapolations of temperature tolerances and environmental
preferences. An overall cooling trend is reflected in the approach of successively
younger assemblages towards the modern west-coast faunal composition.
Comparison of Palaeogene, Neogene, Quaternary, and extant faunas
suggests that extinction, speciation, and migratory colonizations were staggered
as a reflection of differing temperature tolerances, habitat creation-destruction,
and competition. However, a regional mollusc-assemblage zonation scheme must
in time necessarily be supplemented by zonations of ostracode and benthic
foraminiferal assemblages. Planktonic Foraminifera are evidently very scarce in
these littoral deposits, while nannoplankton has not yet been collected, but
efforts to recover these microfossils may aid correlation with deep-sea data and
the global record. Resolution of the tectonic component of the coastal-plain
marine record awaits reliable regional correlations.
We thank Mr Clive Booth and Miss Elsabe Pretorius of the South African
Museum, and Mr Michael Carpenter of the Smithsonian Institution, for the
photographs used in this work.
We are grateful to Mr F. Hoffman of Transhex, for access to the mine
exposures at Hondeklip and Avontuur, and to Mr J. B. Hawthorne and Mr
R. Molyneaux of De Beers Consolidated Diamond Mines for arranging access to
the Koingnaas and Swartlintjies River exposures.
Dr George Steyskal of the Smithsonian Institution kindly checked the new
names and etymologies for correctness.
The paper benefited considerably from the comments and criticisms of Dr
R. N. Kilburn of the Natal Museum, and Dr J. Rogers of the University of Cape
Town.
We thank the Council for Scientific and Industrial Research for financial
assistance during the course of this work, and the Council of the South African
Museum for assistance and hospitality to the first author during several visits.
This work benefited immeasurably from the hospitality and co-operation of,
and discussions with, Dr Q. B. Hendey of the South African Museum, whose
encouragement to both authors is much appreciated.
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