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ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
VOLUME 69 BAND
MUS. COMP. ZOOL.
LIBRARY
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NEW GENERIC NAMES PROPOSED IN THIS VOLUME
Ianiroides Kensley, 1976
Ianisera Kensley, 1976
Inguza Simpson, 1975 ..
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297
71
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LIST OF CONTENTS
Page
BARGHUSEN, H. R.
Notes on the adductor jaw musculature of Venjukovia, a primitive anomodont
therapsid from the Permian of the U.S.S.R. (Published June 1976.) .. errs
BOUILLON, J. see MILLARD, N. A. H.
Ciuver, M. A. see GALTON, P. M.
GALTON, P. M. & CLUveER, M. A.
Anchisaurus capensis (Broom) and a revision of the Anchisauridae (Reptilia,
Saurischia). (Published April 1976.) ae kc = ie oo ed
Harris, J. M.
Pliocene Giraffoidea (Mammalia, Artiodactyla) from the Cape Province. (Published
September 1976.) .. ae a fe, Me ~ ah e yo) eae
HENDEY, Q. B.
The Pliocene fossil occurrences in ‘E’ Quarry, Langebaanweg, South Africa.
(Published April 1976.) _.. a 55 oe aa Ga ae ae AS
KENNEDY, W. J. see KLINGER, H. C.
KENSLEY, B.
Records of mud-prawns (genus Callianassa) from South Africa and Mauritius
(Crustacea, Decapoda, Thalassinidea). (Published December 1975.) .. on 47
KENSLEY, B.
Isopodan and tanaidacean Crustacea from the St Paul and Amsterdam Islands,
southern Indian Ocean. (Published June 1976.) .. ec AS 2: 2. 261
KLINGER, H. C., KENNEDY, W. J. & SIESSER, W. G.
Yabeiceras (Coniacian ammonite) from the Alphard Group off the southern Cape
coast. (Published March 1976.) .. ee a sve oe $e =. Gt
MILLARD, N. A. H. & BOUILLON, J.
Additional hydroids from the Seychelles. (Published December 1975.) .. = 1
Sesser, W. G. see KLINGER, H. C.
Simpson, G. G.
Notes on variation in penguins and on fossil penguins from the Pliocene of
Langebaanweg, Cape Province, South Africa. (Published December 1975.) .. 59
TANKARD, A. J.
Thermally anomalous Late Pleistocene molluscs from the south-western Cape
Province, South Africa. (Published August 1975.) a: ay se 2 17
TANKARD, A. J.
Pleistocene history and coastal morphology of the Ysterfontein—Elands Bay area,
Cape Province. (Published March 1976.).. xe sn ce Ss ro fis)
VAN DEN HEEVER, J. A.
The cranial and cervical muscles of the South African limbless lizard Typhlosdaurus
aurantiacus aurantiacus Peters (Reptilia, Sauria), (Published April 1976.) .. 169
VOLUME 69 PART 1 DECEMBER 1975 ISSN 0303-2515
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BuLLouGu, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FIsCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques.—J. Conch., Paris 88: 100-140.
FiscHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines.—Archs
Zool. exp. gén. 74: 627-634.
Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.—
Ann. Mag. nat. Hist. (13) 2: 309-320. it
Koun, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian
Ocean.— Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
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(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 69 Band
December 1975 Desember
Part 1 Deel
ADDITIONAL HYDROIDS FROM THE SEYCHELLES
by
N. A. H. MILLARD
&
J. BOUILLON
Cape Town Kaapstad
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ADDITIONAL HYDROIDS FROM THE SEYCHELLES
By
N. A. H. MILLARD
South African Museum, Cape Town
&
J. BOUILLON
Universite Libre de Bruxelles
(With 3 figures)
[MS. accepted 16 April 1975]
ABSTRACT
This paper supplements an earlier one on the hydroids of the Seychelles and records ten
more species. In addition three new species are described, namely Egmundella modesta,
Hebella muscensis and Scandia tubitheca. The last two have identical trophosomes but different
gonosomes, and arising from this it becomes necessary to consider Campanularia costata Bale,
1884 and Campanularia corrugata Thornely, 1904 as nomina oblita since these were both
described on sterile material.
CONTENTS
PAGE
Introduction Set ee ce eee 1
EISe Of Species’ i)" a2 We. we 2
Systematic section . . . .. . 3
Acknowledgements . : : ‘ . 14
References . ; : Sts ie s 14
INTRODUCTION
The hydroids described in this paper were collected by the secona author
during an expedition to the Seychelles Archipelago from June to September
1972. This expedition was financed by the Belgian ‘Fonds National de la
Recherche Scientifique’, the Belgian Ministry of Education and Culture, the
“Musée Royal de |’Afrique Centrale’ (Tervuren, Belgium) and the University of
Brussels. The collection is the property of the ‘Musée Royal de l’Afrique
Centrale’, where the types of new species have been deposited.
During a previous mission, in 1966, a large quantity of marine invertebrates
was collected and from the interest their study aroused we were led to consider
another expedition in order to complete our observations and extend them to
some islands of the archipelago never before prospected or scarcely so.
As for the hydroids, one paper has already been published by Millard &
Bouillon (1973) and another by Bouillon (1974). The present account expands
our knowledge of the hydroid fauna of this area.
]
Ann. S. Afr. Mus. 69 (1), 1975: 1-15, 3 figs.
2 ANNALS OF THE SOUTH AFRICAN MUSEUM
LIST OF SPECIES
Additional to those described by Millard & Bouillon (1973), together with
localities. Those species marked with an asterisk will be discussed further in the
pages which follow.
Family Asyncorynidae
Asyncoryne ryniensis Warren
A small fertile colony growing inside a dead Pecten shell, and bearing
several young medusae with cnidophores. Dredged between
‘Silhouette and Mahé.
Family Cladocorynidae
Cladocoryne floccosa Rotch
A large number (over 60) of infertile polyps from Praslin growing on
sponges.
Family Pandeidae
*Amphinema ?rugosum (Mayer)
A small fertile colony from Anse la Mouche.
Family Campanulinidae
*Egmundella modesta sp. nov.
Family Haleciidae
*Campalecium cirratum (Haeckel)
Three rich and fertile colonies from Anse la Mouche and Praslin.
Family Lafoeidae
* Hebella muscensis sp. nov.
* Scandia tubitheca sp. nov.
Family Syntheciidae
* Hincksella corrugata Millard
An infertile colony from Bird Island.
* Hincksella cylindrica pusilla Ritchie
Several stems reaching a maximum height of 8,7 mm from Bird Island.
Family Sertulariidae
* Dynamena obliqua Lamouroux
An infertile colony from Bird Island.
* Sertularella diaphana (Allman)
A fragment of an infertile stem 1,1 cm in length from Praslin.
Sertularia distans (Lamouroux)
An infertile colony from Bird Island.
ADDITIONAL HYDROIDS FROM THE SEYCHELLES 3
Family Plumulariidae
Pycnotheca mirabilis (Allman)
Two infertile stems from Bird Island.
In addition new information is provided for the following species:
Family Bougainvilliidae
* Silhouetta uvacarpa Millard & Bouillon
A fertile colony dredged between Silhouette Island and Beau Vallon
on Mahé Island, from which newly released medusae were obtained.
Family Campanulinidae
*Phialella quadrata (Forbes)
A fertile colony from Anse la Mouche growing on the stem of
Halocordyle disticha.
Family Syntheciidae
*Synthecium patulum (Busk)
Three fertile colonies, one dredged between Silhouette Island and
Beau Vallon on Mahé Island, and two from Anse la Mouche.
SYSTEMATIC SECTION
Family Bougainvilliidae
Silhouetta uvacarpa Millard & Bouillon, 1973
Silhouetta uvacarpa Millard & Bouillon, 1973: 25, fig. 3A—D, pls 2-3.
Description
Newly released medusae reaching a maximum size of approximately
0,9 (depth) x 1,0 mm (diameter) possess four oral tentacles with one dichotomy,
and four marginal tentacles. The branching oral tentacles confirm the inclusion
of this species in the Bougainvilliidae.
Family Pandeidae
Amphinema ?rugosum (Mayer, 1900)
Fig. 1A—-D
Amphinema rugosum: Rees & Russell, 1937: 67, figs 5—6. Russell, 1953: 183, fig. 90, pl. 10 (fig. 3),
pl. 11 (figs 2, 4). Kramp, 1965: 29.
Description
Solitary hydranths and medusa-buds arising separately from a creeping
hydrorhiza. Hydrorhiza with firm perisarc, about 0,03 mm in diameter.
Hydranths up to 3 mm in height, with one row of 10-12 filiform tentacles,
contained within a firm perisarcal tube for most of length. Perisarcal tube
slender and annulated at base, smooth for the rest and widening distally.
4 ANNALS OF THE SOUTH AFRICAN MUSEUM
aOR
F F-H
Fig. 1. Amphinema ?rugosum. A. Part of a colony sketched from a combination of several
damaged hydranths. B—D. Views of medusa-buds from whole mounts, right one viewed from
above, showing two tentacles, hypostome and radial canals.
Egmundella modesta sp. nov. E-H. Views of hydranthophores and nematothecae.
Scale in mm/10.
ner, meagan Fy ~~
ADDITIONAL HYDROIDS FROM THE SEYCHELLES a
Medusa-buds on short pedicels contained in wrinkled perisarc, oval, with
four radial canals and two opposite marginal bulbs bearing coiled tentacles.
No oral tentacles.
Measurements (mm)
Perisarcal tube, height Fy a i bi .. 1,41-2,53
diameter at base .. =< aed ae Ses .. 0,02-0,04
diameter at distal end oe We bs ail .. 0,09-0,13
Medusa-bud, height .. oF = = reaching 0,23
diameter... iy: - ae g reaching 0,20
Remarks
The medusa-buds, with their two stout marginal bulbs, resemble the genus
Amphinema, in which the hydranths of A. dinema and A. rugosum are known
(Rees & Russell 1937). Of the two this material is closer to A. rugosum, which
has stronger perisarc annulated at the base. Both species of medusa occur in the
plankton; A. dinema is rare and A. rugosum very abundant.
Family Campanulinidae
Egmundella modesta sp. nov.
Fig. 1E-H
Lovenella sp.: Millard & Bouillon, 1973: 42, fig. SE-F.
Holotype: an infertile stolonial colony from Anse la Mouche.
Description
Hydrothecae borne singly on the summit of pedicels of variable length.
Pedicel increasing in diameter from base to distal end; with thicker perisarc at
the base and thinner distally ; annulated irregularly, with 2—7 distinct annulations
at base and above this smooth or with faintly corrugated areas or with groups of
annulations, distal end always smooth.
Hydrotheca deep-campanulate, rounded at base, narrowed above this and
widening again at margin. A definite diaphragm present. Operculum of about
eight triangular segments clearly demarcated from thecal wall.
Nematothecae scattered on hydrorhiza, one-chambered, obovate to
globular, sessile, containing a cluster of large nematocysts.
Hydranth with 15-18 tentacles in the few extended individuals present, with
no intertentacular web.
Measurements (mm)
Pedicel, length 23 sf Sd = $3 .. 0,14-0,71
diameter at base .. 3 ds ae ae .. 0,03-0,05
Hydrotheca, depth .. oe xg 5g Y: .. 0,20-0,33
diameter at margin % J a a .. 0,08-0,11
Nematotheca, depth .. aye e 2 of .. 0,04-0,05
maximum diameter. . eS = os mi .. 0,02-0,04
6 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 2. Phialella quadrata. A. Gonotheca containing medusa-buds. B. Hydrotheca.
Campalecium cirratum. C. Hydranthophores with hydranths and gonotheca. D. Gonotheca
with medusa-bud ready to escape. E. Microbasic mastigophore, discharged and undischarged.
Scale: A—D in mm/10, E in mm/100.
ADDITIONAL HYDROIDS FROM THE SEYCHELLES q
Remarks
This species differs from E. amirantensis in the presence of a well-developed
and annulated thecal pedicel and in the sessile nematothecae.
The material is similar to that described by Millard & Bouillon (1973) as
?Lovenella sp. Re-examination of this earlier material has established the
presence of nematophores in the samples from Praslin although in the original
preparation it was not possible to relate them with certainty to the same
hydrorhiza.
The structure of the hydrotheca with its diaphragm and well-demarcated
opercular segments distinguishes E. modesta.from other species of Egmundella.
Indeed the hydrotheca is similar to that of Lovenella and the only character
definitely distinguishing the hydranth generations of these two genera is the
presence of nematothecae in the former.
Phialella quadrata (Forbes, 1848)
Fig. 2A-B
Hypsorophus quadratus: Huvé, 1952: 38, figs 3-7.
?Phialella quadrata: Millard & Bouillon, 1973: 43, fig. SG—J.
Description
Colony stolonial. Hydrothecae similar to those described from Mahé in
1973 and measurements within range.
Gonothecae present, elongated and truncated distally, reaching 0,57 mm
in length and 0,24 mm in maximum diameter, containing two medusa-buds,
one large and one small. Larger medusa-buds deep, with at least two marginal
bulbs and tentacles.
Remarks
The presence of gonothecae, which are exactly like those illustrated by
Huvé, supports the identification of this material.
Family Haleciidae
Campalecium cirratum (Haeckel, 1879)
Fig. 2C-E
Halecium simplex Pictet, 1893: 22, pl. 1 (figs 16-17). Ritchie, 1910: 807, pl. 77 (figs 10-11).
Campalecium medusiferum Torrey, 1902: 48, pl. 3 (figs 26-29). Huvé, 1954: 183, pls 7-9.
Eucheilota cirrata: Brinckmann, 1959: 82, figs 1-3.
Lovenella cirrata: Kramp, 1961: 177; 1968: 80, fig. 215.
Campalecium simplex: Rees & Thursfield, 1965: 112.
Description
Hydrorhiza reticular or forming long unbranched threads on the surface of
sponges and polyzoans, giving rise to hydranthophores at intervals.
8 ANNALS OF THE SOUTH AFRICAN MUSEUM
Hydranthophores usually solitary and with a single terminal hydranth,
occasionally with one or two sympodial branches; often regenerated. Pedicel
constricted at base, of very variable length. Hydrotheca with straight walls
widening to margin, with a circle of refringent nodules marking attachment of
hydranth. Hydranth very large, with 24~30 tentacles and an intertentacular web.
Gonotheca arising from side of pedicel below hydrotheca, curved, widening
to truncated distal end, containing three or four medusa-buds one above the
other.
Oldest medusa-bud with four perradial marginal bulbs of which two bear
tentacles and are slightly larger than the others, and eight adradial statocysts.
Although the margin is rather crumpled, in the largest medusa a marginal cirrus
is clearly visible next to three of the large marginal bulbs, and four interradial
cirri can be identified on rudimentary marginal bulbs.
Large nematocysts (microbasic mastigophores) present in intertentacular
web and in gonophores, 33,6 x 7,2 — 36,0 x 9,0 ». Capsule banana-shaped.
Butt with length approximately equal to that of capsule, bearing a raised spiral
ridge with small spines on the proximal region. Several other types of nematocyst
present in tentacles, but undischarged.
Measurements (mm)
Pedicel, length a “s - - in .. 0,07-0,99
diameter at base .. as a a ff .. 0,04-0,08
Hydrotheca, depth .. = a - Se a 0,02
diameter at margin mt Eas oe e .. 0,10-0,14
Gonotheca, depth eG aa ce a oy .. 0,41-0,77
maximum diameter. . Ss ie — os .. 0,23-0,38
Remarks
Brinckmann (1959) reared medusae of Lovenella cirrata (Haeckel, 1879)
from the polyp Haleciella microtheca Hadzi, 1914, which latter Huvé (1954)
included as a synonym for Campalecium medusiferum Torrey, 1902.
Rees & Thursfield (1965) synonymized Campalecium medusiferum with
Halecium simplex Pictet, 1893 after re-examination of Ritchie’s material (1910)
of the latter species from Mergui. They used the name Campalecium simplex.
However, they gave no description of the gonophores and did not comment on
Ritchie’s statement that the gonangium contained ‘ova to the number of about
six’. Recently the first author, by the courtesy of the Royal Scottish Museum,
Edinburgh, was able to re-examine Ritchie’s slides (nos. 1959. 33. 162-171).
Although it was not possible to decipher details of structure, the gonothecae
clearly contained medusa-buds and not eggs as described and illustrated by
Ritchie. We, therefore, confirm Rees & Thursfield’s synonymy, but point out
that of the available specific names cirratum Haeckel, 1879 antedates simplex
Pictet, 1893, medusiferum Torrey, 1902 and microtheca Hadzi, 1914.
Although the genus name Lovenella Hincks, 1868 has precedence over
ADDITIONAL HYDROIDS FROM THE SEYCHELLES 9
Campalecium Torrey, 1902, we do not feel that the former can be stretched to
contain a Haleciid polyp, and therefore retain the name Campalecium. We are
here faced with a species in which the polyp generation belongs to one family
(Haleciidae) and the medusa to another (Lovenellidae).
The presence of microbasic mastigophores in the species supports the idea
of an affinity between the Haleciidae and the Campanulinidae suggested earlier
by Millard (1975) and based on the presence of a Lovenellid medusa in Campale-
cium and certain resemblances of hydranth and hydrothecal structure in the two
families. These nematocysts, which are by no means common in the Athecata,
occur also in Hydrodendron caciniformis (personal observation of first author)
and Halecium halecinum (Weill 1934) among the Haleciidae, and in Eucheilota
maculata and Eutonina indicans among the Campanulinidae (Werner 1968a,
19685). Werner suggests that their evolutionary forerunners are basitrichous
isorhizas, which are often difficult to distinguish from them and which occur
commonly in the Campanulinidae.
The polyp generation of C. cirratum is known from the Mediterranean, the
Indo-West Pacific (Moluccas, Mergui) and the eastern Pacific (California). This
is the first record from the western Indian Ocean. These records, together with
those of the medusa from the Mediterranean, various parts of the tropical
Atlantic and Malaya (Kramp 1961, 1968), show the species to be circumtropical
in distribution.
Family Lafoeidae
Remarks on Hebella corrugata and H. costata
Campanularia corrugata Thornely, 1904 was described from a sterile
colony from Ceylon. The type material of Campanularia costata Bale, 1884 from
Port Darwin, with which species Billard (1941) synonymizes Hebella corrugata,
was also sterile. Since the original description of these two species all material
delegated to one or the other has been sterile, except for some empty and
probably immature gonothecae described by Billard (1941). In 1973, however,
Millard & Bouillon described gonophores in material from the Seychelles,
which was attributed to Thornely’s species. Since the gonophores were fixed
sporosacs and showed no medusoid characters, the species was transferred from
Hebella to Scandia, thus: Scandia corrugata.
In this second collection from the Seychelles, material is present with an
identical trophosome—it cannot be distinguished from that described in 1973
on measurements or any other character—but with gonothecae containing well-
developed medusa-buds. It is apparent, therefore, that there are two species with
identical trophosomes and different gonosomes, one a Hebella and the other a
Scandia. Since there is no means of telling to which of these Thornely’s and
Bale’s species belong, the only immediate solution is to create two new species,
and to regard Hebella corrugatum (Thornely 1904) and Hebella costata (Bale
1884) as nomina oblita. Sterile material cannot be identified, and all sterile
records attributed to either of the two species should be disregarded.
10 ANNALS OF THE SOUTH AFRICAN MUSEUM
Hebella muscensis* sp. nov.
Fig. 3A-B
Holotype: a fertile colony epizootic on Synthecium sp. from Anse la Mouche.
Description
Hydrotheca tubular, curved to one side, with 5-9 transverse annulations,
with everted margin usually oblique to axis and lower on the shorter, more
concave side. An annular thickening present round base, asymmetrically
developed, pronounced on the shorter, more concave side and often not notice-
able on the other. Pedicel short, not annulated, but occasionally with an indistinct
node separating it from the hydrorhiza.
Gonotheca borne on hydrorhiza on short pedicel, elongated, widening
distally, often slightly curved, usually longer than hydrotheca, with transverse
annulations which may be somewhat irregular, with an operculum of four
segments, containing up to four medusa-buds one above the other. Medusa-bud
with rounded hypostome, at least four marginal tentacles and a varying number
of ocelli (usually eight).
Measurements (mm)
Pedicel length .. a ae el se te .. 0,08-0,17
Hydrotheca, depth, convex side “ a - .. 0,82-1,20
diameter at mouth .. oe a a ve .. 0,35-0,53
diameter/depth ‘ss 3 cia a: ae .. 0,37-0,50
Gonotheca, length .. aS ea ws % .. 1,04-1,42
maximum diameter. . wm rs - me .. 0,38-0,53
Scandia tubitheca sp. nov.
Scandia corrugata: Millard & Bouillon, 1973: 60, fig. 8D—F (fertile colony only).
Holotype: fertile colony from Amirante, Seychelles, epizootic on Synthecium
dentigerum.
Diagnosis
Hydrotheca similar to that of Hebella muscensis. Gonotheca (only male
known) also very similar, but wider distally and not curved, containing a single
gonophore in the form of a fixed sporosac.
Family Syntheciidae
Hincksella corrugata Millard, 1958
Hincksella corrugata Millard, 1958: 181, fig. 5. Gravier, 1970: 116.
Description
Unbranched stems reaching 8,0 mm and with up to 10 hydrothecae.
Structure similar to holotype, but dimensions all slightly less.
* From Anse la Mouche: Bay of flies.
ADDITIONAL HYDROIDS FROM THE SEYCHELLES 11
E
Fig. 3. Hebella muscensis sp. nov. A. Hydrothecae. B. Gonothecae containing medusa-buds.
Synthecium patulum. C. Part of stem with a pair of gonothecae in narrow view. D. Part of
hydrocladium. E. Gonothecae: a smooth one in narrow view, an annulated one in narrow view
and a smooth one in broad view.
Scale in mm/10.
12 ANNALS OF THE SOUTH AFRICAN MUSEUM
Measurements (mm)
Stem, internode length =. . “a a .. 0,41-0,74
Hydrotheca, length abcauline a ss $e .. 0,43-0,53
length adcauline, adnate part ee ae a? .. 0,30-0,33
length adcauline, free part. . .; ea ne .. 0,30-0,38
diameter at margin a Hy sé BY .. 0,3-0,41
Remarks
H. corrugata has been reported from the western Indian Ocean from Natal
to S.E. Madagascar (Gravier 1970). This record extends the range further north.
The gonophores still await discovery.
Hincksella cylindrica pusilla Ritchie, 1910
Sertularella cylindrica var. pusilla Ritchie, 1910: 817, pl. 77 (fig. 9).
Hincksella cylindrica pusilla: Millard, 1964: 22, fig. 6A—D.
Cyclonia pusilla: Hirohito, 1969: 16, fig. 12.
Description
Stems normally unbranched, but one with a single lateral branch. Hydro-
theca with a larger proportion adnate than the South African material described
by Millard (1964), but otherwise very similar. Empty gonothecae present,
arising from within hydrothecae, elongated and tapering distally.
Measurements (mm)
Stem, internode length << i ey 7 .. 0,26-0,59
Hydrotheca, length abcauline sii a ms .. 0,35-0,54
length adcauline, adnate part ae ah Se .. 0,15—0,34
length adcauline, free part. . = Js 2 .. 0,28-0,41
diameter at margin ae a .. 0,12-0,27
Gonotheca, length from fpiecticeal margin ‘oe .. 0,57-0,77
maximum diameter. . ae 7 fe a 2 Q22-835
Remarks
This is the first discovery of gonophores in African material of the sub-
species, and the shape is similar to that of the male gonophores described by
Hirohito from Japan.
Synthecium patulum (Busk, 1852)
Fig. 3C-E
Synthecium orthogonia: Bale, 1888: 767, pl. 17 (figs 1—5).
Synthecium campylocarpum Allman, 1888: 78, pl. 37 (fig. 1).
Synthecium patulum: Billard, 1925: 125, figs 2-3. Millard & Bouillon, 1973: 64, fig. 8J.
Description
Pinnate stems reaching 2,0 cm, with 1-3 pairs of hydrothecae between
successive pairs of hydrocladia.
ADDITIONAL HYDROIDS FROM THE SEYCHELLES 13
Gonothecae arising from within hydrothecae on stem or hydrocladia and
rather different in appearance from those described in 1973, being longer and
more compressed. The transverse annulations, which number up to 11, are
restricted to the centre region and are visible only in side view unless the
gonotheca is empty, when the appearance is similar to Bale’s diagram (1888:
pl. 17 (fig. 5)). Within the same colony are gonothecae which have no annula-
tions at all and are completely smooth. All gonothecae are male.
Measurements (mm)
Hydrotheca, length abcauline os te as .. 0,36—0,50
length adcauline, adnate part i x — .. 0,48-0,63
length adcauline, free part. . f rt Ay .. 0,08—-0,20
diameter at margin .. ie oh - - OP RS ee:
Gonotheca, length .. ae ne a & .. 1,38-1,84
breadth ue ae ae K e ig .. 0,56—0,74
thickness... be rs as - a .. 0,20-0,41
Remarks
Although the gonothecae are more compressed than any previously
illustrated for S. patulum or its synonyms, we do not feel justified in creating a
new species for what is probably a variable feature. It is possible that the shorter
and fatter gonothecae illustrated in 1973 were female.
Family Sertulariidae
Dynamena obliqua Lamouroux, 1816
Pasythea quadridentata var. balei Billard, 1907: 355, fig. 6.
Dynamena obliqua: Millard, 1958: 184, fig. 6A.
Description
Stems reaching 6,6 mm, most of them with hydrothecal pairs ungrouped,
some of them with one or two groups of two pairs. Hydrothecae smaller than
those reported from Mogambique (Billard 1907) and from South Africa
(Millard 1958), but shape very similar. Internal teeth present in most hydro-
thecae, one adcauline and two latero-abcauline.
Measurements (mm)
Hydrotheca, length abcauline Ses _ ii .. 0,20-0,24
length adcauline, adnate part = ay Js .. 0,20-0,24
length adcauline, free part. . ge rif < .. 0,12-0,14
diameter at mouth .. art he ne Si .. 0,07-0,08
14 ANNALS OF THE SOUTH AFRICAN MUSEUM
Sertularella diaphana (Allman, 1886)
Thuiaria diaphana Allman, 1886: 145, pl. 18 (figs 1-3).
Sertularella diaphana: Billard, 1925: 157, figs 22-24, pl. 7 (figs 12-14). Millard, 1958: 188,
fig. 7C—D.
Description
This stem is unusual in that it rotates through 90° half-way up. Both stem
and hydrocladia are more slender than the South African material (Millard 1958)
and are close to Billard’s var. delicata (1925). The hydrocladia have only two or
three hydrothecae to an internode.
Remarks
This species is known from the east coast of Africa, Mauritius and
Madagascar, so its presence in the Seychelles is not unexpected.
ACKNOWLEDGEMENTS
The authors acknowledge with gratitude their indebtedness to Mr Kandi
Jivan Shah and Mr S. Savy, Director of the Department of Agriculture, for their
kindness, co-operation and constant help during the stay of the mission in the
Seychelles Islands.
REFERENCES
ALLMAN, G. J. 1886. Description of Australian, Cape and other Hydroida, mostly new, from
the collection of Miss H. Gatty.—J. Linn. Soc. (Zool.) 19: 132-161.
ALLMAN, G. J. 1888. Report on the Hydroida dredged by H.M.S. Challenger during the years
1873-76. Part IIl.—The Tubularinae, Corymorphinae, Campanularinae, Sertularinae and
Thalamophora.— Rep. Voy. Challenger 1873-76 23(70): 1-90.
BALE, W. M. 1884. Catalogue of the Australian hydroid zoophytes. Sydney: Australian Museum.
Bae, W. M. 1888. On some new and rare Hydroida in the Australian Museum collection. —
Proc. Linn. Soc. N.S.W. (2) 3: 745-799.
BILLARD, A. 1907. Hydroides de Madagascar et du sud-est de l’Afrique. — Archs Zool. exp. gén.
(4) 7: 335-396.
BILLARD, A. 1925. Les Hydroides de l’expédition du Siboga. II. Synthecidae et Sertularidae. —
Siboga Exped. 7b: 117-232.
BILLARD, A. 1941. Note sur les Hydroides: Hebella costata (Bale) et H. corrugata (Thornely). —
Bull. Soc. zool. Fr. 66: 13-15.
BOUILLON, J. 1974. Description de Teissiera milleporoides, nouveau genre et nouvelle espéce
de Zancleidae des Seychelles (Hydrozoaires; Athécates-Anthoméduses), avec une révision
des Hydroides “‘Pteronematoidea’’.— Cah. Biol. mar. 15: 113-154.
BRINCKMANN, A. 1959. Uber den Generationswechsel von Eucheilota cirrata (Haeckel 1879).—
Pubbl. Staz. zool. Napoli 31: 82-89.
GraAvieR, N. 1970. Etude des Hydraires epiphytes des Phanérogames marines de la région de
Tulear (sud-oest de Madagascar).— Rec! Trav. Stn mar. Endoume-Marseille 10: 111-161.
HirouiTo, Emperor of Japan. 1969. Some hydroids of the Amakusa Islands. Tokyo: Imperial
Household.
Huve, P. 1952. Revision des polypes Campanulinides mediterranéens.— Recl Trav. Stn mar.
Endoume 4: 34-47.
Huve, P. 1954. Hydranthea et Campalecium. Genres Mediterraneens aberrants d’Hydroides
de la famille des Haleciides.— Rec! Trav. Stn mar. Endoume 13: 173-192.
KRamp, P. L. 1961. Synopsis of the medusae of the world.—J. mar. biol. Ass. U.K. 40: 7-469.
ADDITIONAL HYDROIDS FROM THE SEYCHELLES 13
KRaAmpP, P. L. 1965. The Hydromedusae of the Pacific and Indian Oceans.— Dana Rep. 63:
1-161.
Kranmp, P. L. 1968. The Hydromedusae of the Pacific and Indian Oceans. Sections II and III. —
Dana Rep. 72: 1-200.
Miciarp, N. A. H. 1958. Hydrozoa from the coasts of Natal and Portuguese East Africa.
Part I. Calyptoblastea.— Ann. S. Afr. Mus. 44: 165-226.
MILLARD, N. A. H. 1964. The Hydrozoa of the south and west coasts of South Africa. Part II.
The Lafoeidae, Syntheciidae and Sertulariidae.— Ann. S. Afr. Mus. 48: 1-56.
MILLARD, N. A. H. 1975. Monograph on the Hydroida of southern Africa.— Ann. S. Afr. Mus.
68: 1-513.
MILLARD, N. A. H. & BourILLon, J. 1973. Hydroids from the Seychelles (Coelenterata). —
Annls Mus. r. Afr. cent. Sér 8vo (Sci. zool.) 206: 1-106.
Pictet, C. 1893. Etude sur les Hydraires de la Baie d’Ambouine.— Revue suisse Zool. 1: 1-64.
Rees, W. J. & RussELL, F. S. 1937. On rearing the hydroids of certain medusae, with an
account of the methods used.—J. mar. biol. Ass. U.K. 22: 61-82.
Rees, W. J. & THURSFIELD, S. 1965. The hydroid collections of James Ritchie.—Proc. R. Soc.
Edinb. (B) 69: 34-220.
Ritcuie, J. 1910. The marine fauna of the Mergui Archipelago, Lower Burma, collected by
Jas. J. Simpson, M.A., B.Sc., and R. N. Rudmose-Brown, D. Sc., University of Aberdeen,
February to May 1907.—The hydroids.—Proc. zool. Soc. Lond. 1910: 799-825.
RUussELL, F. S. 1953. The medusae of the British Isles. Cambridge: University Press.
THORNELY, L. R. 1904. Report on the Hydroida collected by Professor Herdman, at Ceylon,
in 1902.— Rep. Govt Ceylon Pearl Oyster Fish. Gulf Manaar suppl. Rep. 8: 107-126.
Torrey, H. B. 1902. The Hydroida of the Pacific coast of North America, with especial
reference to the species in the collection of the University of California.— Univ. Calif.
Publs Zool. 1: 1-104.
WEILL, R. 1934. Contribution a l’étude des Cnidaires et de leurs nématocystes. I. Recherches
sur les nématocystes. II. Valeur taxonomique du cnidome.—Trayv. Stn zool. Wimereux
10: 1-347; 11: 349-701.
WERNER, B. 1968a. Polypengeneration und Entwicklungsgeschichte von Eucheilota maculata
(Thecata— Leptomedusae). — Helgoldnder wiss. Meeresunters. 18: 136-168.
WERNER, B. 19685. Polypgeneration und Entwicklung von Eutonina indicans (Thecata—
Leptomedusae). — Helgoldnder wiss. Meeresunters. 18: 384-403.
6. SYSTEMATIC papers must conform with the International code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. n., sp. n., comb. n.,
syn. n., etc.
a An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.
Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (figs 8a—b).
Nucula largillierti Philippi, 1861: 87
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers
Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.
In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:
Holotype
SAM-A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach,
Port Elizabeth (33.51S, 25.39E), collected by AS Smith, 15 January 1973.
Note standard form of writing South African Museum registration numbers and of date.
7. SPECIAL HOUSE RULES
Capital initial letters
(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. ‘... the Figure depicting C. namacolus .
*,..in C. namacolus (Fig. 10)...’
(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A. L. du Toit
Von Huene but F. von Huene
(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian
Punctuation should be loose, omitting all not strictly necessary
Reference to the author should be expressed in the third person
Roman numerals should be converted to arabic, except when forming part of the title of a
book or article, such as
‘Revision of the Crustacea. Part VIII. The Amphipoda.’
Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.
N. A. H. MILLARD
. &
J. BOUILLON
ADDITIONAL HYDROIDS FROM THE SEYCHELLES
(OLUME 69 PART 2 AUGUST 1975 ISSN 0303-2515
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(a) Author’s name and = of publication given in text, e.g.:
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Note: no comma separating name eae year
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within each name, with suffixes a, b, etc. to the year for more than one paper by the same
author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a).
For books give title in italics, edition, volume number, place of publication, publisher.
For journal article give title of article, title of journal in italics (abbreviated according to the World list of
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number (only if independently paged) in parentheses, pagination (first and last pages of article).
Examples (note capitalization and punctuation)
BuLLOuGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques.—J. Conch., Paris 88: 100-140.
FiscHer, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines.—Archs
Zool. exp. gén. 74: 627-634.
Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.—
Ann. Mag. nat. Hist. (13) 2: 309-320. :
Konn, A. J. 19606. Spawning behaviour, egg — and larval development in Conus from the Indian
Ocean. — Bull. Bingham oceanogr. Coll. 17 (4): 1-51
THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: ScHULTzE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer.— Denkschr. med.-naturw. Ges. Jena 16: 269-270. é d
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 69 Band
August 1975 Augustus
Part 2 Deel
THERMALLY ANOMALOUS
LATE PLEISTOCENE MOLLUSCS
FROM THE SOUTH-WESTERN CAPE PROVINCE,
SOUTH AFRICA
By
ANTHONY J. TANKARD
Cape Town Kaapstad
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THERMALLY ANOMALOUS LATE PLEISTOCENE MOLLUSCS FROM
THE SOUTH-WESTERN CAPE PROVINCE, SOUTH AFRICA
By
ANTHONY J. TANKARD
South African Museum, Cape Town
(With 3 figures and 2 tables)
[MS. accepted 29 May 1975]
ABSTRACT
Numerous exposures of Late Pleistocene marine sediments occur on the west and south
coasts of the Cape Province. The mollusc fauna, which includes some 124 species, only 3 of
which are extinct, lived in sediments deposited during the last interglacial. Two broad and
contemporaneous facies are recognized: a cool-water open-coast facies, and a warm-water
estuarine-lagoonal facies. Whereas the open-coast facies is characterized by molluscs which
presently live on the adjacent coast, the estuarine-lagoonal facies regularly contains tropical
species which lived far south of their known present-day geographic ranges. Differences in
distribution between the fossil warm-water molluscs and their present-day temperature-
sensitive counterparts have been used to interpret Late Pleistocene climatic change in the
south-western Cape. These deposits are correlated with the Mediterranean Eutyrrhenian.
CONTENTS
PAGE
Introduction é , ‘ é P ; ‘ ’ ily
Geologic Setting . : ; : : : : . 18
Methods . ; ‘ : - ; ‘ : : 20
Palaeoecology . ‘ ‘ : ‘ ; i ; 21
Population Dynamics : é ‘ : : : 25
Predation Z P : ‘ ‘ : ; 29
Teratological Specimens : ; P : , ‘ 31
Depth |: ; , , ‘ ‘ : , ‘ 31
Temperature . : ; i ‘ ; ‘ : 32
Summary ; : ; ; F . 37
Palaeoclimatic Interpretation ‘ ; : 38
Review of Present Climate and Hydrology 3 P ‘ 38
Late Pleistocene Climate . , F : ‘ 40
Discussion i ; F ‘ F ‘ : : 41
Acknowledgements : , ‘ : ¥ ; F 43
References . ‘ : ; ‘ . : . : 43
INTRODUCTION
Isolated fossiliferous, marine sediments occur on the wave-cut platform
and in the sheltered Late Pleistocene embayments from Elands Bay on the west
coast to Knysna on the south coast. The fossils occur in unconsolidated
quartzose and shelly quartzose sands. The invertebrate fauna is essentially
modern in composition and comprises some 150 species which today live in
shallow-water environments. Only three of the mollusc species from the west
17
Ann. S. Afr. Mus. 69 (2), 1975: 17-45, 3 figs., 2 tables
18 ANNALS OF THE SOUTH AFRICAN MUSEUM
coast deposits are not known to be living today. In so far as most of these taxa
are still living, and are inhabitants of shallow water, they provide ideal material
for a palaeoecological study.
The Late Pleistocene fauna of the west coast is broadly divisible into two
distinct, but contemporaneous, ecologic zones. These are, firstly, a cool-water,
open-coast facies characterized by rocky shore and sandy beach assemblages,
and secondly, a warm-water, sheltered embayment facies (estuaries and lagoons).
Whereas the open-coast facies is laterally continuous, the sheltered embayment
facies is restricted in distribution. The open-coast facies is characterized by
molluscs which commonly inhabit the present coast. A striking feature of the
estuarine-lagoonal facies is the association of extant and extralimital thermo-
philic species. (“Thermophilic’ and ‘extralimital’ imply species which occur
outside their normal spawning range.) Examination of their present latitudinal
ranges indicates that a significantly warmer hydroclimate prevailed when those
species were common along the south-western Cape coast. In attempting to
reconstruct the palaeoenvironment, a detailed examination of a south coast
assemblage is necessary. The fossiliferous deposits at Knysna are therefore
included in the study.
Late Pleistocene faunas of open-coast and sheltered-embayment aspects
from southern California are similarly distinctive. Nearly all the dominant
species also inhabit the adjacent coast. But the sheltered embayments also
contain a high proportion of thermophilic molluscs which are found far north
of their present-day geographic range end-points. Several authors have used
these anomalies as a key to Late Pleistocene climatic interpretation (Valentine
1955, 1957, 1961; Valentine & Meade 1961; Addicott & Emerson 1959; Emerson
& Chase 1959; Kern 1971). Valentine (1955) explained the diverse nature of
the fauna by changes in intensity of the oceanic circulation and upwelling,
while the water of the embayments was heated by increased solar radiation. A
recent alternative explanation suggests that the larvae of tropical molluscs were
transported into the cooler areas by periodic local and temporary current
changes, and that they became only temporary, non-breeding, members of the
community (Zinsmeister 1974).
The south-western Cape Province, with its extensive Late Pleistocene
fossiliferous deposits, and its complex present-day ocean current systems, is
well placed to make a contribution to the knowledge of Late Pleistocene
hydroclimates. The purpose of this paper is to describe the mollusc fauna,
particularly between Ysterfontein and Elands Bay, and to examine their
palaeoenvironmental significance. The taxonomy of some of these molluscs
appears elsewhere (Kilburn & Tankard, in press).
GEOLOGIC SETTING
The Pleistocene epoch was characterized by the waxing and waning of
continental ice sheets with sea level oscillating in sympathy. Relative movement
of sea level is shown by the emerged wave-cut platforms, stranded beaches,
i etl
a ee
THERMALLY ANOMALOUS LATE PLEISTOCENE MOLLUSCS
Dar-es- Salaam
\ TANZANIA
St Helena Bay
32°45S |
Paternoster
15 4
Kruispad
wz Vredenburg
2
ny
YASaldanha
aldanha
B
lopefield
Fig. 1. Locality map. The insert (partly after Boss 1969) shows the
distribution of present-day coastal forms: black = _ tropical;
stippled = warm-temperate; area bounded by broken line on west
coast = cold water fauna.
19
20 ANNALS OF THE SOUTH AFRICAN MUSEUM
and marine terraces. Research on Barbados has suggested the existence of three
transgressions in the last interglacial (Eem): +6 m (BI) at 122 ka (i.e. 122 000
years ago); —13 m (BIJ) at 103 ka; —13 m (BIII) at 82 ka (Broecker et al. 1968).
These three peaks are confirmed by oxygen isotope studies of deep-sea cores
(Shackleton & Opdyke 1973).
Along the west coast of South Africa a thin mantle of Late Pleistocene
marine sediments overlies earlier Pleistocene, Tertiary, and pre-Tertiary rocks.
On the granitic headlands between St. Helena Bay and Ysterfontein a narrow
seaward-sloping wave-eroded platform is encountered. St. Helena Bay has
formed on Malmesbury rock which weathers more rapidly than the granite
and gives a negative relief. The open-coast fossil assemblages were collected
from marine deposits overlying the granite platforms, and from exposures in
the coastal flats bordering St. Helena Bay.
The last interglacial shorelines parallel the present coast very closely,
with two exceptions. Whereas the present coastline is interrupted only by
Saldanha Bay and its southerly offshoot, Langebaan Lagoon, the Late
Pleistocene sea extended up the Berg River valley and along Verlorevlei to
form two prominent estuaries. Verlorevlei is now separated from the sea by a
bar of Palaeozoic sandstone at | m a.s.]. (above mean sea level). These sheltered
Late Pleistocene estuaries and lagoons were of considerable palaeogeographic
importance since all fossil sites within them contain a significant proportion
of thermophilic mollusc species.
The greatest observed thickness of the open-coast facies exceeds 5m at
Velddrif; the greatest observed thickness of the estuarine facies is 2,2 m at
Verlorevlei. The open-coast facies consists of medium-grained quartzose sand
and comminuted shell. Frequently there are shell beds and banks with little
detrital quartz, indicating a slow supply of detrital sediment. A series of
emerged breaker-bars between Velddrif and Laaiplek parallel the modern
coast. These are composed largely of shell material.
In all the sheltered embayments the fossils are found in fine sands. At
Verlorevlei the marine horizon is overlain by coarse, poorly-sorted colluvium.
At Churchhaven the marine sediments are carbonate cemented and form an
erosional bench just above high-water level. Most of the sites examined, both
open-coast and sheltered embayment, are covered with a veneer of wind-blown
sand.
METHODS
The absolute density of fossil specimens of each molluscan species at any
particular site is difficult to determine because of the sampling problems
inherent on the size of the shell. For instance, ‘Rissoa’ capensis is a small
gastropod (usually less than 3mm) and would frequently number in the
hundreds from just 100 g of sediment, while the large (approximately 200 mm)
Panopea glycymeris would occur at approximately 1 m intervals. Bearing in
mind that the present study is a palaeoecological one, and that the larger
THERMALLY ANOMALOUS LATE PLEISTOCENE MOLLUSCS 21
molluscs have been better studied with respect to taxonomy and ecology,
emphasis has in all cases been placed on the macro-molluscs. A further
source of error arises from the differential fragmentation of shells. Bivalve
shells are more easily broken than gastropod shells, and in the living assem-
blage they may have been far more abundant than the faunal list suggests.
Table 1 should be taken only as an approximation to the original community
structure.
The procedures adopted in drafting Table 1 are as follows:
1. It was desirable to sample as small an area as possible to obtain not
only the absolute density of each species, but also to obtain restricted samples
for size-frequency analyses. In most cases a quadrat size of | square metre
has been found adequate, although quadrat size may have to be adjusted up or
down depending upon the relative abundance of specimens.
2. For a shell to be counted as an individual it must be nearly complete,
or so nearly so that the remainder could not be identified and counted separately.
Left and right valves of each bivalve species were counted separately, and the
highest count taken as the total number of individuals of that species in that
quadrat.
3. Each species has been recorded in Table 1 as percentage frequencies
in the sample where more than 50 individuals were counted. Where a total of
less than 50 individuals were counted they are recorded in Table 1 as ‘x’.
No attempt has been made to relate species to sediment texture since every
mollusc at some time must have been living among already dead and fragmented
shells. The substrate would thus consist of quartzose sand and bioclastic
material ranging in size from complete shells to finely comminuted fragments.
PALAEOECOLOGY
In this paper past extensions of tropical and subtropical mollusc geographic
ranges are used as a basis fcr interpreting Late Pleistocene palaeotemperature
changes. These inferences are based only on fossils of still extant species, and
the assumption (Durham 1950) that stenothermal organisms are in general
more critically limited by minimum temperatures than by maximum tempera-
tures. The validity of such palaeotemperature inferences depends on the fossils
being preserved in the sediments in which they once lived.
In describing the relationship between the fossils, after death, and the
sedimentary environment, we define the following types of fossil assemblages:
1. Life assemblage: disturbance after death negligible (Hallam 1960).
2. Death assemblage:
(i) Indigenous: organic remains disturbed after death but not trans-
ported very far (Hallam 1960).
(ii) Transported: organic remains introduced from a neighbouring
contemporaneous or older environment.
3. Mixed assemblage: this comprises any combinations of the above
ANNALS OF THE SOUTH AFRICAN MUSEUM
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THERMALLY ANOMALOUS LATE PLEISTOCENE MOLLUSCS 25
possibilities, and is the general case (Hallam 1960). The status of this assemblage
is clarified by describing it, for instance, as a mixed life and indigenous death
assemblage. In the present study this is the commonest case.
POPULATION DYNAMICS
Boucot (1953), Olson (1957), Craig & Hallam (1963), and Craig & Oertel
(1966) have attempted to discriminate between life and indigenous death
assemblages on the one hand and transported death assemblages on the other,
by using size-frequency distributions. According to Boucot an indigenous death
assemblage is characterized by a positively skewed distribution (large number
of small forms), while negative skewness or a normal distribution characterizes
the transported death assemblage. He suggested that negative skewness resulted
from winnowing of the smaller shells. Craig & Oertel (1966) question this and
maintain that the shape of the size-frequency distribution in a fossil population
depends principally upon the growth-rate and mortality-rate of the relevant
species. In this way negative skewness could arise from a decreasing growth-rate
with constant mortality-rate which would concentrate the older age-classes
in a few size-classes. Craig and Oertel suggest the following options:
Growth-rate Mortality-rate Size-frequency distribution
decreasing constant negative skewness
constant decreasing positive skewness
constant increasing flattening of curve, possibly
negative skewness
constant constant mirror image of living popul-
ation by dead population
Growth-rate and mortality-rate complement each other when one decreases and
the other increases, but cancel each other if both increase or decrease.
Higher mortality-rates which favour large populations may result from
a fluctuating environment (Valentine 1971). Mortality is affected by nutrients,
temperature, and salinity changes. In general, invertebrates have higher
mortality-rates in the early stages of life, but the rate may be lower in some
species than others (Craig & Oertel 1966). Environmental conditions in estuaries
and lagoons would be expected to fluctuate widely and rapidly. They would
be expected to show a variable salinity range due to evaporation and influx
of fresh water, and a high diurnal temperature range. Furthermore, seasonal
upwelling of cold water along the Cape west coast leads to marked instability
of the environment on the open coast too.
Although growth-rate is an important factor in this type of study, it is
one of the attributes about which there is little information. There is evidence,
however, that most bivalves maintain a slightly decreasing, but nearly linear,
growth-rate throughout life (Craig & Oertel 1966). This has been shown to be
the case for Cardium edule, Tapes japonica, Dosinia exolata, and Venus striatula
(Kristensen 1959; Wilbur & Yonge 1964).
26 ANNALS OF THE SOUTH AFRICAN MUSEUM
The most likely effect of the unstable environmental characteristics of the
west and south coasts of the Cape would be a high mortality-rate among the
juvenile molluscs. Assuming a constant growth-rate for the bivalves, the inter-
play of mortality-rate and growth-rate should lead to a positively skewed
size-frequency distribution for a life assemblage, or an indigenous death
assemblage.
Besides the effect of growth-rate and mortality-rate on the shape of the
histogram, the assemblage may be affected by post-death mechanical change
such as sorting or winnowing by currents (Boucot 1953) and selective frag-
mentation, and solution of the smaller or thinner shells. Experience with the
west coast fossils shows that crushing and fracturing is of primary importance,
and affects the bivalves more than the gastropods.
The large number of fragmented shells in both the open-coast facies and
estuarine-lagoonal facies sediments suggests death assemblages that have
undergone considerable modification by wave-action. Whereas the open-coast
bivalves are usually disarticulated, those in the estuarine-lagoonal sediments
show a high degree of articulation. These sheltered embayment sites contain
epifaunal and infaunal molluscs, some of which are preserved in their living
positions. Size-frequency distributions for some of the bivalves are shown in
Figure 2. Since the warm-water element inhabited the Late Pleistocene lagoons
and estuaries, samples from those environments have been analysed in most
detail.
A single sample from a known high-energy open-coast site was examined in
detail. Figure 2A shows the size-frequency distribution for Venerupis senegalensis
from the Velddrif site. The field-setting suggests a transported death assemblage
in which vigorous wave-action piled shell debris up to form a breaker-bar.
In these deposits the thinner shells have generally been fragmented. The
V. senegalensis population is composed of thick-shelled forms. Its estuarine
ecomorph, on the other hand, has a thinner shell and is of more constant
morphology. The Velddrif V. senegalensis shows marked negative skewness
(—0,82). It has an articulation ratio less than 0,05. Articulation ratio is defined
as the ratio of complete shells/} (RV+LYV).
Size-frequency distributions of bivalves from the estuarine-lagoonal facies
differ from the pattern of the Velddrif example. Tellina madagascariensis from
Verlorevlei (Fig. 2C) and Churchhaven (Fig. 2H) tends to have a flattened
histogram. The Verlorevlei Tellina has a size-peak at 64-66 mm, and the
Churchhaven specimens at 56-60 mm. While the narrower size range of the
Verlorevlei material (50-86 mm) suggests a transported death assemblage, the
greater range of the Churchhaven material (16-74 mm; more juveniles) suggests
an indigenous death assemblage. That this argument can be misleading is shown
by their articulation ratios of 1,46 and 0,47 respectively. At both of these
localities 7. madagascariensis is associated with comminuted shells and dis-
articulated valves of other species. 7. madagascariensis at Verlorevlei was
observed in a nearly horizontal attitude as were the other bivalves. But whereas
THERMALLY ANOMALOUS LATE PLEISTOCENE MOLLUSCS
A
Venerupis senegalensis LV
Velddrif (9)
N=50
4 «18 22
e
26
30 34 38 42
Width (mm)
Tellina madagascariensis
Verloreviei (3)
50 54 58 62 66
18 Ee nm
16 Gastrana tadoa
$ Verloreviei (3)
N= 74
Loripes liratula
Kruispad (12)
N=71
Dosinia lupinus
Churchhaven (21)
18 22 26
42
30
RV
Art. rat. 146
70 74 78 82
LV
Art. rat. 256
46 50
RV
Art. rat (9.05
RV
Art. rat. Q13
34 382
Art rat. <g05
46
86
RV
Art. rat. 040
Dosinia lupinus
Verlorevlei (1)
&. 0% 6.22. 25 <u
Width (mm)
104 Dosinia lupinus RV
Verloreviei (3) Art. rat. Qlu
N=80
RV
Art. rat. <Q05
Kruispad (12)
N=71
6
No.
4
2
Gree 1g 18 22? 2p a0 34 38
14
12
10
Tellina madagascariensis LV
No81 Churchhaven(21) Art. rat. 0,47
6 N=57
4
2
1G; - 24, -32. AD 48, 6 56. G4 72
J
Loripes liratula
Knysna LV
8 N= 89 Art. rat. €9,05
16
14
12
No,!0:
8
6
4
2
3 5 7 9 11 Bie ai 7
Fig. 2. Size-frequency distribution of selected bivalve shells.
27
28 ANNALS OF THE SOUTH AFRICAN MUSEUM
most bivalves burrow with the shell vertical, Te//ina burrows rapidly and settles
in a horizontal position. For this reason the posterior end of the shell is strongly
flexed to the right so as to broaden the radius of curvature of the siphons to
minimize the current flow constriction. In a small quadrat at Verlorevlei
29 articulated shells were observed. Of these 26 had the right valve uppermost,
i.e. flexure upwards, and with the posterior end slightly raised. Many of the
T. madagascariensis shells still had the ligamental material attached across
both valves as a powdery residue. Clearly these animals must have died in their
life positions where the weight of sediment prevented opening of the shells
after relaxation of the adductor muscles. If they had opened after death and
subsequently closed again due to increased sediment load, the ligament would
have broken.
At Verlorevlei the 7. madagascariensis (site 3) size-frequency distribution
can be compared with those of Gastrana matadoa and Dosinia lupinus (Fig. 2B,
D, E). Dosinia lupinus at site 1 (Fig. 2B) has a positively skewed (+-0,37)
histogram with an articulation ratio of 0,40. The field occurrence suggests that
reworking of the sediment has taken place, although the shells are still situated
close to their original life habitat. Although the articulation ratio is high,
relative displacement of each valve is common. At site 3 the D. /upinus histogram
is bell-shaped (Fig. 2D). Here the articulation ratio is only 0,10. The bell-shaped
distribution suggests less winnowing of the small specimens. Both D. lupinus
populations peak at 22-24 mm. From site 1 to site 3 at Verlorevlei there is a
tendency for a greater spread of size-ranges of D. lupinus: 16-34 mm at site 1
and 640 mm at site 3. Conceivably the histogram for D. /upinus (site 1) was
originally bell-shaped but winnowing may have removed the smaller sizes.
This is borne out to an extent by the increase in relative proportions of Loripes
liratula in the assemblage, from 9 per cent at site 1 to 26,4 per cent at site 3.
The size-range of L. liratula 3-17 mm (Fig. 2G, J) coincides with the juvenile
fraction of D. Jupinus. Because of the close similarity in shape of D. lupinus
and L. liratula, equal-size specimens of each species would be expected to be
hydraulically equivalent.
The size-frequency distribution of Dosinia lupinus at Churchhaven has a
tail towards the larger specimens (Fig. 21), suggesting winnowing. It has a
narrow size-range, 14-42 mm, and peaks at 28-30 mm. The low articulation
ratio (0,13) suggests reworking.
At Kruispad the Dosinia lupinus population again has a wide size-range,
640 mm (Fig. 2F), but an articulation ratio less than 0,05. Furthermore, the
histogram has two peaks, at 16-18 mm and 28-30 mm. The bimodal distribution
could be explained by extinction of one living population, followed by fresh
recruitment when environmental conditions improved. The site is 15 km up
the Berg River and could possibly have been influenced by sudden influxes of
fresh water. But Loripes liratula at this site appears to represent a single popula-
tion: the histogram is bell-shaped and peaks at 12-13 mm. Its low articulation
ratio (< 0,05) may be meaningless since this species does not have prominent
THERMALLY ANOMALOUS LATE PLEISTOCENE MOLLUSCS 29
dentition, and seldom in this study were articulated valves encountered. For
example, Loripes liratula from Knysna has an articulation ratio less than 0,05
although juveniles dominate. Craig (1967) found that of five species he examined,
only Divaricella quadrisulcata tended toward a normal distribution. (Divaricella
and Loripes are both Lucinidae.)
Gastrana matadoa (Fig. 2E) at Verlorevlei is positively skewed and has a
high articulation ratio (2,56). This high articulation ratio, compared with that of
Dosinia lupinus from the same site, could be due to the more robust dentition
of the former, but the greater proportion of juveniles of G. matadoa suggests
it probably is closer to a life assemblage.
In general, the sign of skewness alone as used by Boucot (1953) is insufficient
to distinguish between indigenous and transported death assemblages. The
field-setting shows the Velddrif breaker-bar deposit to contain a transported
death assemblage. It has strong negative skewness and a low articulation ratio.
This tail towards the left has been produced by winnowing of the finer fractions,
and effectively displacing the mode towards the coarser size-grades. The other
size-frequency distributions are suggestive of indigenous death assemblages
which have in most cases been slightly reworked. In Figure 2B the lack of
juveniles of Dosinia lupinus coincides with low numbers of the hydraulically
equivalent Loripes liratula.
Other indications that the shells are found in the sediment in which they
once lived include relatively high articulation ratios of the larger shells. Smaller
shells, e.g. Loripes liratula, tend to have weaker hinge attachments, while
juveniles of other species may have been subjected to predation. Tellina
madagascariensis at Verlorevlei appears to be in a life orientation, and also
has the ligamental material still attached. At Kruispad a bed contains Panopea
glycymeris still in the life orientation. The shells were all articulated, all posterior
end upwards, and all on the same horizontal plane, i.e. they had all burrowed
a siphon-length below the sediment-water interface. At most sites, left and right
valves were present in equal proportions.
In conclusion, the field-setting shows that open-coast assemblages are all
transported death assemblages, but transportation has been only local. The
faunas of the sheltered environments are mixed life and indigenous death
assemblages. Taken as a whole the fauna of the open-coast facies and the
estuarine-lagoonal facies are consistent with the inferred Late Pleistocene
environments.
PREDATION
Many of the bivalve shells are punctured by countersunk borings, 1-2 mm
in diameter, made by predatory gastropods. The bored bivalves include: Dosinia
lupinus, Tellina madagascariensis, Gastrana matadoa, Venerupis senegalensis,
Venerupis dura, and Ostrea algoensis. Crepidula capensis and Natica genuana
were the most frequently bored gastropods.
At Churchhaven 17 per cent of Dosinia lupinus and 17 per cent of Tellina
30 ANNALS OF THE SOUTH AFRICAN MUSEUM
madagascariensis were bored. In the case of the former an equal number of
right and left valves were bored, as would be expected in view of the fact that
it burrows in a vertical position. Analysis of predation on 7. madagascariensis
reveals a very different pattern. Successfully bored valves fell into two size-
classes (Fig. 2H). The first group comprised juveniles with a size range
20-22 mm, and the second group adults with a size range 46-64 mm. Only
right valves of adults were bored, which is to be expected because this species
lives buried in sediment in a horizontal position and with right valve uppermost.
Craig (1967) found that the right valve of Tellina radiata was also preferentially
bored. The second group of T. madagascariensis comprised juveniles in which
left valves were preferentially bored, although a few right valves were also
bored. The preferential boring of juvenile left valves could also be the result
of the burrowing characteristics. Tellina is a rapid burrower (Stanley 1970).
Depth of burrowing would be controlled by length of the siphons, and for this
reason the juveniles would presumably live at shallower depths than the adults
and would possibly burrow more slowly. Subsequent current scour would
possibly reach only the juveniles and flip them over to leave them left valve
uppermost. This also suggests that, besides predation, the juvenile bivalves
would be more susceptible to environmental changes since they live closer
to the surface.
Most of the bored bivalve shells have only one hole, which is not surprising
since only one puncture is necessary to kill the animal. In a few cases two holes
per shell were encountered. As far as Tellina madagascariensis is concerned
the borings occur most frequently in the antero-dorsal half of the shell (Fig. 3).
It was also observed that all bored specimens of T. madagascariensis at Church-
haven consisted of separate valves, while four bored articulated shells of Dosinia
lupinus were observed.
Of the gastropods, Crepidula capensis was the most extensively bored. Again
there was usually only one hole per shell, although up to three were recorded.
The reason why Crepidula capensis is so susceptible to predation is because
it lives exposed at the surface where it is an easy prey. Unlike the bivalves,
Crepidula does not appear to have any preferred area for boring (Fig. 3).
Fig. 3. Predation: distribution of borings on Tellina madagascariensis
(left), and Crepidula capensis (right).
THERMALLY ANOMALOUS LATE PLEISTOCENE MOLLUSCS 31
Crepidula capensis from the shell beds north of Laaiplek have smaller bored
holes (less than 1 mm) than other C. capensis or the bivalves.
It is difficult to isolate the species that would have been the predator,
although it seems probable that the borings were made by a gastropod. The
most likely predator would appear to be the buccinid Burnupena papyracea.
Where many bivalves were found to be bored at Churchhaven, B. papyracea was
very prominent. At Verlorevlei there was a total absence of B. papyracea and
no sign of predation on bivalves.
TERATOLOGICAL SPECIMENS
Teratological specimens are those that are outside the normal range of
variation of a species (Ager 1963). At most sites an occasional aberrant form
was observed, and it was found generally that Gastrana matadoa was the most
susceptible to damage during life. This was very apparent at Verlorevlei (site 3)
where 27 per cent of G. matadoa were in some way deformed, compared with
less than 2 per cent for Te/lina madagascariensis. Perhaps this reflects the degree
of adaption of G matadoa to its niche, and that the environment at Verlorevlei
in particular did not favour this species.
DEPTH
Regional geomorphic analysis suggests that the depth of water reflected
in the strata of all open-coast and sheltered embayment sites examined could
not have exceeded 5 m. The Late Pleistocene estuarine-lagoonal facies is
dominated by intertidal deposits, and the open-coast facies by beach deposits.
At Velddrif there is a good exposure of a breaker-bar with a washover-fan
the top of which is probably a close approximation to high water spring tide.
At Churchhaven and Kraalbaai Callianassa burrows and crab-burrows are
still preserved, indicating an intertidal or shallow subtidal environment. The
ostracod fauna, too, is indicative of intertidal conditions.
Very few of the 124 species listed in Table 1 suggest water depths greater
than 25 m. Most of these species live today in intertidal and the uppermost
sublittoral zones. Less than 4 per cent of the fauna lives today in water deeper
than 25 m.
Of the open-coast assemblages Nassarius speciosus is today usually dredged,
although it is sometimes found intertidally. No more than a few isolated shells
were found. Erycina subradiata has been encountered at 26 m (Barnard 1964).
Other species have a wide depth range although they are common in the inter-
tidal zone. Nearly all species of Patella encountered at open-coast sites are
intertidal. Only P. miniata, P. tabularis, P. compressa are infratidal (Branch
1971). Other intertidal molluscs include: Littorina knysnaensis (above
HWN), Marginella capensis (low tide down), Bullia digitalis (follows the tide),
B. laevissima, Thais cingulata, Burnupena papyracea (below mid-tide), Perna perna
(mid-tide), Aulacomya ater (mid-tide to low tide on rocky shores), Choromytilus
meridionalis, Dosinia lupinus, Donax serra (burrows in surf beaches below
32 ANNALS OF THE SOUTH AFRICAN MUSEUM
mid-tide), Venus verrucosa (found on the surface at mid-tide). Depth ranges
are given by Day (1969).
Of the estuarine-lagoonal facies molluscs Ostrea algoensis prefers depths of
25-200 m, Tellina ponsonbyi prefers off-shore waters to depths of 95 m, and
Theora alfredensis has been dredged from depths below 70 m (Barnard 1964).
Ostrea algoensis and Theora alfredensis are common in this facies. Only one
Tellina ponsonbyi valve was found at Churchhaven, suggesting that this specimen
strayed into shallow water.
Most of the sheltered-embayment molluscs prefer very shallow or intertidal
water. These include the gastropods Patella spp., Littorina knysnaensis (HWN),
Turritella capensis (quiet water shallower than 3 m), Oxystele variegata,
Burnupena papyracea, Nassarius kraussianus (mud-banks and weed-beds of
estuaries); the bivalves Loripes liratula (muddy sand at low tide), Solen capensis
(muddy sand of estuaries), Venus verrucosa, Choromytilus meridionalis, Perna
perna, Aulacomya ater, Tellina madagascariensis (extensive infratidally), Dosinia
lupinus, Donax serra and Panopea glycymeris.
None of the constituents of the open-coast or estuarine-lagoonal facies
required water depths greater than about 25 m. Modern bathymetric ranges for
the species suggests maximum Late Pleistocene water depths at these sites of the
order of 0-5 m. This is consistent with the geomorphic evidence.
TEMPERATURE
Since the primary purpose of this paper is a discussion of a southerly
migration of tropical and subtropical mollusc species in the last interglacial,
the most important criteria are those indicative of temperature. Sea surface
temperature inferences may be made by comparing geographic ranges of living
molluscan species with those of their fossil counterparts. Although the fossil
fauna of the west coast is distinctly modern in character, the estuarine-lagoonal
facies assemblages regularly contain several species whose present-day dis-
tribution is restricted to tropical and subtropical waters. The modern geographic
range end-points of the thermophilic species that live further north on the west
African coast are separated by 2 000 km from the fossil occurrences in the
south-western Cape, so that the fossil assemblages contain species that do not
today live in association. Northward migration of the extralimital species was
probably induced by deteriorating environmental conditions. Since the geo-
graphic ranges of recent molluscs seem to be determined mainly by temperature
(Valentine 1955), it would appear that falling temperature was the compelling
factor. It could also be argued that lower temperatures accompanied the advance
of the Weischelian ice-sheets, in which case falling sea level would have drained
the Late Pleistocene estuaries and lagoons. But with reflooding of these
environments in Recent time, the thermophilic molluscs did not return.
Mollusc assemblages in the estuarine-lagoonal facies of the west and
south coasts contain a total of more than 20 species that do not sustain
populations in those areas today. West coast sites contain 15 such species,
33
THERMALLY ANOMALOUS LATE PLEISTOCENE MOLLUSCS
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ANNALS OF THE SOUTH AFRICAN MUSEUM
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THERMALLY ANOMALOUS LATE PLEISTOCENE MOLLUSCS 35
mainly bivalves. These extralimital molluscs, their present-day geographic
ranges, and minimum temperature tolerances are shown in Table 2.
It is often difficult to assess the minimum temperature tolerance of living
molluscs. Zinsmeister (1974) divides the normal biogeographic range of a
taxon into a spawning range, and a non-spawning range. The modern geographic
range of Ostrea atherstonei, from Saldanha to Bushmans River, implies’ a
minimum temperature tolerance of about 13°C. But it appears that present
temperatures in Saldanha Bay are too low for spawning (Korringa 1956).
Spawning experiments in this area have confirmed this, and suggest that the
oyster larvae may actually originate in an area of higher water temperature.
It is possible that several species of molluscs extend their geographic range
end-points by inhabiting local pockets of warm water such as estuaries. If this
is the case then the temperature minima shown in Table 2 may actually be too
low. It is not always clear from the literature whether the range end-points
are open-coast, estuaries or embayments. Other difficulties arise from taxonomic
problems. For instance, Macoma ordinaria and M. crawfordi may be geographic
variants of the Mediterranean M. cumana.
1. West coast estuarine-lagoonal facies
Because of physiographic changes at Verlorevlei and the lower reaches
of the Berg River, present temperature ranges would be meaningless standards
against which to measure Late Pleistocene changes. The present shoreline of
Langebaan Lagoon has changed little since the last interglacial, and present
temperatures at Churchhaven could be used as an approximation for the other
sites as well. Day (1959) gives a surface temperature range for Churchhaven
of 13,5°C to 37°C (HW) and 38,5°C (LW).
The fossil assemblage at Verlorevlei (site 3) is represented by 23 species
of molluscs, 6 of which are indicative of warm water. These 6 thermophilic
species constitute 42 per cent of the total individuals. The modern geographic
range of Loripes liratula (26,4 per cent of the fauna) is Mauritania to Angola
(Nicklés 1950), where its minimum temperature requirement would be 17-18°C.
Ostrea stentina (2,7 per cent), Tellina madagascariensis (8,1 per cent), Macoma
tricostata (< 1 per cent), and Venerupis dura (< 1 per cent) are also tropical
west African species and could not tolerate temperatures below about 17°C.
Gastrana matadoa (4,1 per cent) extends into water as cool as 14°C. Accepting
that stenothermal organisms are more critically limited by minimum tem-
peratures than by maximum temperatures (Durham 1950), it would appear
that the minimum temperature in this Late Pleistocene estuary must have been
17-18°C, or at least 3°C warmer than present Churchhaven surface temperatures.
At Kruispad (site 12) 8 of the 31 species prefer warm water, and constitute
47,9 per cent of the total individuals. Of these Nuculana bicuspidata (3,2 per cent),
Loripes liratula (15,4 per cent), and Leporimetis hanleyi (1,0 per cent) have a
minimum temperature requirement of 17-18°C. A significant constituent of
this assemblage is Panopea glycymeris. Although it forms less than 1 per cent
36 ANNALS OF THE SOUTH AFRICAN MUSEUM
of the total individuals, it is nevertheless very common (because it is a large
animal, about 200 mm, and is always found in life orientation, a shell count in
a 1 m quadrat would be unlikely to yield more than one or two specimens).
Panopea glycymeris lives in areas with a temperature range 20-27°C (Kensley
1974), although it could probably tolerate cooler water. This assemblage is thus
indicative of a temperature minimum of about 18°C. At Bloemendal (site 11)
Loripes liratula and Leporimetis hanleyi are common.
Three sites examined along the shore of Langebaan Lagoon yielded a
significant proportion of thermophilic molluscs: Geelbek (site 19; 7 species in
26 constituting 46,0 per cent of the fauna), Skrywershoek (site 20; 6 species in
19 constituting 32,9 per cent of the fauna), and Churchhaven (site 21/22;
7 species in 31 constituting 28,1 per cent of the fauna in the lower unit; 7 species
in 27 constituting 35,5 per cent of the fauna in the upper unit). The dominant
thermophilic mollusc content at these sites is as follows:
Geelbek Skrywershoek — Churchhaven
Lower Upper
7 Zo Zo 7
Ostrea atherstonei am | 1,0 5,0
Loripes liratula 20,4 14,1 1,0 4,1
Mactra ovalina 1,0 ee ZS if
Tellina madagascariensis 10,2 9,4 23,4 33
Leporimetis hanleyi _ < 1,0 —- —
Venerupis dura 4,1 12 — —
All of these species, except Ostrea atherstonei and Mactra ovalina, have a
minimum temperature requirement of 17°C. Ostrea atherstonei is found living
today in Saldanha Bay, but temperatures are apparently too low for spawning.
It is possible that this species has a minimum temperature tolerance of 14°C.
The floor of the present lagoon is underlain by about 3 million metric tons of O.
atherstonei shells, with little detrital sediment, suggesting optimum temperatures
for breeding, and hence probably greatly in excess of 14°C. Mactra ovalina
has a minimum requirement of 19°C. A minimum temperature in the Late
Pleistocene lagoon of about 18°C is indicated, about 4-5°C warmer than the
present surface temperature minimum at Churchhaven.
2. South coast estuarine—lagoonal facies
A Late Pleistocene site at Knysna was the only one on the south coast
examined in detail. But examination of material in collections of the South
African Museum shows that very similar fossil faunas exist at Sedgefield, and
Groot Brak and Klein Brak estuaries. The fossil assemblage at the Klein Brak
estuary contains Panopea glycymeris, but their shells are, on average, much
smaller than the Kruispad specimens (Kensley 1974).
THERMALLY ANOMALOUS LATE PLEISTOCENE MOLLUSCS 37
The Knysna assemblage contains 47 molluscan species, of which 6 are
extralimital and which constitute 60,4 per cent of the individuals. But these 6
species include the extinct subspecies Cantharidus suarezensis fultoni (9,8 per
cent) and Cerithium scabridum rufonodulosum (16,3 per cent). Although the
geographic ranges of the species is shown in Table 2, it is obviously not certain
that the extinct subspecies had the same temperature tolerances. Their Recent
relatives Cantharidus suarezensis suarezensis and Cerithium scabridum have
minimum temperature requirements of 19°C and 24°C respectively. Atys
cylindrica (< 1 per cent) has a minimum temperature requirement of 21°C.
Of the bivalves Felania diaphana (3 per cent), Tellina madagascariensis (< 1 per
cent), Leporimetis hanleyi (< 1 per cent) and Venerupis dura (< 1 per cent) all
suggest temperature minima of 17°C. Loripes liratula (57,5 per cent) also
suggests a temperature minimum of 17°C, but the fact that it occurs in such
great numbers, and the fact that all growth stages are present (Fig. 2J), suggest
optimum conditions. Together the evidence suggests a temperature minimum
in excess of 17°C for the Late Pleistocene estuary. Day ef al. (1952) gives a
temperature range at the railway bridge of 12-24°C.
3. West coast open-coast facies
The present-day temperature range on the adjacent open coast is about
13-15°C (Shannon 1966). The fossil assemblage contains only three species
which prefer warm water, but which never constitute more than | per cent of
any assemblage: Cypraea algoensis, Marginella piperata and Scissodesma
spengleri. The marked paucity of warm-water species indicates that Late
Pleistocene nearshore water temperatures were not very different from the
present.
SUMMARY
Before discussing the palaeoclimatic significance of the extralimital
molluscan species it may be as well briefly to summarize the evidence.
1. All fossiliferous Late Pleistocene estuarine-lagoonal facies deposits
contain extralimital species which usually constitute more than 30 per cent
of the individuals of each assemblage, and which indicate minimum water
temperatures 4-6°C warmer than the present-day estuaries and lagoons, with
minimum temperatures of about 18°C. Kanakoff & Emerson (1959) suggest
temperatures in excess of 19°C for similar Californian occurrences.
2. All growth stages of the thermophilic molluscs are present (Fig. 2).
The importance of this is that it implies that temperatures were such that the
normal spawning range of each mollusc is represented. Normal spawning range
is characterized by a population which continually maintains its numbers
(Zinsmeister 1974). The large number of individuals and presence of all growth
stages show that the thermophilic taxa formed self-sustaining populations that
were adequately adapted to the depositional environment.
3. The estuarine-lagoonal facies contain mixed life and indigenous death
38 ANNALS OF THE SOUTH AFRICAN MUSEUM
assemblages of molluscs. These molluscs obviously lived in environments
reflected in the sediments, and have suffered little post-death transportation.
4. The sediments and their mollusc fossils indicate water depths in general
less than 5 m.
5. The fact that sediments with high proportions of extralimital molluscan
species characterize most Late Pleistocene estuary and lagoonal situations,
that these molluscs formed self-sustaining populations and inhabited very
shallow water, suggests that the marine transgression to 6 m in the last inter-
glacial was a major event during a climatic optimum.
Zinsmeister (1974) has suggested that similar, well documented occurrences
of thermophilic molluscs in Californian Late Pleistocene embayments represent
only temporary, non-breeding, members of the community. He believes that
periodic local and temporary current changes introduced tropical mollusc larvae
into areas of cooler water, and that these molluscs represent only temporary
members of the community since temperatures would have been too low for
them to maintain self-sustaining populations.
Zinsmeister’s arguments stem from the fact that previous studies paid
scant attention to the population dynamics, and consequently they fail to
prove the presence of self-sustaining populations. Similar occurrences in the
south-western Cape support the view that these faunas are the result of a warmer
climate which must have affected the whole world. Age-wise these deposits
are also similar and probably coincide with a high climatic peak (substage Se)
and sea level up to 7 m higher than present at 120 ka (Shackleton 1969).
6. Configuration of the coastline was probably very important. Today
a low Palaeozoic rock bar just above high-tide level keeps the sea out of
Verlorevlei. A slight rise of sea level would create radical changes at Verlorevlei.
Likewise the lower reaches of the Berg River formed a prominent estuary due
to flooding by last interglacial high sea levels. Normally one would expect to
find an overlap of geographic ranges of tropical and temperate species. In
an overlap area the tropical species would be restricted to inshore, protected
environments, and temperate species would live in the cooler open coast sites
(Emerson 1956).
PALAEOCLIMATIC INTERPRETATION
REVIEW OF PRESENT CLIMATE AND HYDROLOGY
The south-western Cape has a Mediterranean-type climate. Hot, dry
summers are the result of the dominant anticyclones in those months, while
depressions associated with westerly winds bring rainfall in winter.
Hart & Currie (1960) have summarized the effect of the wind system on
the climate of the south-western Cape Province. A subtropical high-pressure
system is centred between 26° and 30°S. To the south this high-pressure system
borders on the ‘westerlies’ causing a steep pressure gradient. The south-easterly
winds of the south-western Cape are the result of winds blowing anticyclonically
THERMALLY ANOMALOUS LATE PLEISTOCENE MOLLUSCS 39
around this high-pressure system. In summer the centre of the anticyclone lies
at about 30°S and brings strong south-easterly winds to the south-western
regions, but in winter it moves northwards to 26°S. The westerly wind system
follows the anticyclone northward and the southern Cape is then frequented
by depressions which bring rain from the south-west Atlantic, although the
Namib Desert is still influenced by the Trade Wind belt.
The west coast of southern Africa, like the west coasts of other countries
in similar latitudes, is characterized by a linear belt of centres of upwelling of
cold subsurface water (Central Water) from Liideritz to the Cape Peninsula
(Fig. 1 insert). There is a north-south isotherm lineation with the coldest and
least saline water near the coast (Shannon 1966). This upwelling, the Benguela
Current, varies in intensity depending upon the wind system and local
topography. The upwelling phenomenon arises from the displacement of
surface water northwards and off-shore by the south-easterly wind system.
Cooler subsurface water wells up to replace this warmer water. The result
of this active upwelling is a complex system with tongues of cold water
alternating with intrusions of warmer oceanic waters, and all diverging to the
north-west (Bang 1971). Bang defines the Benguela Current as the area east of a
belt of off-shore divergence within which the oceanic processes are dominated
by short-term atmospheric interactions.
During the winter months when the anticyclone centre moves northward,
weakening of the southerly wind component results in weakening of the
upwelling system also. With less upwelling the surface waters over the inner
part of the continental shelf are warmer by 2°C, but off-shore a drop in
temperature tends to minimize this affect. With weakening of the Benguela
Current system and the greater prominence of westerly and north-westerly
winds, a southward-flowing inshore counter-current develops. Shannon (1966)
mentions a predominantly southward-flowing counter-current between
Lambert’s Bay and Cape Point which is present during all seasons, but most
marked in winter. A southward-flowing counter-current carries ‘seeboontijies’
from Angola and driftwood from the Orange River southwards (Wagner &
Merensky 1928). Surface temperatures at the Orange mouth rise noticeably
when the north-westerly winds blow.
The highest surface-water temperatures are encountered in the lagoons
and estuaries which are protected from the effects of upwelling. The temperature
range in Langebaan Lagoon, 10-39°C (Day 1959), contrasts markedly with
that of the near-by open sea, 13-15°C (Shannon 1966). During the summer
months temperatures in Langebaan Lagoon are at a maximum, while those
of the open sea drop.
The currents off the east and south coasts, the Mozambique Current and
Agulhas Current, have been studied by Clowes (1950), Orren (1963, 1966),
and Darbyshire (1964). Both of these currents are southward extensions of the
great South Equatorial Current which flows westwards across the Indian Ocean.
At 26°S the Mozambique Current is met by the southern branch of the South
40 ANNALS OF THE SOUTH AFRICAN MUSEUM
Equatorial Current, which is divided by Madagascar, and they combine to
form the Agulhas Current. The Agulhas Current is finally deflected by the
Agulhas Bank (Clowes 1950) (Fig. 1).
Upwelling does take place off Cape Agulhas but it is a geostrophic
upwelling, i.e. the upwelling varies with the velocity of the current. Here the
Agulhas Current is weakest and Agulhas water retreats northwards in winter,
thus allowing Central Water to reach the surface. In summer, when the Agulhas
Current is flowing at its strongest, it overrides the Central Water (Darbyshire
1964). Sometimes in summer the warm Agulhas Current moves away from the
south coast and allows cold Central Water to replace it inshore, and within
a day or two the temperature may fall by as much as 10°C (Day 1963). Schell
(1968) describes the occasional penetration of the Agulhas Current round the
Cape Peninsula into the South Atlantic.
LATE PLEISTOCENE CLIMATE
It would be expected that the intensity, and probably position, of the
anticyclones would vary with the solar radiation. A climatic optimum at 120 ka
produced temperatures warmer than at any other time in the last 120 ka
(Shackleton 1969). The anticyclone would have moved south of latitude 30°S
(Van Zinderen Bakker 1967: fig. 6). This would lead to the linear west coast
upwelling belt moving south in response, and the winter-rainfall area would
be more strictly restricted to the south-western Cape. Such a southward move-
ment would be accompanied by more pronounced upwelling off the west
coast than at present, and would operate over a longer period of time just
as happens on the Namib Desert coast. Since the Central Water originated
in the Southern Ocean, and since the Antarctic ice-sheet was essentially stable
throughout the Pleistocene (Mercer 1968), it is unlikely that the upwelling water
would have been any colder than the present. This is confirmed by the fossil
molluscs from the open-coast facies which suggest little temperature change.
Addicott & Emerson (1959) and Valentine (1955) have a!so suggested intensified
nearshore upwelling and a poleward expansion of isotherms to explain their
molluse faunas.
Eustatic rise of sea level would have changed the configuration of the
coast by forming salt-water estuaries at Verlorevlei and the Berg River, as
well as at numerous sites along the south coast. The present coast is not as
embayed as the Late Pleistocene coast would have been. Thermally anomalous
assemblages existed contemporaneously only in the vicinity of the protected
bays where the increased solar radiation of the last interglacial heated the
surface water considerably.
Periodic current changes permitted the introduction of tropical mollusc
larvae into these pockets of warm water. The intensification of atmospheric
circulation that led to more pronounced upwelling presumably also affected
the inshore counter-current which possibly flowed more strongly than at present.
Isaacs & Sette (1959) described anomalous wind fields in the Pacific area during
THERMALLY ANOMALOUS LATE PLEISTOCENE MOLLUSCS 4]
1957 and 1958 which so changed the oceanic circulation that tropical taxa
were found far north of their expected ranges. This demonstrates that circulation
changes as proposed in this paper do happen at present, albeit less frequently
and less pronounced than is here suggested.
On the east coast with intensification of the air circulation intensification
of the present summer conditions would also be anticipated. Presumably the
Agulhas Current flowed more strongly than today and continually overrode
the Central Water, perhaps with frequent eddies of Agulhas water around the
Cape Peninsula. This would limit the likelihood of sudden incursions of cold
water, such as presently affect the Knysna estuary in summer and cause
temperatures to drop between 10 and 15°C (Korringa 1956).
These possible changes in the oceanic circulation during the hyperthermal
period would have had far-reaching effects on the distribution of molluscs.
Because of the low migratory ability of molluscs in the post-larval stage, their
geographic distribution depends primarily on the dispersion of larvae. Distribu-
tion of the planktonic larvae depends upon ocean currents. About 85 per cent
of tropical marine molluscs have a free-swimming pelagic larval stage which
can exist, on average, three to four weeks before settling (Zinsmeister 1974).
Dietrich (1935 in Korringa 1956) gives the velocity of the Agulhas Current as
50 km per day. At this rate larvae could be transported 1 000 km before
metamorphosis. Larvae can apparently withstand sudden changes of
temperature; Korringa mentions a change from 25° to 2°C which did not
adversely affect oyster larvae.
A general southward movement of isotherms on the west and south coast is
envisaged. This would have brought the tropical mollusc zone closer to the south-
western Cape. Periodic expansions of the warm-water isotherms coupled with an
intensified inshore counter-current on the west coast would have enabled
tropical mollusc larvae to pass the cold Benguela barrier. If the larvae reached
the sheltered estuaries and lagoons of the south-west coast, increased solar
heating (substage Se) would have allowed these molluscs to sustain their
populations and persist there even after the open-coast thermal barrier became
impassable again. Those thermophilic larvae that reached metamorphosis
on the open coast may well have survived, but there is no evidence that they
were able to sustain their populations. South coast estuarine facies show that
larvae from the tropical west African coast were able to pass the Cape Pen-
insula: e.g. Tellina madagascariensis, Loripes liratula, Leporimetis hanleyi,
Venerupis dura, Panopea glycymeris. The Agulhas Current was also able to
transport Indo-Pacific mollusc larvae around the Peninsula to the west coast
sites. In these sheltered embayments warm-water taxa survived as relicts.
DISCUSSION
The occurrence of thermophilic molluscs in Late Pleistocene sediments
of the south-western Cape far beyond their present-day geographic range
end-points is not unique. Similar occurrences in California have been extensively
42 ANNALS OF THE SOUTH AFRICAN MUSEUM
studied, and the same conditions appear to have operated in the Miocene
where the configuration of the coastline was very different from the present.
The San Joaquin basin was a protected embayment in the Late Miocene, and
there relict faunas persisted long after the temperate faunas had spread south-
ward along the open coast (Addicott & Vedder 1963). The thermally anomalous
Late Pleistocene molluscs also lived in shallow protected embayments (Addicott
& Emerson 1959). The only significant point of difference between the two
regions is that the open-coast facies of the Cape coast contains molluscs which
nearly all live on the adjacent coast, while the Californian open-coast facies
contains a fauna that reflects cooler water than the present. Valentine (1955)
proposed that during the last interglacial of California upwelling was intensified,
while at the same time the warmer water of the sheltered embayments was
derived from increased solar radiation. He also suggested a general warming
of the oceanic waters by increased solar heating.
‘ The Late Pleistocene estuarine-lagoonal facies of the Cape Province are
attributed to a eustatic rise of sea level which changed the configuration of the
coast and produced a great number of estuaries where the effects of seasonal
upwelling were excluded. A southward movement of the south Atlantic anti-
cyclone at this time shortened the cold-water barrier on the western open
coast by causing a southward shift of the Benguela Current, and possibly
strengthened the inshore counter-current, so that thermophilic mollusc larvae
were able to migrate southwards where they could sustain their populations
in the solar-heated estuaries and lagoons. It would appear that the present
temperatures are too low for breeding.
This history contrasts with that of the Early Pleistocene when sea
temperatures on the open coast were much warmer than at present. The last
of these warm-water open-coast episodes is associated with the 45-50 m trans-
gression complex of the Namaqualand coast (Carrington & Kensley 1969).
Striostrea margaritacea was common on the open coast, forming the so-called
‘oyster line’ (Haughton 1931). This oyster requires a minimum water temperature
of 25°C in summer (Korringa 1956). The 45-50 m transgression complex
sediments were probably pre-glacial Pleistocene. Such warm conditions must
have been in response to the warmer conditions prevailing in the northern
hemisphere, since the Antarctic ice-sheet is thought to have been stable through-
out the Pleistocene (Mercer 1968). Warming of the northern hemisphere would
have moved the intertropical convergence farther south than even its Late
Pleistocene position. This would have moved the South Atlantic anticyclone
south, and with it the belt of upwelling, bringing tropical waters down the
Namaqualand coast. But Striostrea margaritacea is totally absent from Early
Pleistocene deposits in the Saldanha area, suggesting that cool water, and
upwelling, were still dominant there.
If the model for a southward shift of the anticyclonic system in the last
interglacial is correct, then one would expect the opposite trend during the
Weischelian. Lamb (1961) suggests that Ice Age circulation was marked by
THERMALLY ANOMALOUS LATE PLEISTOCENE MOLLUSCS 43
intensified circulation of the belt of westerlies, and greater mobility in the
subtropical anticyclones, which generates upwelling systems such as the Benguela
Current. Van Zinderen Bakker (1967, in press) presents evidence to show that
during hypothermal periods the influence of the anticyclones and the Benguela
Current would have shifted northwards to the equatorial regions. Displacement
of the westerlies towards lower latitudes would have expanded the winter-
rainfall area of the southern Cape, and would have caused a northerly shift
of the Namib Desert. Tankard & Schweitzer (1974) have demonstrated, from
Die Kelders cave sediments, that wetter conditions in the Weischelian coincided
with a cooler period. Butzer (1973) has found a similar record in the Nelson
Bay cave. A northward migration of the belt of westerlies would have resulted
in a longer rainy season, if not year-round precipitation.
Finally, this study suggests a possible correlation of the last interglacial
deposits of the Cape Province with the Eutyrrhenian of the Mediterranean.
Although warping of the Eutyrrhenian shorelines is universal (Richards 1962),
Bonifay & Mars (1959) have attempted to restore these shorelines to their
original elevations. They attribute the Eutyrrhenian shoreline to a transgression
to 2-3 m a.s.l. Chronologically the Eutyrrhenian shoreline and last interglacial
shorelines of the south-western Cape appear to be similar.
The Strombus fauna of the Eutyrrhenian deposits is characterized by
Strombus bubonius and other species typical of Senegal and west Africa
(Richards 1962) which suggest warmer hydroclimates than today. Like the
warm water fauna of the south-western Cape, the Strombus fauna can also be
attributed to poleward expansion of isotherms, and once in the Mediterranean
the molluscs remained a relict fauna which survived in water heated by increased
solar radiation.
ACKNOWLEDGEMENTS
The writer wishes to thank Mr R. N. Kilburn for his assistance in
identifying some of the molluscs and Mr V. Branco who assisted with the
illustrations. This research project was supported by a grant from the
5.1.R.
REFERENCES
AppicotTrT, W. O. & EMERSON, W. K. 1959. Late Pleistocene invertebrates from Punta Cabras,
Baja California, Mexico.—Am. Mus. Novit. 1925: 1-33
AppicoTt, W. O. & VEDDER, J. G. 1963. Paleotemperature inferences from late Miocene
mollusks in the San Luis Obispo—Bakersfield area, California. — Prof. Pap. U.S. geol. Sury.
475C: 63-68.
Acer, D. V. 1963. Principles of Paleoecology. New York: McGraw-Hill.
BANG, N. D. 1971. The southern Benguela Current region in February, 1966: Part II. Bathy-
thermography and air-sea interactions. — Deep-sea Res. 18: 209-224.
BARNARD, K. H. 1964. Contributions to the knowledge of South African marine Mollusca.
Part V. Lamellibranchiata.— Ann. S. Afr. Mus. 47: 361-593.
BoniFAy, E. & Mars, P. 1959. Le Tyrrhénien dans le cadre de la chronologie quaternaire
méditerranéenne. — Bull. Soc. géol. Fr. (7) 1: 62-78.
a4 ANNALS OF THE SOUTH AFRICAN MUSEUM
Boss, K. J. 1969. The subfamily Tellininae in South African Waters (Bivalvia, Mollusca).— Bull.
Mus. Comp. Zool. 138: 81-162.
BoucotT, A. J. 1953. Life and death assemblages among fossils.—Am. J. Sci. 251: 25-40.
BRANCH, G. M. 1971. The ecology of Patella Linnaeus from the Cape Peninsula, South Africa.
1. Zonation, movements and feeding.— Zoologica Afr. 6: 1-38.
BROECKER, W. S., THURBER, D. L., GODDARD, J., Ku, T-L., MATTHEWS, R. K. & MESOLELLA,
K. J. 1968. Milankovitch hypothesis supported by precise dating of coral reefs and deep-
sea sediments.— Science 159: 297-300.
Butzer, K. W. 1973. Geology of Nelson Bay Cave, Robberg, South Africa.—S. Afr. archaeol.
Bull. 28: 97-110.
CARRINGTON, A. J. & KENSLEY, B. F. 1969. Pleistocene molluscs from the Namaqualand
coast.— Ann. S. Afr. Mus. 52: 189-223.
CLowes, A. J. 1950. An introduction to the nadihony of South African waters.—Investl.
Rep. Fish. Mar. biol. Sury. Div. Un. S. Afr. (12): 1-42.
CraiG, G. Y. 1967. Size-frequency distributions of living and dead populations of pelecypods
from Bimini, Bahamas, B.W.I.—J. Geo/. 75: 34-45.
CralGc, G. Y. & HALLAM, A. 1963. Size-frequency and growth-ring analyses of Mytilus edulis
and Cardium edule, and their palaeoecological significance.— Palaeontology 6: 731-750.
Craic, G. Y. & OERTEL, G. 1966. Deterministic models of living and fossil populations of
animals.— Q. J. geol. Soc. Lond., 122: 315-355.
DARBYSHIRE, J. 1964. A hydrological investigation of the Agulhas Current area.— Deep-sea
Res. 11: 781-815.
Day, J. H. 1959. The biology of Langebaan Lagoon: a study of the effect of shelter from wave
action. — Trans. r. Soc. S. Afr. 35: 475-547.
Day, J. H. 1963. The complexity of the biotic environment.—Publs. Syst. Ass. (5): 31-49.
Day, J. H. 1969. A guide to the marine life of South African shores. Cape Town: A. A.
Balkema.
Day, J. H., MILLarp, N. A. H., & Harrison, A. D. 1952. The ecology of South African
estuaries. Part III, Knysna: a clear open estuary.—Trans. r. Soc. S. Afr. 33: 367-413.
DuruHaM, J. W. 1950. Cenozoic marine climates of the Pacific coast. — Bull. geol. Soc. Am. 61:
1243-1264.
EmersSON, W. K. 1956. Pleistocene invertebrates from Punta China, Baja California, Mexico.
— Bull. Am, Nat. Hist. 111: 313-342.
EMERSON, W. K. & Cuase, E. P. 1959. Pleistocene mollusks from Tecolote Creek, San Diego,
California.— Mem. S. Diego. Soc. nat. Hist. 12: 335-346.
GRANT, U.S. IV & GALE, H.R. 1931. Catalogue of the marine Pliocene and Pleistocene Mollusca
of California and adjacent regions. — Mem. S. Diego Soc. nat. Hist. 1: 1-1036.
HALLAM, A. 1960. A sedimentary and faunal study of the Blue Lias of Dorset and Glamorgan.
—Phil. Trans. r. Soc. Lond. B 243: 1-44.
Hart, T. J. & Currie, R. I. 1960. The Benguela Current.— Discovery Rep. 31: 127-297.
HAUGHTON, S. H. 1931. The late Tertiary and Recent deposits of the west coast of South
Africa.—Trans. geol. Soc. S. Afr. 34: 19-57.
Isaacs, J. D. & Sette, O. E. 1959. Unusual conditions in the Pacific. — Science 129: 787-788.
KANAKOFF, G. P. & EMERSON, W. K. 1959. Late Pleistocene invertebrates of the Newport
Bay area, California.— Contr. Sci. 31: 1-47.
KENSLEY, B. F. 1973. Sea-shells of southern Africa—gastropods. Cape Town. Maskew Miller.
KENSLEY, B. F. 1974. The status of the Plio-Pleistocene Panopea in Southern Africa (Mollusca,
Bivalvia, Hiatellidae).—Ann. S. Afr. Mus. 65: 199-215.
KERN, J. P. 1971. Paleoenvironmental analysis of a Late Pleistocene estuary in southern
California.—J. Paleont. 45: 810-823.
KILBurn, R. N. 1971a. On some species of the families Tonnidae, Hipponicidae, Buccinidae,
Columbariidae, Fasciolariidae, Psammobiidae and Mactridae (Mollusca) in South African
waters. Ann. Natal Mus. 20: 483-497.
KiILBuRN, R. N. 19716. A revision of the littoral Conidae (Mollusca: Gastropoda) of the
Cape Province.—Ann. Natal Mus. 21: 37-54.
KILBurn, R. N. & TANKARD, A. J. 1975. Pleistocene molluscs from the west and south coasts
of the Cape Province, South Africa. Ann. S. Afr. Mus. 67: 183-226.
KorrinGA, P. 1956. Oyster culture in South Africa.—Investl. Rep. Div. Sea Fish., S. Afr.
20: 1-86.
THERMALLY ANOMALOUS LATE PLEISTOCENE MOLLUSCS 45
KRISTENSEN, J. 1959. The coastal waters of the Netherlands as an environment of Mollusca
life. — Basteria 23 (Suppl.): 18-55.
Lams, H. H. 1961. Fundamentals of climate. Jn: Nairn, A.E.M. ed. Descriptive palaeo-
climatology: 8-44. New York: Interscience Publishers.
Mercer, J. H. 1968. Antarctic ice and Sangamon sea level.—Pub/. int. Ass. scient. Hydrol
79: 217-225.
NIcKLEs, M. 1950. Mollusques testacés marins de la céte occidentale d’ Afrique. Paris: Lechevalier
OLson, E. C. 1957. Size-frequency distributions in samples of extinct organisms.—J. Geol. 65:
309-333.
OrrEN, M. J. 1963. Hydrological observations in the South West Indian Ocean. —Jnvestl. Rep.
Div. Sea Fish., S. Afr. 45: 1-61.
Orren, M. J. 1966. Hydrology of the South West Indian Ocean. —Jnvestl. Rep. Div. Sea Fish.,
S. Afr. 55: 1-35. 7
RicHarps, H. G. 1962. Studies on the marine Pleistocene:
Part 1. The marine Pleistocene of the Americas and Europe.
Part 2. The marine Pleistocene mollusks of eastern North America.—
Trans. Am. phil. Soc. 52 (3): 1-141.
SCHELL, I. I. 1968. On the relation between the winds off Southwest Africa and the Benguela
Current and Agulhas Current penetration in the South Atlantic.— Dr. hydrogr. Z. 21:
109-117.
SHACKLETON, N. J. 1969. The last interglacial in the marine and terrestrial records.—Proc. r.
Soc. Lond. B. 174: 135-154.
SHACKLETON, N. J. & OppykeE, N. D. 1973. Oxygen isotope and palaeomagnetic stratigraphy
of equatorial Pacific core V 28-238: Oxygen isotope temperatures and ice volumes on a
10° year and 10° year scale.— Quatern. Res. 3: 39-55.
SHANNON, L. V. 1966. Hydrology of the south and west coasts of South Africa.—Jnvestl.
Rep. Div. Sea Fish., S. Afr. 58: 1-22.
STANLEY, S. M. 1970. Relation of shell form to life habits of the Bivalvia (Mollusca).—Mem
geol. Soc. Am. 125: 1-296.
SVERDRUP, H. U., JOHNSON, M. W. & FLEMING, R. H. 1942. The oceans, their physics, chemistry,
and general biology. New York: Prentice-Hall.
TANKARD, A. J. & SCHWEITZER, F. R. 1974. The geology of Die Kelders cave and environs:
a palaeoenvironmental study.—S. Afr. J. Sci. 70: 365-369.
VALENTINE, J. W. 1955. Upwellirig and thermally anomalous Pacific coast Pleistocene molluscan
faunas.—Am. J. Sci. 253: 462-474.
VALENTINE, J. W. 1957. Late Pleistocene faunas from the northwestern coast of Baja California,
Mexico.— Mem. S. Diego Soc. nat. Hist. 12: 289-308.
VALENTINE, J. W. 1961. Paleoecologic molluscan geography of the Californian Pleistocene. —
Univ. Calif. Publs geol. Sci. 34: 309-442.
VALENTINE, J. W. 1971. Resource supply and species diversity patterns.—Lethaia 4: 51-61.
VALENTINE, J. W. & MEADE, R. F. 1961. California Pleistocene paleotemperature.— Univ.
Calif. Publs Bull. Dep. Geol. 40: 1-46.
VAN ZINDEREN BAKKER, E. M. 1967. Upper Pleistocene and Holocene stratigraphy and ecology
on the basis of vegetation changes in sub-Saharan Africa. Jn: Bishop, W. W. and Clark,
J. D. eds. Background to evolution in Africa: 125-147. Chicago: University of Chicago
Press.
VAN ZINDEREN BAKKER, E. M. (in press). The origin and palaeoenvironment of the Namib
Desert biome. —J. Biogeogr.
WAGNER, P. E. & MERENSKY, H. 1928. The diamond deposits on the coast of Little Namaqua-
land.—Trans. geol. Soc. S. Afr. 31: 1-35.
WILBur, K. M. & YONGE, C. M. eds. 1964. Physiology of Mollusca. Vol. 1. New York: Academic
Press.
ZINSMEISTER, W. J. 1974. A new interpretation of thermally anomalous molluscan assemblages
of the California Pleistocene. —J. Paleont. 48: 84-94.
6. SYSTEMATIC papers must conform with the International code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. n., sp. n., comb. n.,
syn. n., etc.
An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.
Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (figs 8a—b).
Nucula largillierti Philippi, 1861: 87
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers
Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.
In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes ; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:
Holotype
SAM-A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach,
Port Elizabeth (33.51S, 25.39E), collected by A. Smith, 15 January 1973.
Note standard form of writing South African Museum registration numbers and of date.
7. SPECIAL HOUSE RULES
Capital initial letters
(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. ‘... the Figure depicting C. namacolus .. .’
*...in C. namacolus (Fig. 10)...’
(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
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ANTHONY J. TANKARD
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LATE PLEISTOCENE MOLLUSCS
FROM THE SOUTH-WESTERN CAPE PROVINCE,
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VOLUME 69 PART 3 DECEMBER 1975
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BuLLouaGu, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan,
FIscHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques.—J. Conch., Paris 88: 100-140.
FISCHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines.—Archs
Zool. exp. gén. 74: 627-634.
Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.—
Ann, Mag. nat. Hist. (13) 2: 309-320.
Koun, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian
Ocean. —Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
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ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 69 Band
December 1975 Desember
Part 3 Deel
RECORDS OF MUD-PRAWNS (GENUS
CALLIANASSA) FROM SOUTH AFRICA AND
MAURITIUS (CRUSTACEA, DECAPODA,
THALASSINIDEA)
By
BRIAN KENSLEY
Cape Town Kaapstad
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RECORDS OF MUD-PRAWNS (GENUS CALLIANASSA)
FROM SOUTH AFRICA AND MAURITIUS
(CRUSTACEA, DECAPODA, THALASSINIDEA)
By
BRIAN KENSLEY
South African Museum, Cape Town
(With 5 figures)
[MS. accepted 17 June 1975]
ABSTRACT
A new species of Callianassa is described from Port Alfred, South Africa. Three species
from Mauritius, viz. C. mauritiana, C. armata, and C. indica are figured and earlier descriptions
are supplemented. The latter two species are new records for Mauritius. C. kraussi is recorded
as a Pleistocene fossil from Table Bay and Port Elizabeth.
CONTENTS
PAGE
Introduction . : : : : Z : 4 Al
Systematic account : : : : : - 48
Acknowledgements Oe ee . See 7
References. : F , F ; : prey
INTRODUCTION
When two specimens of a large Callianassa from the Kowie River at Port
Alfred were submitted to the South African Museum for identification, it was
found that they belonged to none of the six species previously recorded from
South Africa (Kensley 1974). Use of Barnard’s key (1950: 505) ran the species
down to C. martensi Miers. In an attempt to solve this problem of identification,
two species of mud-prawn were obtained on loan from the Mauritius Institute,
a few specimens were collected in Mauritius by D. Hatton and given to the
South African Museum, and the type specimens of the two species of Callianassa
previously recorded from Mauritius were obtained on loan from the British
Museum.
As a result, the specimens from the Kowie River were found to be
undescribed. The specimens from Mauritius were found to be new records for
the island, and it was thought useful to figure them and to supply brief descrip-
tions. Also, it was thought desirable to supplement the description of Callianassa
mauritiana. The other species from Mauritius, viz. C. martensi Miers, has
recently been very thoroughly redescribed (Tirmizi 1974).
In the accompanying figures, all dimensions are in millimetres.
47
Ann. S. Afr. Mus. 69 (3), 1976: 47-58, 5 figs.
48 ANNALS OF THE SOUTH AFRICAN MUSEUM
SYSTEMATIC ACCOUNT
Callianassa armata A. M. Edwards
Fig. 1A-H
Callianassa armata A. M. Edwards, 1870: 90, pl. 1. De Man, 1902: 754; 1928a: 109.
Description
3 Carapace with spinose acutely triangular rostrum reaching to distal
end of cornea; well-defined antero-lateral spine present.
Eyestalks reaching to end of basal peduncular segment of antennule,
distally slightly curved away from midline; cornea situated dorso-laterally at
about midpoint of outer eyestalk margin; outer proximal part of eyestalk
bearing three or four tiny spinules.
Antennular peduncle three-segmented, second segment about half length
of third. Two distal antennal peduncular segments slender, subequal, basal
segment bearing distal spine.
Third maxilliped with propodus, carpus, merus, and ischium moderately
expanded; propodus with notch on antero-distal margin; ischium bearing on
inner surface a marked distal crest armed with about twelve teeth of varying
sizes, proximal portion bearing three separated spines.
Larger cheliped with dactylus strongly curved, cutting edge uneven, with
no well-marked teeth or tubercles; propodus with upper margin bearing three
strong spines; cutting edge of thumb with single blunt proximal tubercle,
ventral margin proximally slightly dentate; carpus shorter than propodal palm,
ventral margin bearing nine spines; merus with two spines on dorsal margin,
seven on ventral margin; ischium armed with row of ten spines, increasing in
size distally.
Finger and thumb of smaller chela gaping, cutting edge of finger unarmed,
that of thumb with blunt tubercle at about midpoint; upper margin of palm
bearing three spines; carpus armed with four spines on ventral margin; merus
bearing three spines on dorsal margin, four on ventral margin; ischium with
row of fourteen spines, distal four longer than more proximal spines.
Propodus of third pereiopod with posterior lobe not very marked, evenly
rounded. Telson broader than long, with semicircular seta-bearing ridge in
proximal third, and two very faint radiating rounded ridges not quite reaching
distal margin.
Uropodal exopod markedly bipartite, anterior portion about half length
of posterior, two portions separated by a curved ridge bearing a sharp spine
proximally; endopod lanceolate with rounded tubercle at base.
Material
1 3, Mauritius Institute 956. Carapace length (including rostrum) 28,4 mm.
Total length 98 mm.
RECORDS OF MUD-PRAWNS FROM SOUTH AFRICA AND MAURITIUS 49
Fig. 1. Callianassa armata 3
A. Anterior carapace, eyestalks, and antennae in dorsal view. B. Smaller cheliped. C. Larger
cheliped. D. Third maxilliped, with inner view of ischium. E. Third pereiopod. F. Telson
and uropod. G. First pleopod. H. Second pleopod.
50 ANNALS OF THE SOUTH AFRICAN MUSEUM
Previous Records
Fiji Islands; Ternate, India.
Remarks
Callianassa armata was described from a female of 125 mm from the Fiji
Islands, while the second specimen was a juvenile female from Ternate, India.
The present specimen, being a male, may account for a few discrepancies with
the original description. For example, De Man (1928qa) in his key to the species
of the subgenus Callichirus described the uropodal exopod as lacking a spine—
this spine is obviously present in the male. In the latter, the sixth pleonal seg-
ment is two and a half times the length of the telson, while in the Fijian female
this segment is three times the length of the telson.
Callianassa indica de Man
Fig. 2A-E
Callianassa indica de Man, 1905: 605; 1928a:100, 160, pl. 17 (fig. 26).
Description
2 Rostrum very short, obtusely triangular, antero-lateral angles rounded,
bearing tufts of setae.
Cornea dorso-lateral; eyestalks with rounded apex armed with three or
four very small blunt tubercles.
Propodus of third maxilliped as long as broad, inner face of ischium
bearing curved row of spines, those of distal part of row closely packed, unequal,
longer spines alternating with three or four short spines, more proximal spines
separate and subequal.
Larger cheliped with dactylus strongly curved, cutting edge bearing two
strong tubercles at about midpoint, thumb of propodus proximally finely
denticulate; palm of propodus one and a half times longer than dactylus,
ventral margin bearing about nine small serrations; ventral margin of carpus
evenly convex; ventral margin of merus armed with ten small teeth; ventral
margin of ischium bearing about twelve small teeth, increasing in size distally.
Smaller cheliped with finger and thumb gaping, equal in length to palm
of propodus; carpus and merus subequal in length, unarmed.
Material
1 2. Mauritius Institute 948, Black River, Mauritius. Carapace length
(including rostrum) 22,5 mm. Total length 84 mm.
Previous Records
Kangeang Reef, Bay of Kankamaraan, East Indies.
RECORDS OF MUD-PRAWNS FROM SOUTH AFRICA AND MAURITIUS 51
Fig. 2. Callianassa indica 9
A. Anterior carapace, eyestalks, and antennae in dorsal view. B. Larger cheliped. C. Smaller
cheliped. D. Third pereiopod. E. Third maxilliped, with inner view of ischium.
Remarks
This is the second record of a species that was described from a single
male, total length 90 mm, which lacked the larger cheliped. The above figures
and description, although of a mature female, supplement De Man’s description.
Callianassa mauritiana Miers
Fig. 3A-H
Callianassa mauritiana Miers, 1882: 341; 1884: 15, pl. 1 (fig. 2). Nobili, 1906: 106, figs 5, 6.
De Man, 1928a: 99; 19285: 10, pl. 2 (fig. 4). Michel, 1974: 256.
Description
In spite of the descriptions given by Miers, Nobili, and De Man, it was felt
that additional figures of this rarely recorded species would be useful. The
figures were done from the male and female syntypes from the British Museum
(Natural History) collection.
52 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 3. Callianassa mauritiana
A. Anterior carapace, eystalks, and antennae in dorsal view. B. Telson and uropod.
C. Third maxilliped, with inner view of ischium. D. Third pereiopod. E. First pleopod 3.
F. Second pleopod 3. G. First pleopod 2. H. Second pleopod 9.
RECORDS OF MUD-PRAWNS FROM SOUTH AFRICA AND MAURITIUS 53
Eyestalk with cornea situated dorsally, slightly distal to midpoint, with
tiny but distinct rounded tubercle distal to cornea.
Third pereiopod with propodus having a curved posterior lobe.
Uropodal exopod bipartite, anterior portion only slightly shorter than
posterior, with spine at base. This spine only developed in adult.
Material
Mauritius, SAM-A13636 3 carapace length 12,0 mm total length —
2 11,3 mm 43 mm
2 9,0 mm 31 mm
Previous Records
Mauritius; Red Sea.
Remarks
Miers (1882: 341) expressed the opinion that the detached large chela
in the container with the two syntypes probably did not belong to either of
the specimens, but to a distinct and larger individual. This need not necessarily
be so, as the species seems to be characterized by a disproportionally massive
larger cheliped. In an immature male with a carapace length of 12,0 mm the
larger cheliped has a length of 40 mm.
Callianassa pixii sp. 0.
Fig. 4A-H, 5A-K
Description
2 Carapace with oval shield well defined; rostrum short, acute, triangular,
with median keel running on to carapace; front tridentate, lateral teeth only
slightly shorter than rostrum, also slightly keeled. Eyestalks contiguous for
whole length, tapering from about mid-length to acute apex; cornea situated
dorsally in distal half, hardly visible. Eyestalks reaching to end of first anten-
nular peduncle segment.
Third peduncular segment of antennule slightly more than twice length of
second. Antennal peduncle not quite reaching distal end of antennular peduncle,
two distal segments subequal.
Mandibular palp three-segmented, distal segment set with numerous
setae, equal in length to two proximal segments together; incisor portion weakly
chitinized, with about thirteen small teeth on cutting edge; molar portion a
blunt flattened structure set with plumose setae.
First maxilla, palp slender, distal portion flexed, median edges of lobes
densely fringed with setae.
Second maxilla also densely fringed with setae.
Epipod of first maxilliped with posterior portion longer and broader than
anterior part.
54 ANNALS OF THE SOUTH AFRICAN MUSEUM
Second maxilliped pediform, exopod slender, leaf-like, endopod with
apical segment bearing cluster of short, stiff setae, rest of appendage fringed
with long setae.
Third maxilliped pediform, lacking exopod, three distal segments together
slightly shorter than two proximal segments; merus and ischium fringed with
long setae, together four and a half times longer than wide; inner face bearing
another fringe of setae; propodus and carpus each with dense bristle pad on
inner face.
First pereiopods subequal, finger and thumb slightly shorter than palm
of propodus, dactylus curved only at tip, cutting edge entire, thumb of propodus
Fig. 4. Callianassa pixii sp. n.
A. Anterior carapace, eyestalks, and antennae in dorsal view. B. Telson and uropod.
C. Mandible. D. First maxilla. E. Second maxilla. F. First maxilliped. G. Second maxilliped.
H. Third maxilliped, inner view.
ee Tae
RECORDS OF MUD-PRAWNS FROM SOUTH AFRICA AND MAURITIUS 355
Fig. 5. Callianassa pixii sp. n.
A. Larger cheliped 3. B. Smaller cheliped g. C. Third pereiopod. D. Second pereiopod.
E. Fourth pereiopod. F. Fifth pereiopod. G. First pleopod ¢. H. Second pleopod ¢.
I. First pleopod 2. J. Second pleopod 9. K. Third pleopod 3.
56 ANNALS OF THE SOUTH AFRICAN MUSEUM
with large triangular tooth at midpoint of cutting edge, followed proximally
by fine denticulations; carpus about half length of palm of propodus, upper
margin entire; merus longer than carpus; ischium with about six tiny denticu-
lations on lower margin.
Second pereiopod chelate, with dactylus, propodus, and carpus together
subequal to merus in length, cutting margins of dactylus and propodus straight,
all segments bearing numerous elongate setae.
Third pereiopod with posterior lobe of propodus oblique-truncate.
Fourth pereiopod with propodus having distal thumb-like lobe slightly
shorter than dactylus, and therefore incompletely chelate; dactylus and propodus
set with numerous short stiff setae.
Fifth pereiopod chelate, dactylus and propodal thumb apically spooned,
set with patches of short dense setae.
First pleopod uniramous, longer distal portion flexed at right angle to
basal portion, bearing clumps of elongate setae.
Second pleopod biramous, rami subequal in length, inner ramus bearing
small lobe on distal half.
Telson broader than long, distal margin concave.
Uropodal exopod bipartite, anterior portion only slightly shorter than
posterior, with spine at base, both portions fringed with short setae; endopod
oval, shorter than exopod, with rounded longitudinal ridge.
3 Larger cheliped subequal in length to smaller cheliped, but propodus
and carpus broader; finger and thumb shorter than palm of propodus; dactylus
only distally hooked, with blunt tubercle near base of cutting edge; thumb of
propodus with slight notch and fine denticulations proximal to blunt tubercle
at about midpoint of cutting edge; upper margin of palm finely denticulate;
carpus with single tooth on upper margin; lower margin of ischium with two
or three barely discernible teeth.
Smaller cheliped similar to first pereiopods of female, but possessing single
tooth on upper margin of carpus.
First pleopod about half length of second pleopod, uniramous, two-
segmented, distal segment with three slight tuberculations.
Second pleopod biramous, outer ramus half length of inner, latter with
trilobed apex, middle lobe largest.
Material
Holotype 9 SAM-A13637 carapace length 27,0 mm total length 85 mm.
Allotype ¢ SAM-A13637 carapace length 26,0 mm (posterior pleon and
telson missing). Kowie River estuary, Cape Province, South Africa.
Remarks
Callianassa pixii belongs to that group of species of the subgenus
Callichirus which is characterized by the possession of a pediform third
maxilliped. Of this group, C. pixii most closely resembles C. guineensis de
RECORDS OF MUD-PRAWNS FROM SOUTH AFRICA AND MAURITIUS 57
Man, described from the Gold Coast (De Man 19286). Several differences
make the separation of these two species quite simple.
Although the third maxillipeds are very similar, in Callianassa pixii the
merus and ischium have a more definite curvature than in the west African
species. In the latter, the antennae are distinctly shorter and more slender
than the antennules, eyestalks are not contiguous, and the cornea of the eyes
are large and distinct. In C. pixii the antennae are only slightly shorter than
the antennules and of similar thickness. The eyestalks are contiguous and the
cornea small and hardly defined. The telson and uropodal endopods of the
two species are similar, but the uropodal exopod in C. guineensis is unevenly
bipartite, while in the Kowie species the two portions of the exopod are of
similar length and breadth. De Man’s specimen of C. guineensis was a juvenile,
thus a comparison of the cheliped structure is of little value.
The species is named for Mr Pixie John, well-known local figure of Port
Alfred.
Callianassa kraussi Stebbing
Callianassa kraussi Barnard, 1950: 506. Kensley, 1974: 277.
Remarks
Numerous dactyli and propodi of the larger chelipeds of this species have
been collected from Pleistocene raised beaches at Swartkops, near Port Eliza-
beth, and at Milnerton, Table Bay. Several are larger and more robust than those
of the largest known living specimens of the species.
ACKNOWLEDGEMENTS
My sincere thanks are due to the following for making material available
for study: Dr R. W. Ingle, of the British Museum (Natural History) for the
loan of type material of Callianassa martensi and C. mauritiana; Dr C. Miche
of the Mauritius Institute, for the loan of specimens of C. armata and C. indica;
Dr A. T. Forbes for donating the specimens of C. pixii to the South African
Museum, and for useful information; and Mr D. Hatton, for the donation of
specimens of C. mauritiana to the South African Museum.
I am grateful to Professor Nasima Tirmizi of the University of Karachi,
and Dr R. W. Ingle, for reading the manuscript of this paper and for many
useful comments and criticisms.
REFERENCES
BARNARD, K. H. 1950. Descriptive catalogue of the South African Decapod Crustacea (crabs
and shrimps).— Ann. S. Afr. Mus. 38: 1-837.
Epwarbs, A. M. 1870, Révision du genre Callianassa (Leach) et description de plusieurs
espéces nouvelles de ce groupe.— Nouv. Archs Mus. Hist. nat., Paris 6: 75-102.
KeEnsLEY, B. F. 1974. The genus Callianassa (Crustacea, Decapoda, Thalassinidea) from the
west coast of South Africa with a key to South African species.— Ann. S. Afr. Mus. 62:
265-278.
58 ANNALS OF THE SOUTH AFRICAN MUSEUM
MAN, J. G. pe. 1902. Die von Herrn Professor Kiikenthal im Indischen Archipel gesammelten
Dekapoden und Stomatopoden.—Abh. senckenb. naturforsch. Ges. 25: 267-929.
Man, J. G. DE. 1905. Diagnoses of new species of macrurous decapod Crustacea from the
Siboga-Expedition. — Tijdschr. ned. dierk. Vereen. (2) 9(3/4): 587-614.
Man, J. G. Dg. 1928a. The Decapoda of the Siboga-Expedition. Part VII. The Thalassinidae
and Callianassidae collected by Siboga-Expedition with some remarks on the
Laomediidae.— Siboga Exped. monogr. 39a.6: 1-187.
MAN, J. G. pe. 19285. A contribution to the knowledge of twenty-two species and three
varieties of the genus Callianassa Leach.— Capita zool. 2(6): 1-56.
MICHEL, C. 1974. Notes on Marine Biology studies made in Mauritius.— Bull. Mauritius Inst. -
7(2): 1-284.
Miers, E. J. 1882. On some Crustaceans collected at the Mauritius.—Proc. zool. Soc. Lond.
1882: 339-342.
Miers, E. J. 1884. On some crustaceans from Mauritius. —Proc. zool. Soc. Lond. 1884: 10-17.
Nosiui, G. 1906. Faune carcinologique de la Mer Rouge. Décapodes et Stomatopodes. —
Annls Sci. nat. 4: 1-347.
Tirmizi, N. M. 1974. A description of Callianassa martensi Miers, 1884 (Decapoda, Thalas-
sinidea) and its occurrence in the northern Arabian Sea.— Crustaceana 26: 286-292.
6. SYSTEMATIC papers must conform with the Jnternational code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. n., sp. n., comb. n.,
syD. n., etc.
An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.
Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (figs 8a—b).
Nucula largillierti Philippi, 1861: 87
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers
Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.
In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
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BuLLouGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
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Zool. exp. gén. 74: 627-634.
Kogn, A. J. 19602. oy. Gastropoda) in the Trincomalee region of Ceylon.—
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Kon, A. J. 19605. poe eg he= egg masses and larval development in Conus from the Indian
Senn cole tie gham ceeanonr. Col. 17 ra, Gastropoda marina, Bivalvie. In: Scumzza, 1. Zooloniache
ollusca: fo) marina, In:
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ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 69 Band
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Part 4 Deel
NOTES ON VARIATION IN PENGUINS AND ON
PossllL PENGUINS FROM THE PLIOCENE OF
LANGEBAANWEG, CAPE PROVINCE,
SOUTH AFRICA
By
GEORGE GAYLORD SIMPSON
Cape Town Kaapstad
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NOTES ON VARIATION IN PENGUINS AND ON FOSSIL PENGUINS
FROM THE PLIOCENE OF LANGEBAANWEG, CAPE PROVINCE,
SOUTH AFRICA
By
GEORGE GAYLORD SIMPSON
The Simroe Foundation, 5151 East Holmes Street, Tucson, Arizona 85711
(With 5 figures and 4 tables)
[MS. accepted 25 June 1975]
ABSTRACT
Improved statistical data on limb bones of Recent Spheniscus demersus agree with previous
suggestions that adaptively functional elements in penguins have comparatively slight variation.
These data assist in the interpretation of isolated fossil bones. Additional specimens from the
Pliocene of Langebaanweg indicate that Spheniscus predemersus Simpson, 1971, was incorrectly
referred to Spheniscus, and the species is placed in a new genus, Jnguza. A second, larger species
occurs at Langebaanweg, but available material does not warrant further identification or
diagnosis.
CONTENTS
PAGE
HAILOGUCHION © ey te) ee an ee ee eevee s. =, Oo
Comparative data on variation in penguin limb bones . . 60
Rossilsiirom,Langebaanwer. (2/02 9:1 = = «8. =) = 163
AcknowledeementSy |, 0) mmr fo eet ==) fo eee Le
IRCLELCHICCS Ich et Sarre NER Peseta: me te See oe
INTRODUCTION
These notes consist of two parts. First, a statistical study is made of some
dimensions of limb bones of Spheniscus demersus, not only to increase the data
for variation in penguins generally, but also and particularly to broaden the basis
for sorting and identifying isolated and often incomplete fossil penguin bones.
Next, the considerably enlarged suite of fossil penguin bones from Langebaanweg
is studied. It is reasonably sure that many of these bones, and especially a partial
but characteristic tarsometatarsus, belong to the species previously described as
Spheniscus predemersus (Simpson 1971la). The tarsometatarsus excludes the
species from Spheniscus and indicates reference to a hitherto unnamed and
undefined genus. That genus is technically established later in this paper after
the basis for its distinction and for the reference of predemersus to it has been
established, but to simplify things the new combination Jnguza predemersus
will be used throughout.
All fossil specimens here discussed are in the South African Museum.
Reference numbers for all begin SAM-PQ-L, followed by a serial number, but
for brevity they are here given only as L plus the number, which suffices for
identification.
59
Ann. S. Afr. Mus. 69 (4), 1976: 59-72, 5 figs, 4 tables.
60 ANNALS OF THE SOUTH AFRICAN MUSEUM
All measurements are in millimetres. Other conventions and abbreviations
are explained as needed.
COMPARATIVE DATA ON VARIATION IN PENGUIN LIMB BONES
Many museums now have some osteological material of penguins, but few
have samples of unified origin and large enough to give good estimates of |
population variation. The most nearly adequate statistical data are those in
Simpson (1946) for specimens, mostly of Aptenodytes patagonicus, in the
American Museum of Natural History, relatively few specimens, not sexed,
and without precise locality data. Now better data can be given for another
species, Spheniscus demersus, the African black-footed or jackass penguin.
The population represented is that of Robben and Dyer Islands, one north-
west and the other south-east of Cape Town. Although the islands are some
distance apart, their penguin populations are essentially unified as birds banded
on one may turn up on the other. The birds in this sample were caught in an oil-
slick and taken for treatment by the South African National Foundation for the
Conservation of Coastal Birds (SANCCOB) but failed to survive. (Many oiled
birds do recover after treatment.) The dead birds were turned over to the South
African Museum, where their wing and leg bones were macerated. The left limb
bones are in that Museum, and the right limb bones of the same birds were kindly
sent by Dr Q. B. Hendey to the Simroe Foundation for study by the author.
It is conceivable that the sample is biased by the fact that these birds
died from oiling, but that is improbable. There is no apparent reason why
mortality of adult birds from oiling would be correlated with small differences
in the sizes of their limb bones.
Ten of the thirty birds involved are recorded as juvenile, and these were
not included in the statistics even though some of them have apparently fully
ossified limb bones. Three not recorded as juvenile do not have fully adult
ossification and were therefore also excluded. One specimen consisted of wings
only and was excluded because it is desirable to have wing and leg measurements
all on the same individual. One more was disregarded because of poor preser-
vation and one because of pathology (exostoses on some of the bones). Nineteen
specimens were thus measured. Of these, five were recorded as males, one
doubtfully. The doubtful specimen does happen to be smaller than the four
certain males, but it was considered incorrect to exclude it from the sample on
that account. The fourteen females measured are all recorded without question
as to sex.
The dimensions measured were selected primarily for their potential useful-
ness in judging variation and proportions in single bones, especially in fossil
specimens at hand. They are as follows:
Humerus:
a. Maximum longitudinal dimension.
b. Width of shaft about one-third of distance distal to head. In this
species this is taken at the minimum width of the shaft.
NOTES ON VARIATION IN PENGUINS 61
c. Width of shaft about two-thirds of distance distal to head.
d. From the radial condyle to the longest distal process.
Femur:
a. From the hollow between the head and trochanter to that between
distal condyles.
6. Proximal width.
c. Distal width.
Tibiotarsus:
a. From the proximal articulation (excluding the crest) to the hollow
between distal condyles.
b. Distal width.
Tarsometatarsus:
a. Length on third metatarsal (proximal convexity to distal groove).
b. Width of distal end of third metatarsal.
c. Length on fourth metatarsal (to distal groove).
The following statistics are given in Tables 1, 2 and 4:
N —number of specimens.
OR — observed range in sample.
X —mean and standard error.
S —Standard deviation and standard error.
V —coefficient of variation and standard error.
The males of this living species are in general larger in mean sample esti-
mates than the females, ‘humerus d’ being the only dimension of which this is
not true. However the differences are slight and there is large overlap in all the
observed ranges, still more in the probable population ranges. These bones in
this species cannot be reliably sexed on the basis of size.
The humerus is distinctly more variable in males than in females of this
sample. That could be due to sampling error, but probably is not. Its functional
significance, if any, is not clear. The tarsometatarsus is also somewhat more
variable in males than in females, but here the difference is less and is quite
probably due to sampling error. No sexual difference in variation is indicated
for femur or tibiotarsus.
The coefficients of variation are in general quite small. The mean of the
24 coefficients in Table | is only 3,63. The variation in functionally adaptive
dimensions of birds tends to be low, suggesting that these characters are subject
to effective stabilizing or centripetal selection. It is interesting that this is true
of wing bones in penguins, aqueous fliers, as well as in aerial fliers.
An aid to sorting isolated bones is provided by the ratios of measurements
in associated bones of individuals of one species. Some data from the Spheniscus
demersus sample are given in Table 2. There is no evident sexual distinction in
this respect. These ratios are likely to be different in different species, but they
probably will be close to those for S. demersus in species of approximately the
same size. There is a tendency in Recent species, at least, for the humerus to be
62 ANNALS OF THE SOUTH AFRICAN MUSEUM
TABLE 1
Statistics on Some Limb Bones of Spheniscus demersus
N OR xX N) V
Males
Humerus
a 5 64,0-69,5 67,82 + 1,06 2,37 + 0,75 3,50 + 1,11
b 5 10,0-11,5 10,64 + 0,33 0,74 + 0,24 6,99 + 2,21
c 5 11,7—-13,4 12,44 + 0,36 0,80 + 0,25 6,40 + 2,02
d 5 19,9-22,5 21,56 +-0,50 1,12 + 0,35 5,20 + 1,65
Femur
a 5 65,6-70,0 68,32 + 0,73 1,64 + 0,52 2,40 + 0,76
b 5 14,5-15,5 15,04 + 0,20 0,44 + 0,14 2,96 + 0,94
c 5 13,9-15,0 14,42 + 0,22 0,50 + 0,16 3,45 + 1,09
Tibiotarsus
a 5) 96,7-104,4 101,16 + 1,28 2,87 + 0,91 2,84 + 0,90
b 5 13,1-14,3 13,86 + 0,21 0,46 + 0,15 3,33 + 1,05
Tarsometatarsus
a 5 28,9-32,8 31,26 + 0,66 1,47 + 0,47 4,72 + 1,49
b 5 6,3-7,1 6,76 + 0,15 0,34 + 0,11 5,08 + 1,61
é 5 26,0—28,5 27,22 + 0,44 0,98 + 0,31 3,61 + 1,14
Females
Humerus
a 14 63,9-69,3 66,66 + 0,51 1,91 + 0,36 2,87 + 0,54
b 14 10,0-10,8 10,43 + 0,07 0,28 + 0,05 2,64 + 0,50
Cc 14 11,3-12,7 11,94 + 0,09 0,35 + 0,07 PHN ae OLS)
d 14 20,9-22,9 21,66 + 0,14 0,51 + 0,10 2,34 + 0,44
Femur
a 14 61,9-68,9 65,94 + 0,43 1,61 + 0,30 2,44 + 0,46
b 14 14,1-15,5 14,79 + 0,12 0,44 + 0,08 3,01 + 0,57
c 14 13,6—15,3 14,27 + 0,12 0,45 + 0,09 3,19 + 0,60
Tibiotarsus
a 14 91,8-102,6 97,54 + 0,88 3,29 + 0,62 3,37 + 0,64
b 14 13,1-14,3 13,71 + 0,11 0,42 + 0,08 3,06 + 0,58
Tarsometatarsus
a 14 28,9-32,8 30,79 + 0,28 1,05 + 0,20 3,39 + 0,64
b 14 6,5—7,4 6,74 + 0,07 0,26 + 0,05 3,93 + 0,74
c 14 25,4-28,0 26,55 + 0,24 0,92 + 0,17 3,45 + 0,65 -
Symbols are explained in the text.
TABLE 2
Ratios of Some Dimensions in Spheniscus demersus
Humerus a Humerus a Femur a
Femur a Tarsometatarsus a Tarsometatarsus a
N OR x N OR 574 N OR x
Males. =.)) 2) 311795" 10:96—L.08- 7099 57 2312-2 21F SB 2a 5 2,10-2,27 2,19
Females 14 0,98-1,04 1,01
Symbols are explained in the text.
14 2,09-2,28 2,17 14 2,08-2,27 2,14
longer relative to the femur and tarsometatarsus the larger the species (Simpson
1946: table 8).
Since limb bones cannot be sexed by size in S. demersus and this is
apparently usually but not necessarily always true in penguins, statistics were
also calculated for some dimensions in a sample of S. demersus with the most
NOTES ON VARIATION IN PENGUINS 63
probable ratio of males to females, that is, equal numbers of the two sexes.
This includes the five males available and five females taken at random (using
a table of random numbers applied to the serial numbers of these specimens).
The dimensions were selected for comparison with available specimens of fossil
humeri and femora from Langebaanweg. Results are given in Table 4.
FOSSILS FROM LANGEBAANWEG
Penguins from the Pliocene of Langebaanweg were previously described by
the author (Simpson 1971a) on the basis of three humeri, one essentially com-
plete and two fragmentary, a partial tibiotarsus, two complete femora, and one
pedal phalanx. The humeri were referred to a then new species as Spheniscus
predemersus (Fig. 1). It will be shown below that the generic ascription was
almost certainly erroneous, and the name Jnguza predemersus will be used here.
Later discoveries have added greatly to the available materials, although
even now they are not wholly adequate. Most of the bones of the wings and legs
are represented, although some, unfortunately including the tarsometatarsus,
only by imperfect specimens. The provenience is described in Hendey (1974)
and Dr Hendey has added information in personal communication. The speci-
mens are from the ‘E’ Quarry at Langebaanweg, and for the most part from the
area designated as ‘East Stream’ (Hendey 1974: fig. 3). All are from the Vars-
water Formation, and with one exception they are from ‘Bed 2’ of Hendey (1974:
table 4), which he is now proposing to call the ‘Quartzose Sand Member’. That
member is comprised of deposits accumulated in and adjacent to an estuary
and its fauna is made up largely of terrestrial vertebrates. Besides the penguins,
aquatic or amphibious elements are represented by an otter, Enhydriodon
africanus, and a seal, Prionodelphis capensis. These are at present the only fossil
penguins found in a predominantly non-marine association. The one exception
referred to above is from the Gravel Member (=Bed 1), the basal unit of the
Varswater Formation in which marine fossils are predominant.
This deposit is the type of the proposed South African provincial land
mammal stage and age Langebaanian, Pliocene in age and tentatively correlated
with the Astian of Europe and the Rexroadian of North America (Hendey 1974:
table 8).
Renewed study of the systematics of these fossil penguins with enlarged
samples is based on the humeri, femora, tibiotarsi, and tarsometatarsi. The
other bones, although numerous, are less characteristic and do not add to con-
clusions based on these bones.
Measurements are given in Table 3. The dimensions are the same as those
specified in the text above and used in Tables | and 2. Statistics derived from
these measurements are given in Table 4 and there compared with statistics
for a sample of Recent Spheniscus demersus made to include equal numbers of
the two sexes, as noted in previous text. These dimensions, and therefore the
measurements, differ somewhat from those previously used for the smaller
sample (Simpson 197la: table 1). For ‘humerus 5’ and ‘humerus c’ the co-
64 ANNALS OF THE SOUTH AFRICAN MUSEUM
IVULT ALLL LLL LPL LLL LLL
Fig.1. Inguza predemersus, type, humerus SAM-PQ-L6510. Ventral, dorsal and postaxial
views. Scale in mm.
NOTES ON VARIATION IN PENGUINS
TABLE 3
65
Measurements of some Limb Bones of Fossil Penguins from Langebaanweg
Specimen No. Dimension
Humerus
a b c d
L6510, holotype of
Inguza predemersus 59,0 8,6 10,0 17,8
L123010 57,5 9,6 10,9 18,9
L14853 . — 8,7 11,1 18,4
L12887A — 9,4 ile 19,3
122952 . — 10,0 Ale? =
L21928 . — 9,8 — —
Femur
a b c
L23002 . 59,8 — 13,3
L22983 . 61,5 — 13,4
Wer R ca ne ls 62,3 13,9 13,9
|e PERN Be 0 et a oe — 14,4 —
L12524A — — 14,5
L13154 . TSA 18,1 16,3
£3656: 79,5 — 17,6
L13066A — 18,7 —
Tibiotarsus
a b
123012... 85,2 11,5
122950 . ca. 92 3
Tarsometatarsus
a b c
L23018 . YA 5,3 24,2
L22974 . — 4,7 —
L22985 . — ca. 74 —
Symbols are explained in the text.
TABLE 4
Some Statistical Data on a Sample with Equal Numbers of Males and Females of Spheniscus
demersus and on Available Fossils from Langebaanweg
Sample Dimension N OR x
Spheniscus Humerusb 10 10,0-11,5 10,59 + 0,16
demersus Humerus c 10 =‘11,7-13,4 12,24+ 0,16
Humerusd 10 19,9-22,5 21,63 + 0,25
Femur a 10 61,9-70,0 66,63 + 0,79
Femur c 10 13,6-15,0 14,19 + 0,14
Inguza
predemersus Humerus } 6 8,6-10,0 9,35 + 0,24
Humerus c S$ 10,0-11,7 10,98 + 0,28
Humerus d 4 17,8-19,3 18,60 + 0,32
Femur a 3 59,8-62,3 61,20 + 0,74
Femur c 4 13,3-14,5 13,78 + 0,28
Mixed sample, all measureable bones from Langebaanweg
Femur a 5 59,8-79,5 67,64 + 4,02
Femur c 6 13,3-17,6 14,83 + 0,71
Symbols are explained in the text.
S
0,52 + 0,12
0,62 + 0,12
0,80 + 0,18
2,50 + 0,56
0,44 + 0,10
0,58 + 0,17
0,62 + 0,29
0,65 + 0,23
1,28 + 0,52
0,55 + 0,19
9,00 + 2,01
1,74 + 0,50
Vv
4,87 + 1,09
4,21 + 0,94
3,72 + 0,83
3,76 + 0,84
3,12 + 0,70
6,19 + 1,79
5,67 + 1,27
3,48 + 1,23
2,09 + 0,85
7,26 + 2,57
13,31 + 2,98
11,75.2,3,39
66 ANNALS OF THE SOUTH AFRICAN MUSEUM
efficients of variation are somewhat higher than for S. demersus or, so far as
data are available, most other Recent penguins. However the difference is not
great enough to indicate that it is probably not due either to sampling error or
to somewhat greater real variability within a single species. For ‘humerus a’,
generally a less variable dimension, V is not statistically different from that of
S. demersus. Except for the moderate variation in these dimensions and in the
ratios among them, these humeri do not differ appreciably in morphology. |
They are therefore all tentatively referred to Inguza predemersus, the more
common but, as will now appear, not the only species of penguins in this fauna.
There are seven tibiotarsi in the collection but only one is complete, and
although three others are nearly so only one other gives approximate comparable
measurements. The variation in size is considerable, but again not enough to
indicate reliably that more than one species is represented. The two most
complete specimens, L23012 and L22950, are within a size range appropriate
for association with humeri referred to Jnguza predemersus.
The situation regarding the femora is quite different. For the six femora
affording one or more useful measurements the coefficients of variation for all
taken together are decidedly too high to be derived from a single species of
penguins. It is also evident in Table 3 that they fall into two quite distinct size
groups. The smaller group, L23002, L22983, L22117, L22954 and L12524<A, is
of appropriate size for association with the humeri referred to Jnguza pre-
demersus (Fig. 2). The ratio of the mean for ‘humerus a’ (two specimens) to the
mean for ‘femur a’ (three specimens) is 0,95. In Spheniscus demersus it is 0,99
(Table 2), not significantly different. The femora L13154 and L3656 give a
corresponding ratio of 0,75, which is significantly different, and these femora
surely belong to a second, larger species, to which L13066 also belongs (Fig. 3).
‘Femur a’ for specimens referred to Jnguza predemersus has unusually
slight variation and ‘femur c’ unusually large variation. However N is small
for both (3 and 4, respectively), standard errors are correspondingly large, and ©
the differences of values of V from those found in Spheniscus demersus are not
significant.
The specimen mentioned above as having come from the Gravel Member
is L21628, the distal part of a femur. The end is too abraded or corroded for
useful measurement, but this bone agrees closely with the larger specimens from
the Quartzose Sand Member. Dr Hendey (pers. com.) notes that ‘The Gravel
Member does include derived fossils, probably of Miocene Age’. It is, however,
probable that this fragment represents the same species as the larger one in the
overlying member.
There are 16 partial tarsometatarsi in the collection, but most of these
are scraps, especially distal condyles, from which little can be learned. The most
extensive, L23018 (Fig. 4), includes the third and fourth tarsometatarsals and
the dorsal, but not the plantar, part of the proximal end. The ratio for mean
‘humerus a’ of two Inguza predemersus to ‘tarsometatarsus a’ of this specimen
is 2,14, which suggests that these animals were of quite closely the same size
NOTES ON VARIATION IN PENGUINS 67
113 114 115 116 117
2
l
I
Fig. 2. Inguza predemersus, femur SAM—PQ-L22117. Anterior and posterior views.
and very likely of the same species. L23401, a fourth tarsometatarsal with some
adjacent bone, and L22974, most of a third tarsometatarsal with some adjacent
bone, also may be referred to this species, to which most of the lesser fragments
probably belong. There are, however, some scraps, notably L22985, the distal
part of the third tarsometatarsal and some adjacent bone, and L23402, approxi-
mately the same but even more poorly preserved, which belong to a definitely
distinct and larger species (Fig. 5). The species could well be the same as that
represented by the larger femora.
On L23018 it can be determined that the medial intermetatarsal foramen
(or inner proximal foramen of Zusi 1975) is larger than the lateral foramen and
slightly more distal. It opens on the plantar side distal to the medial (inner)
68 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 3. Spheniscidae indet. (left) and Inguza predemersus (right), femora SAM-—PQ-L13154
and SAM-PQ-L22117. Anterior views.
8
|
j
mm
|
6
1
5
1
4
l
3
|
112
1
|
NOTES ON VARIATION IN PENGUINS 69
115 116 117 118
Fig. 4. Inguza predemersus, tarsometatarsus SAM-—PQ-L23018. Dorsal and plantar views.
calcaneal ridge. These characters definitely exclude reference to the genus
Spheniscus in which the medial foramen is relatively smaller, is somewhat more
proximal, and opens on the medial side of the medial calcaneal ridge. (See
Zusi 1975, who shows that the latter arrangement occurs in Eudyptes; I have
confirmed that it also occurs and is apparently invariable in Spheniscus
demersus.) Among Recent penguins the genera Aptenodytes and Pygoscelis have
the arrangement of intermetatarsal foramina more as in this fossil. The Recent
species of those genera are decidedly larger than the fossil, even in Pygoscelis.
The tarsometatarsus is also less elongate, the foramina relatively less proximal,
70 ANNALS OF THE SOUTH AFRICAN MUSEUM
8
1
7
1
6
1
:
Fig. 5. Spheniscidae indet. (left) and Inguza predemersus (right), tarsometatarsi
SAM-PQ-L22985 and SAM-—PQ-L23018. Dorsal views.
more nearly in a transverse line, and more nearly equal in size. Reference of
the fossil to Pygoscelis or Aptenodytes is not tenable.
It seems open to little question that the holotype humerus of Jnguza prede-
mersus, Originally referred to Spheniscus, and the tarsometatarsus, now available
and added to the hypodigm, belong to the same species and that the species
neither belongs in nor was ancestral to Spheniscus. The humerus really differs
little from that of Spheniscus demersus. Apart from smaller size, it has the
shaft slightly more sigmoid and it lacks a preaxial angle or tubercle; it now
follows that these slight differences are associated with a tarsometatarsus surely
generically distinct from Spheniscus. They therefore now appear to be generic
characters although that would hardly be tenable on the basis of the humeri
alone. The author has been unable to find a previously named extinct genus to
which this species can be reasonably referred. Duntroonornis from the early
Oligocene of New Zealand may come as close as any, but its less elongate
tarsometatarsus, smaller and more unequal foramina, and perhaps the obliquity
of the metatarsals, if that is not caused by crushing of the only known specimen,
distinguish it (see Simpson 19714). ‘Spheniscus’ predemersus must therefore be
referred to a new, extinct genus which will now be formally proposed. It is
improbable that this genus is ancestral or close to any living penguins, but it
NOTES ON VARIATION IN PENGUINS 71
may conjecturally have some special relationship with Pygoscelis or perhaps
even Aptenodytes.
It is probably impossible and certainly inadvisable to identify or name
the second, larger species from Langebaanweg on the basis of specimens now
in hand. It is certainly distinct from /nguza predemersus, but the available
specimens are otherwise not diagnostic.
Technical validation of the new generic name and revision of the specific
name follow.
Order SPHENISCIFORMES
Family Spheniscidae
Inguza, gen. n.
Etymology: ‘Inguza’ is given by McLachlan & Liversidge (1970) as a ‘native
[South African] name’ for penguins. Greek derivatives appropriate for penguins
have become rather overdone and repetitious. Native African languages do not
have gender in the Latin sense, so this name is arbitrarily designated as
masculine.
Type-species: Spheniscus predemersus Simpson, 1971.
Included species: Type only.
Known distribution: Langebaanian, Pliocene, in the Quartzose Sand Member
of the Varswater Formation at Langebaanweg, Cape Province, Republic of
South Africa.
Diagnosis: Humerus with shaft narrower proximally, somewhat sigmoid;
tricipital fossa strongly bipartite; preaxial angulation absent. Tarsometatarsus
elongate; intermetatarsal foramina proximal; medial foramen larger than lateral
foramen, slightly more distal, opening distal to the medial calcaneal ridge on
the plantar surface; third and fourth metatarsals straight.
Inguza predemersus (Simpson, 1971)
Spheniscus predemersus Simpson, 1971a: 1144.
Etymology: Pre+- demersus, as older than known Spheniscus demersus and
erroneously believed to be specially related to the latter.
Holotype: SAM-PQ-L6510, left humerus, essentially complete.
Present Hypodigm: The type and the following:
L23010, complete humerus, L14853, L12887A, L22952, and L21928,
partial humeri.
L22117A, complete femur, L23002, L22983, L122954, and L12524A,
partial femora.
L23012, complete, and L22930, nearly complete, tibiotarsi.
L23018 and L22974, partial tarsometatarsi.
72 ANNALS OF THE SOUTH AFRICAN MUSEUM
Numerous other bones, mostly fragmentary, almost certainly belong to
this species but have not yet entered explicitly into the present concepts
and diagnoses of the genus and species.
Known distribution: As for the genus.
Diagnosis: Only known species of Jnguza. Measurements in Table 3.
ACKNOWLEDGEMENTS
I am greatly indebted to the South African Museum and to Dr Q. B. Hendey
who provided the material for the study of variation in Spheniscus demersus,
arranged the loan of the fossil specimens, gave some further information, and
reviewed the manuscript of this paper. The study has been performed at the
Simroe Foundation under the joint support from that foundation and the
Department of Geosciences of the University of Arizona, Tucson, U.S.A.
The illustrations are by Mr S. Kannemeyer of the South African Museum.
REFERENCES
HENDEY, Q. B. 1974. The late Cenozoic Carnivora of the south-western Cape Province. — Ann.
S. Afr. Mus. 63: 1-369.
McLAcuLan, G. R. & LiversipGe, R., revisers. 1970. Roberts birds of South Africa. Cape
Town: Central News Agency.
Simpson, G. G. 1946. Fossil penguins.— Bull. Amer. Mus. nat. Hist. 87: 1-99.
Simpson, G. G. 1971a. Fossil penguin from the late Cenozoic of South Africa.— Science 171:
1144-1145.
Simpson, G. G. 19715. A review of the pre-Pliocene penguins of New Zealand.— Bull. Amer.
Mus. nat. Hist. 144: 319-378.
Zusi, R. L. 1975. An interpretation of skull structure in penguins.* Jn: STONEHOUSE, B., ed.
The biology of penguins: 59-84. London and Basingstoke: Macmillan.
*This study also refers to bones other than the skull.
6. SYSTEMATIC papers must conform with the International code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. n., sp. n., comb. n.,
syn. n., etc.
An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.
Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (figs 8a—b).
Nucula largillierti Philippi, 1861: 87
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
comma separates author’s name and year
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figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers
Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.
In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:
Holotype
SAM-A13535 in the South African Museum, Cape Town. Adult pone from mid-tide region, King’s Beach,
Port Elizabeth (33.51S, 25.39E), collected by A. Smith, 15 January 1973
Note standard form of writing South African Museum registration ree and of date.
7. SPECIAL HOUSE RULES
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(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. ‘... the Figure depicting C. namacolus...’
*,..in C. namacolus (Fig. 10)...’
(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A. L. du Toit
Von Huene but F. von Huene
(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian
Punctuation should be loose, omitting all not strictly necessary
Reference to the author should be expressed in the third person
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book or article, such as
“Revision of the Crustacea. Part VIII. The Amphipoda.’
Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.
GEORGE GAYLORD SIMPSON
NOTES ON VARIATION IN PENGUINS AND ON
FOSSIL PENGUINS FROM THE PLIOCENE OF
LANGEBAANWEG, CAPE PROVINCE,
SOUTH AFRICA
VOLUME 69 PART 5 MARCH 1976 ISSN 0303-2515
(SAF Clapehrn/
MUS. COMP. ZOOL.
| LIBRARY
MAY 26 1976
ot eT’
OF THE SOUTH AFRICAN
MUSEUM
CAPE TOWN
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Examples (note capitalization and punctuation)
BuLLouGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FiscHer, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques.—J. Conch., Paris 88: 100-140.
FISCHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. —Archs
Zool. exp. gén. 74: 627-634.
Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. —
Ann. Mag. nat. Hist. (13) 2: 309-320. :
Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. —
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
TuHiELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. — Denkschr. med.-naturw. Ges. Jena 16: 269-270. ae
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 69 Band
March 1976 Maart
Part 5 Deel
LEISTOCENE HISTORY AND COASTAL MORPHOLOGY
OF THE YSTERFONTEIN-ELANDS BAY AREA,
CAPE PROVINCE
By
ANTHONY J. TANKARD
Cape Town Kaapstad
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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap
PLEISTOCENE HISTORY AND COASTAL MORPHOLOGY OF THE
YSTERFONTEIN-ELANDS BAY AREA, CAPE PROVINCE
By
ANTHONY J. TANKARD
South African Museum, Cape Town
(With 25 figures and 3 tables)
[MS accepted 4 September 1975]
ABSTRACT
The south-western Cape has been unstable throughout the Neogene and much of the
Pleistocene so that an Early Pleistocene shoreline at 10 m a.s.]. correlates with the 45-50 m
transgression complex of the Namaqualand coast. There were three sea-level peaks in the last
interglacial: at 6,3 m a.s.l., 2-3,5 m a.s.l., and 0 m a.s.l. which are correlated with the
Tyrrhenian II and Tyrrhenian I of Mallorca. No evidence was found of Holocene sea levels
much higher than the present. The Weichselian lowering of sea level was accompanied by
maximum dune activity.
CONTENTS
PAGE
Introduction. ‘ ; 73
Regional geomorphology and pre- Pleistocene geology : TS)
Tectonic setting . , i ; ‘ ‘ 77
Early Pleistocene marine , deposits ‘ : , : ; 80
The 13 m shoreline : : : : : ; 84
Late Pleistocene marine history : ; ; : : 84
Open-coast facies ; : , : F : ; 85
Rocky shores ‘ F ; F F : ‘ 85
Exposed sandy shores. ; : ; ; : 87
Estuarine-lagoonal facies . ‘ : : : : 98
Verlorevlei . é ‘ ; é , : : 98
Berg River . : é ; ‘ : : : 99
Saldanha Bay. : : : : ‘ : 2 «£02
Langebaan Lagoon : : : : : = «103
Age of the deposits . ; ‘ : : ! LOD
Recent. c : : 3 : : : : , 110
Fluvial sediments : . : : ; ‘ a 0
Aeolianites . : : : : ees
Proposed new lithostratigraphic names : : . ae Ss
Conclusions and synthesis. ‘ : ; : aie!
Acknowledgments ‘ : ; : : : 2 116
References . : ; : : : : : ee OLE
INTRODUCTION
The late Cenozoic history of the South African coast is related to the
history of the sea level. In the Pleistocene sea level fluctuated in sympathy with
the repeated waxing and waning of northern hemisphere ice sheets, and palaeo-
geographic studies show that coastal lowlands all over the world have been
subjected to periods of alternating submergence and emergence.
73
Ann. S. Afr. Mus. 69 (5), 1976: 73-119, 25 figs, 3 tables.
74
ANNALS OF THE SOUTH AFRICAN MUSEUM
Elands
Bay
n
\
;
‘
{
ot
LL been
Noo
AFRICA
ape Town
32°30'S
St. Helena Bay
arcyp 4
32°45S
Slippers
Cape
Columbine J Paternoster
be!
Ls
Vredenburg
wy
%»
335 aldanha, Blguwaterbaai
Vf m
1,
Gyp ce poedjespunt 6)
16g-917/ Saldanha
North Head ““ 2 Borngat Bay
Elandspunt — iS
m
° ys"
Jutbaai
South Head C/ Yr ‘ pares
%
33°5)
Y opefield
Kraalbaal
Churchhaven Q eo
Skrywershoekl4 Geelbek Ss
e
(23) OAbrahamskraal 2
Gyp7— +
Ysterfontein
Fig. 1. Locality map showing sampling sites (solid circles). Tidal range is
shown in brackets; evaporite deposits are referred to as ‘Gyp’.
PLEISTOCENE HISTORY AND COASTAL MORPHOLOGY 75
Previous literature on the Pleistocene marine history of the coastal areas of
the south-western Cape is limited to a few texts, mostly of a geomorphic nature.
Krige (1927) published the first major study. He found evidence for two general
emergences: shorelines at 15-18 m he called the Major Emergence, and those
at 6-9 m the Minor Emergence, which was thought to be of Recent origin.
In the Saldanha area he identified the Major Emergence at 6-12 m and proposed
that the area had been downwarped. Haughton (1931) briefly described the
geology and palaeontology. Mabbutt (1956, 1957) claimed marine terraces
at 91 m, 45-60 m, 20 m, 13 m, 9 m, and 7,5 m a.s.l. (above mean sea level).
All of the terraces above 9 m he identified as marine terraces on the basis of
flattish surfaces. The most detailed geological approach to the problem has
been that of Parker (1968). Davies (1973) found evidence of transgressions to
9 m a.s.l. (Eem) and 1,5 m a.s.l. (Recent).
This paper describes the results of an investigation into the Pleistocene
history of the coastal area between Elands Bay and Ysterfontein (Fig. 1). All
altitudes are recorded as height above mean sea level (a.s.1.). Tidal amplitudes
are shown in Figure 1. Mean wave height is 3 m, and the highest recorded
waves were 9,5 m (Pomeroy 1965). Ocean swell is predominantly south-westerly.
Terms such as ‘last interglacial’, ‘last glacial’, ‘Eem’, ‘Weichselian’, etc. are
used purely as time units based on European glacial events.
REGIONAL GEOMORPHOLOGY AND PRE-PLEISTOCENE GEOLOGY
The south-western Cape has a Mediterranean-type climate with hot, dry
summers and mild winters. The temperature range at Cape Columbine is
19-11°C. Average annual rainfall is 262 mm with a winter maximum. The
coastal waters have a temperature range of 13-15°C (Shannon 1966). The
west coast is characterized by active upwelling of cold subsurface water in the
summer months. An evergreen Cape Macchia vegetation predominates and
grasses are rare so that the surface is never adequately protected against wind
erosion. Soils are generally skeletal.
A lowlying area, drained by the Berg River and its tributaries, lies between
the Hottentots Holland Mountains and the Atlantic Ocean, and reaches a
width of 110 km at latitude 33°S. The higher eastern part of this lowland is
known as the Swartland, and the lower western part as the Sandveld. It has
been suggested (Talbot 1947; Mabbutt 1956) that the Swartland was planed
by a shallow sea during the Pleistocene. Other than its plain-like surface there
is no evidence to support this claim. The Swartland and Sandveld are charac-
terized by thick accumulations of Late Pleistocene and Recent dune sands.
Typical of an aeolian landscape, the Sout River has many ponds and marshes.
The pre-Tertiary geology (Fig. 2) comprises rocks of latest Precambrian
and Early Palaeozoic age. The oldest rock assemblage, the geosynclinical
Malmesbury Group, consists of fine-grained greywackes and slates with associ-
ated phyllites, quartzites, and felspathic grits (Truswell 1970). On the farm
76 ANNALS OF THE SOUTH AFRICAN MUSEUM
St. Helena
B
Pleistocene dune & : s :
marine sands and limestones ens: ‘j Cape granites
e Supergroup Malmesbury Group:
rece _ 4 ___ shale and phyllite
Fig. 2. Comparison of geological map with ERTS-—1 photograph shows the effect of lithology
on topography.
Drommelvlei north of Hopefield (32.46 S, 18.25 E) a local consortium drilling
for oil penetrated 3 300 m of subhorizontally bedded shales and siltstones
without reaching the base. In the earliest Cambrian granites were emplaced
into the core of the Malmesbury anticline in two phases of similar age. The
earlier phase of intrusion is marked by a medium- to coarse-grained biotite
granite, and is followed by a finer-grained quartz-porphyry. There are diorites
with gabbro xenoliths at Ysterfontein.
The Cape Supergroup generally follows unconformably upon the Malmes-
bury Group and the Cape Granites. In the study area the Cape Supergroup
mainly comprises mature conglomeratic sandstones overlying red thinly bedded
siltstones, shales and sandstones which are exposed along the southern shore
of Verlorevlei.
Tertiary strata consist of sediments deposited during transgressions in
the Miocene and Pliocene. The Miocene Saldanha Formation consists of micro-
sphorite and phosphatic sandstone. The Pliocene Varswater Formation com-
PLEISTOCENE HISTORY AND COASTAL MORPHOLOGY WL
prises marine sands and gravels, pelletal phosphoritic sands, and estuarine
and fluviatile sands, silts, and peats, with a rich mammalian fauna (Tankard
1974 and in press).
Figure 2 shows the influence of rock type on the regional geomorphology,
where the geology is compared with an ERTS-1 satellite photograph. The
western part of the area is marked by the north-west trending Darling and
Vredenburg plutons which form a hilly terrain. In the north and east the resistant
Cape Supergroup rocks form mountain ranges. From Elands Bay the con-
glomeratic sandstones form a south-east trending range which forms a natural
groyne at Cape Deseada. The northern boundary of this sandstone block is a
fault-scarp. Less resistant Malmesbury rocks form a negative relief. The Berg
River follows the strike of the Malmesbury geosyncline. Whereas the granite
and sandstone feature rocky shores, the St. Helena Bay coastline on the Malmes-
bury Group is entirely sandy. That Malmesbury rock underlies this entire low
coastal plain is shown by the frequent inclusions of that rock in the Pleistocene
marine deposits between Slippers Bay and Cape Deseada.
From a point only 28 km west of Cape Columbine and trending south-
south-west is the Cape submarine canyon (Simpson & Forder 1968). This
canyon has probably affected sedimentation along the St. Helena Bay coastline
by acting as a sediment drain.
TECTONIC SETTING
Several lines of evidence prompt the conclusion that the west coast of the
Cape Province was probably more unstable than the south coast in the Cenozoic.
The Agulhas Arch forms a divide between the broad and gently sloping conti-
nental shelf to the east where the continental margin is controlled by transform
faulting, and the narrow and steeply sloping shelf to the west where tensional
faulting has been the controlling factor (Dingle & Scrutton 1974). The shelf
break east of the Agulhas Arch is at a normal depth, lying between 120 and
180 m (Dingle 1973a). West of the Agulhas Arch the nature of the shelf break
is variable. West of the Cape Peninsula it has an average depth of 450 m, but
west of the Orange River a depth of 200 m (Dingle 1973b). The west coast
shelf break is one of the deepest in the world (Shepard 1963). Simpson (1971)
noted the variation in depth of the west coast shelf break and attributed it to
differential warping of the continental margin. A double shelf break which
appears in places off the west coast (Fig. 3) and which was apparently in exis-
tence during late Lower Tertiary times (Dingle 1973) suggests a long history
of instability.
The Agulhas Arch is a NW-SE striking antiform (Dingle 1973a) coinciding
with the structural trend (NNW) of Late Precambrian origin. Early Tertiary
intrusive dykes follow these same structural trends (Kréner 1973). On the farm
Dikdoorn on the Groen River an intrusive melilite basalt has been dated at
38,5 million years old (ZSA 56). In the Bogenfels area of South West Africa
78 ANNALS OF THE SOUTH AFRICAN MUSEUM
Luderitz
\
\
=< A Klinghardt
ae \ Mountains
\
\
|
3
- \
i :
range
; River
se
oe)
‘Oe
\
am
t x
: \
oP chine
1 i)
\ =
4 1
\ \
~ \o
ea
©.
‘&
100Km
Se. me Agulhas
\. \
Fig. 3. Coastline configuration relative to tilt-axis. Note deep shelf-break which also deepens
to the south.
phonolitic lavas of the Klinghardt volcanism have been dated at 35,7 million
years old (ZSA 53). Early Tertiary igneous intrusives are also encountered
offshore on the Agulhas Arch (Dingle & Gentle 1972).
Tilting probably took place about an axis or ‘hinge line’ which tended
to follow the NNW Precambrian structural lineament and continued through
the Agulhas Arch. The relative position of the ‘hinge line’ has been drawn in
PLEISTOCENE HISTORY AND COASTAL MORPHOLOGY 79
Figure 3 by connecting the Agulhas intrusives with the Klinghardt extrusives.
The axis passes west of the Dikdoorn intrusives. In reality there probably
would not have been a simple well-defined ‘hinge line’ but rather an axial-zone
about which tilting and differential warping would have taken place. It is
suggested that such tilting and warping west of the axis have lowered the shelf
break to abnormal depths. If the shelf west of the Agulhas Arch has tilted in
relation to that east of it, it would be expected that the sedimentation to the
west would be the more complex. Dingle (1971a, 1973a, 19736) has shown just
that. He has subdivided the Tertiary succession on both shelves by correlation
of unconformities which he recognized on seismic reflection profiles. A major
intra-Tertiary unconformity separates Lower from Upper Tertiary on both
shelves. In addition Dingle (19735) found that the Upper Tertiary of the west
shelf is further subdivisible by an unconformity into lower T3 beds (upper
middle Miocene) and upper T4 beds (Pliocene).
Examination of onshore Neogene sediments between Cape Town and
Saldanha has confirmed Dingle’s subdivision of the shelf succession. Tankard
(1974) has documented Middle Miocene deposits overlain unconformably by
Pliocene deposits. These Miocene deposits could possibly be attributed to a
marine transgression which resulted from accretion of mid-ocean ridges at
that time (Tankard in press). Regression of the high Middle Miocene sea con-
tinued through the Late Miocene and Pliocene, and the transgression complex
sediments were for a time subaerially exposed. But tilting again dropped the
area below sea level, causing a local transgression during a time of world-wide
regression. This second transgression resulted in the accumulation of the
Pliocene Varswater Formation.
Tankard (in press) noted that Neogene sediments have beach zones ranging
in altitude from 9 m a.s.l. at Ysterplaat (Cape Town) to 56 m a.s.]. in the Sal-
danha area. But northwards in South West Africa and eastwards along the
Cape south and east coasts, Tertiary marine strata are found at considerably
higher elevations, suggesting relative sagging and tilting of the Cape west
coast since the Miocene. The highest Pleistocene marine sediments in the
Saldanha area are now at 10 m a.s.l. The highest beaches in Namaqualand
are at 98 m a.s.l. (Carrington & Kensley 1969). Furthermore, the mollusc
fauna from the Saldanha 10 m beach correlates most closely with that of the
45-50 m transgression complex of the Namaqualand coast north of Hondeklip
Bay. This fauna includes: Fissurella robusta, Purpura praecingulata, Triumphis
dilemma, the large Perna perna, and Petricola prava.
Tilting about the ‘hinge line’ shown in Figure 3 readily explains the lower
elevation of the Early Pleistocene shoreline in the Saldanha area. Between the
Olifants River mouth and Cape Agulhas the coastline forms a ‘bulge’, and the
coastline between Cape Town and Saldanha is parallel to and furthest removed
from the axis, so that for any degree of tilting shorelines in this area would be
found at lower elevations than those north of the Olifants River where the
coastline and axis almost coincide. Tilting alone would not affect elevation
80 ANNALS OF THE SOUTH AFRICAN MUSEUM
of the raised beaches north of the Olifants River much, but differential warping,
and the local position of the axis certainly could displace them.
Dingle (19735) has shown the existence of an outer shelf bar (T3) on the
edge of the west coast continental shelf. Assuming that this bar formed close
to sea level, it is possible to compute approximate rates of sinking since the
end of the Miocene. Dingle calculated a rate of sinking | m/27 000 years west
of the Orange River, and | m/12 000 years west of the Olifants River. If the
10 m beach at Saldanha is equivalent to the 45-50 m transgression complex on
the Namaqualand coast, and is of a late Early Pleistocene age, say | million
years, an approximate and average rate of sinking since that time can be com-
puted. The average rate of subsidence at a distance from the axis equivalent
to that of the shelf edge west of the Olifants River is 1 m/13 000 years. This
agrees well with Dingle’s estimate since the Miocene. The fact that last inter-
glacial shorelines on the west coast show little variation from the Late Pleisto-
cene 7 m datum implies that subsidence has either been episodic or has been
operating at a decreasing rate. Little, if any, subsidence has occurred in the
last 120 000-130 000 years.
To summarize: volcanic activity from the Agulhas Arch to South West
Africa in the Late Eocene/Early Oligocene shows instability at least in the
Early Tertiary. Tilting of the west coast relative to the south coast has resulted
in a deeper shelf break in the west, and a more complex sedimentary history.
The effect of tilting on onshore deposits is maximized in the Saldanha area
which is furthest removed from the ‘hinge line’. Although tilting has taken
place since the Early Tertiary, and although post-Miocene and post-Early
Pleistocene rates of subsidence appear to be in agreement, field evidence suggests
that the west coast has been essentially stable in the Late Pleistocene. The
southward deepening shelf break suggests that the rate of subsidence has been
greatest in the south. Russell (1964) believes that most of the South African
coastline is unstable.
EARLY PLEISTOCENE MARINE DEPOSITS
Deposits of presumed Early Pleistocene age occur sporadically around
Saldanha Bay: on either side of Hoedjiespunt peninsula, Elandspunt, and 2,1 km
north-east of Langebaan (Fig. 1). The underlying platform on Hoedjiespunt
is uneven, ranging in elevation from 6,2 m to 8,9 m a.s.l. At the Bomgat on
Hoedjiespunt peninsula the Early Pleistocene deposit is separated from the
granite floor by a lens of Miocene microsphorite which occupies a bedrock
depression. The contact between the microsphorite and the limestone is an
erosional one. On the South Head, at Elandspunt, the eroded uneven surface
of the granite is at 7 m a.s.l]. The Early Pleistocene deposit north of Langebaan
rests on the surface of an older marine limestone which is probably of Neogene
age (not an aeolianite as thought by Davies (1973)). Here the contact is at
PLEISTOCENE HISTORY AND COASTAL MORPHOLOGY 81
9.5 ma.s.l. A terrace at the same altitude may indicate a greater lateral extent
of the Early Pleistocene deposits in the Langebaan area.
The relationship of the Early Pleistocene marine limestone at the Bomgat
to the underlying microsphorite and the overlying Langebaan limestone is
shown in Figure 4. The limestone contains wave-generated granite boulders
at the base of 1,5 m of coquina. The matrix consists of micrite, which indicates
a sheltered environment. Although articulated Perna perna would also suggest
a sheltered environment, much fragmented shell debris is indicative of periods
of higher energy marine conditions. Parker (1968) found that calcareous
material constituted more than 95 per cent of the limestone.
The base of the marine limestone on the northern side of Hoedjiespunt
peninsula (behind the Sea Harvest factory) is characterized by wave-generated
granite boulders. The limestone consists partly of unconsolidated shell deposits,
and partly of coquina and microcoquina. The marine limestone is overlain by
approximately 20 m of aeolian limestone. The microcoquina is a medium-
grained skeletal lime grainstone with a drusy calcite cement. Detrital shell
grains form in excess of 95 per cent of the rock, and the remainder is quartz.
The quartz and shell grains are generally well rounded but corroded. The shell
grains include foraminifer and echinoid spine fragments. Although the lime-
Calcrete rubble
Langebaan limestone
Bredasdorp
Formation
Early Pleistocene SOE?
shelly marine
limestone
Miocene transqressive
complex 5 Saldanha
Phosphorite and granite | Formation
boulders
Late Pleistocene boulder
beach with interstitial coarse
shelly quartzose sands
Height above mean sea level (m)
Granite bedrock
Fig. 4. Geological section at Bomgat, Hoedjiespunt.
82 ANNALS OF THE SOUTH AFRICAN MUSEUM
rs 7 "e io
me as ote ok
. \ ieee »
Bas SES REN Was
em “a “ _-
ne sey Sty ou
Mt wal ye,
Le
<
Fig. 5. Shelly limestone on granite platform, Elandspunt.
stone rests on a granite platform, thin-sections show a total absence of felspar
grains. This is characteristic of a high-energy beach environment. The base
of the coquina behind the Sea Harvest factory is in part phosphate mineralized
by phosphate-rich solutions derived from the Miocene phosphorite at the
Bomgat on the other side of the Hoedjiespunt peninsula. Even at the Bomgat
the lower part of the Early Pleistocene limestone shows phosphate
mineralization.
Wave-generated granite boulders occur at the base of the coquina at.
Elandspunt. The coquina consists of much disarticulated and fragmented
bivalve fragments, and less fragmented gastropod remains. The coquina, shown
in Figure 5, is very different from Late Pleistocene deposits (Fig. 11).
The Early Pleistocene mollusc fauna is shown in Table |. The nature of the
outcrop prevented a shell count being made at the Bomgat and Elandspunt
sites. The fauna as a whole is indicative of shallow-water, mainly intertidal,
conditions. Although Bullia annulata is commonly dredged today, it is some-
times found intertidally. All of the species of Patella are intertidal, except
P. tabularis which is infratidal. Littorina knysnaensis is found most abundantly
above HWS (high water spring). The barnacle Balanus amphitrite, which still
encrusts the limestone bedrock at the Langebaan site, is usually indicative of the
infratidal zone.
The Hoedjiespunt molluscs indicate, in general, the close proximity of a
rocky shore. But whereas the Bomgat assemblage is characterized by abundant
PLEISTOCENE HISTORY AND COASTAL MORPHOLOGY
TABLE 1
Early Pleistocene Invertebrate Fauna
GASTROPODA
Haliotis midae Linnaeus .
Fissurella robusta (Haughton) .
Helcion cf. pruinosus (Krauss)
Patella argenvillei Krauss
P. barbara Linnaeus
P. cochlear Born
P. granatina Linnaeus
P. granularis Linnaeus
P. oculus Born
P. tabularis Krauss
P. variabilis Krauss :
Gibbula rosea (Gmelin) .
Turbo cidaris Gmelin
Oxystele trigina (Chemnitz)
Littorina knysnaensis Philippi .
*Rissoa’ capensis Sowerby
Solariella undata Sowerby :
Cerithidea bifurcata Kilburn & Tankard
Crepidula sp. : :
Thais cingulata (Linnaeus)
Purpura praecingulata (Haughton)
Burnupena papyracea cincta (R6ding)
Triumphis dilemma Kilburn & Tankard
Bullia annulata (Lamarck)
B. digitalis Meuschen
B. laevissima (Gmelin)
Nassarius capensis (Dunker)
N. scopularcus Barnard .
N. speciosus Adams
Fusus sp.
Peristernia pemeatula (Lamarck)
Marginella capensis Krauss
Marginella piperata Hinds
Marginella sp.
Cythara amplexa (Gould)
Conus mozambicus mozambicus Hwass
BIVALVIA
Aulacomya ater (Molina)
Perna perna (Linnaeus)
Gryphaea sp. :
Ostrea atherstonei Newton
Mysella convexa (Gould)
Tellimya trigona Barnard
Carditella capensis Smith
Thecalia concamerata Bruguiére
Eucrassatella sp. :
Lutraria lutraria (Linnaeus)
Tivela tomlini Haughton E
Petricola prava Kilburn & Tankard .
CIRRIPEDIA
Balanus amphitrite Darwin
Total individuals
(x = present but not counted. )
Bomgat
Sea
Harvest
— We
Elands-
punt
83
Langebaan
7%
<1
i Nae
he PO PR BR DR
A A
AAA A
[a ee ee NN
A
—
tN
ty
o
84 ANNALS OF THE SOUTH AFRICAN MUSEUM
Perna perna, the Sea Harvest assemblage has fewer bivalves and abounds in
patellids. Although rock-dwelling forms are common at the Langebaan site,
Patella is generally absent. Sand-dwelling forms predominate there. Further-
more, species such as Cerithidea bifurcata and Thecalia concamerata are indica-
tive of sheltered mud flats.
Assigning these deposits to any age with confidence is difficult. Parker
(1968) reports two MC dates for the Elandspunt site: 41 100 B.P. (Pta—097),
and greater than 49 500 B.P. (Pta—098). Besides the fact that the “C technique
is unreliable on carbonate of this age, there is also a distinct possibility that
the molluscs were not collected in situ from the limestone. The number of
extinct molluscs in the fauna seems to preclude a very young age. These include
Fissurella robusta, Cerithidea bifurcata, Crepidula sp., Purpura praecingulata,
Triumphis dilemma, Tivela tomlini, and Petricola prava. The Crepidula possibly
originated from a C. porcellana-type ancestor (Kilburn & Tankard 1975).
Stratigraphically the fauna resembles the 45-50 m transgression complex
fauna of the Namaqualand coast. It has in common Fissurella robusta, Purpura
praecingulata, Triumphis dilemma, the large form (20 cm) of Perna perna, and
Petricola prava. The absence of Donax haughtoni and Striostrea margaritacea,
45-50 m transgression complex zone fossils, is attributed to the higher energy
environment and colder water conditions in the Saldanha area.
Carrington & Kensley (1969) have assigned an Early Pleistocene age to
the Namaqualand 45-50 m transgression complex. This is based on altimetric
correlation with the Moroccan succession, the number of extinct species, the
low degree of lithification, and the fresh appearance of the shells. They also
note that this was the last warm water fauna. Assuming that they are correct
in assigning these deposits to the Early Pleistocene, the rate of tilting, already
discussed, agrees well with Dingle’s (1973) estimate.
THE 13 m SHORELINE
At the southern entrance of the Elandsberg tunnel at Cape Deseada there
is a prominent horizon of large wave-generated boulders banked against the
foot of the cliff at 13 m a.s.l. Another horizon of well-rounded cobbles and
boulders occurs beneath talus material on the southern shore of Verlorevlei
at 13 m a.s.l. No fossils are associated with these two horizons. This shoreline
is pre-Late Pleistocene since it is considerably higher than the Late Pleistocene
sea level datum. Possibly it correlates with the 10 m Early Pleistocene deposits
of the Saldanha area. If younger than that, then the most likely correlation
would be with the Namaqualand 17-21 m transgression complex which Carring-
ton & Kensley (1969) assign to the Middle Pleistocene.
LATE PLEISTOCENE MARINE HISTORY
The geomorphology and mollusc fossils indicate several depositional
environments. An open-coast facies includes rocky shores and exposed sandy
PLEISTOCENE HISTORY AND COASTAL MORPHOLOGY 85
shores. An estuarine-lagoonal facies is identified in the vicinity of Saldanha
Bay—Langebaan Lagoon, Berg River, and Verlorevlei. At Velddrif (Figs 1,
10) lagoonal deposits are situated between breaker-bars of the open-coast
facies. The mollusc fauna of the open-coast facies includes rock and sand-
dwelling forms which still inhabit the adjacent coast. But the estuarine-lagoonal
facies regularly contains several species which today live in tropical waters to
the north.
Description of the marine deposits between Ysterfontein and Elands Bay
will be discussed according to the dominant depositional environment which
also coincides closely with a regional approach. The mollusc fauna has been
described elsewhere (Tankard 1975).
Open-coast facies
Rocky shores
Formation of a platform results from waves breaking in shallow water
and the movement of detritus across the rock surface. All bottom loss of
energy takes place shoreward of the surf base, which Dietz (1963) defines as
the greatest depth where the waves begin to peak appreciably during storms.
The depth of vigorous abrasion is thus 1,5 times the wave height. The abrasion
platform is a function of the energy generated by the average waves. The
depth of the abrasion base off the Cape Peninsula is 15 m (B. W. Flemming
in press). Wave energy is reflected from slopes greater than 30° so that erosion
is slow, while maximum erosion takes place on slopes less than 15° (N. C.
Flemming 1965). Examples of this are, firstly, the Chapmans Peak coast of the
Cape Peninsula where a very steep slope and consequent reflection of wave
energy is associated with little erosion. Secondly, in the Saldanha area slopes
are low and vigorous abrasion has resulted in a well-defined platform.
In the Saldanha area mean wave height is 3 m (Pomeroy 1965), suggesting
that most vigorous abrasion will take place shallower than 4,5 m. Davies (1973)
attempts to identify Holocene platforms which are related to sea levels at
5,4 m, 3,9 m, and 1,8 m a.s.l. The platform on the seaward side of the Vreden-
burg pluton varies in altitude up to 10,7 m (Visser & Schoch 1973), so that
the relief is about 7 m. It is likely that the platform was eroded during a single
stationary sea level. It is highly unlikely that short-term oscillations of sea
level would have left a permanent record on a platform of low relief.
Table Mountain sandstone forms a natural groyne at Cape Deseada
where structually controlled abrasion has formed a narrow platform. The
outer margin of the platform is 4,7 m a.s.l., and the inner margin 6,5 m a.s.l.
The most widespread platform is that bordering the Vredenburg pluton,
broken only by the deep entrance to Saldanha Bay. At Hoedjiespunt bedded
Miocene phosphorite lies in a bedrock depression with a contact at 5,25 m
a.s.l. This suggests that the platform may be an old feature that originated in
the Miocene. Tankard & Schweitzer (1974) have shown that a similar planation
86 ANNALS OF THE SOUTH AFRICAN MUSEUM
surface at 7-8 m a.s.l. at Die Kelders on the southern Cape coast is overlain
by Neogene limestone. But Early Pleistocene limestones in the Saldanha area
suggest that there the platform has been remodelled in the Pleistocene, and that
it is a composite feature. ‘Fresh’ stacks on the North Head platform possibly
originated in the Late Pleistocene. The present sea does not appear to be cutting
a notch or platform at mean sea level. Instead the rocky shore plunges vertically
about 3-4 m before flattening out on to a sandy bottom.
Parker (1968) has mentioned two beach ramparts which parallel the present
coast on the North Head and South Head platforms. Both ramparts, or storm
beaches, are composed of well-rounded granite boulders with a matrix of
coarse sand and comminuted shell. Slight imbrication of the boulders suggests
that storm waves have played little part in forming these ridges. The outer
rampart on North Head is a composite feature. It rises from the shoreline
with a concave surface to a vegetated crest 7 m a.s.]. It thus has a fossil as well
as a modern component. The second rampart (Fig. 6) peaks at 12,1 m a.s.l.,
and is approximately 40-50 m behind the lower and outer rampart. The flat
between them is underlain by coquina and shelly sand typical of an upper
foreshore accretionary unit. It abuts against the inner rampart. Pans have
developed behind the ramparts.
The crests of these ramparts, like breaker-bars, probably relate to MHWS,
which at St. Helena Bay is 0,75 m above MSL. The two ramparts thus reflect
Fig. 6. Excavation through inner storm beach on North Head platform. Elevation of the
crest is 12 maz.s.].
PLEISTOCENE HISTORY AND COASTAL MORPHOLOGY 87
sea levels at 6,25 m and 11,35 m a.s.l. The 7 m rampart would most likely be
of last interglacial age since it agrees with the Late Pleistocene datum, but
the 12 m rampart would be older, and of unknown age.
Shell deposits are extensive on the platform and Tankard (1975, table 1)
has tabulated the mollusc assemblages at two sites, namely Paternoster and
North Head. Parker (1968) has listed the molluscs from other sites. A hori-
zontally bedded shelly sand is exposed below the Paternoster lighthouse. The
bedding takes the form of layers of complete shells separated by layers of
comminuted shell. Whole shells constitute 25-30 per cent of the organic remains.
Gastropods constitute 96 per cent of the assemblage. Most species are rock-
dwelling intertidal forms. Only Patella compressa of the six patellids is infra-
tidal (Branch 1971). Conus scitulus and Cypraea algoensis are common.
The Paternoster deposits are typical of the marine sediments covering the
platform. Rock-dwelling forms are always in excess of 90 per cent, and gastro-
pods predominate. On the North Head platform at the entrance to Saldanha
Bay occasional (< 1%) and stunted Ostrea atherstonei valves occur. There
are also seal bones. Barnacles are common. Due to maximum exposure to
the swell this part of the platform has a great amount of comminuted shell
(> 95°). Beach rounded boulders and cobbles are common, as well as frag-
ments of coquina.
These shell beds regularly occur up to 6 maz.s.l. This and the outer rampart
suggest a Late Pleistocene shoreline dominated by a 6,25 m sea level.
Several localities have features which suggest lower sea levels than that at
6,25 m a.s.]. There is an intertidal occurrence of beachrock at the northern end of
Jutbaai. It is a medium-grained skeletal lime grainstone. The lath-shaped,
well-rounded shell detritus constitutes 95 per cent of the rock. The absence of
felspar is characteristic of a beach environment. The shell and echinoid spine
detritus shows a preferred orientation, and is drusy calcite cemented. Coquina
is preserved at two further localities amongst beach boulders at 1,6 m a.s.l.
and 3,9 m a.s.l.
An horizon of bedded wave-generated boulders is situated within the
Bomgat at 3,5 m a.s.l. The Bomgat is an extension of a gully through the
granite platform through which waves surge high above mean sea level. The
cave deposit was probably formed above high tide level and would have been
related to a sea level between present datum and 2 m a.s.]. A single “C date
(3 500 B.P.: Mr B. Flemming, pers. comm.) derived from mollusc shell collected
in situ from the boulder beach horizon suggests that this feature probably dates
to the Holocene.
Exposed sandy shores
Waves tend to break further from the shore with decreasing gradient so
that there is also a decreasing amount of energy available for shoreline erosion
(N. C. Flemming 1965). With loss of energy in crossing the shallow floor,
88 ANNALS OF THE SOUTH AFRICAN MUSEUM
coarse stirred-up particles are deposited and these lead to formation of breaker-
bars (Holmes 1965). Since the slope of the sea floor adjacent to the Berg River
is only | in 400, one would expect accretion units to develop. But this area is
relatively sediment-starved since perennial runoff is restricted to the Berg River,
which contributes only a small quantity of sediment, and the Cape submarine
canyon possibly acts as a sediment drain.
Late Pleistocene sandy shore deposits are most prominent between Slippers
Bay (west of Berg River mouth) and Cape Deseada. The present shoreline
forms a smooth curve. At Cape Deseada, where the coastal plain is narrowest,
aeolian sands tend to form a concave profile. Further south the coastal plain
becomes more complex.
1. Evaporite deposits
Gypsum deposits are frequently encountered on the flats behind the
coastal dunes (Fig. |). Generally, these evaporite deposits are less than 6 m
in altitude and are believed to be mainly of last interglacial age. The evaporite
deposits north of Dwarskersbos (Fig. |, Gyp 2) and Ysterfontein (Gyp 7)
are probably still forming today since they are associated with salinas. (A
salina is a modern salt pan on an arid coast.) Visser and Schoch (1973) record
several CaSO,.2H,O determinations on these deposits. The gypsum content
of the evaporite deposits usually exceeds 80 per cent. The gypsum usually
occurs in uneven masses of fine-grained texture. Elevations above sea level
are as follows: Cape Deseada (Gyp 1) 4,5-5 m; north of Dwarskersbos (Gyp 2)
3,5-4,5 m; Kruispad (Gyp 3) 3,5 m; St. Helena Bay area (Gyp 4 & 5) 3 m;
Naval Academy (Gyp 6) 2 m; Ysterfontein (Gyp 7) 2 m.
The Cape Deseada occurrence (Gyp 1) was the only one examined in
detail. Here the coastal plain is backed by 180 m high sandstone cliffs. The
sedimentary succession of the terrace from bottom to top is as follows:
(i) shelly calcareous sands overlying the sandstone platform;
(ii) rhythmically laminated (2 mm _ units) carbonate-gypsum-halite units
(Fig. 1);
(111) a thin layer of saline mud;
(iv) modern dune sands which also form extensive coastal dunes.
The shelly calcareous quartzose sands at the base are attributed to a trans-
gression to 6-7 m a.s.l. The mollusc assemblage is indicative of an intertidal
sandy beach, except for Choromytilus which is a rock-dweller. Solen capensis
suggests a sheltered environment which, adjacent to the present high energy
beach, could only have prevailed in a lagoonal environment on the leeward
side of a breaker-bar. An extensive emerged bar which once formed a barrier-
beach follows the St. Helena Bay coastline. In the Cape Deseada region a
salina must have developed behind the bar. Flooding of the salina would have
occurred at spring tides. Precipitation from the stranded brine led to formation
of the evaporite. The evaporite is now about 0,75 m thick and would have
PLEISTOCENE HISTORY AND COASTAL MORPHOLOGY 89
Fig. 7. Cape Deseada evaporite deposit.
required a sea-water column of 45 m for its formation (by extrapolation from
statistics quoted by Hsii 1972).
The evaporite contains a microfauna suggestive of a complex history of
formation. The microfauna is summarized in Figure 8 and Table 2. The fora-
minifers, which constitute only 5 per cent of the assemblage, are all abraded.
In contrast, the ostracodes (95 per cent of the assemblage) are thin-shelled,
show no signs of abrasion, and are frequently still articulated. All growth
stages are also encountered. Candona sp., Pionocypris assimilis, and Cypri-
dopsis ochracea are all freshwater species. Aurila dayii suggests a more saline
environment as found in estuaries and lagoons. Cyprideis cf. limbocostata is
an inhabitant of brackish water, salt lagoons and marsh environments. Species
of Cyprideis have both smooth forms and nodose forms, but the number of
nodose dimorphs decreases with decreasing salinity (Benson 1961). The smooth
tests of Cyprideis cf. limbocostata confirms the freshwater nature of the
assemblage.
Since freshwater ostracodes are scarce in waters more saline than 2%,
(Benson 1961), the Cape Deseada assemblage must reflect low salinities con-
sidering the abundance of these ostracodes. But gypsum will only precipitate
from a brine at salinity of about 117%,, 3,35 times that of normal sea-water.
The apparent contradiction between the ostracode evidence and the gypsum
evidence suggests a complex history of sea-water inflow, evaporation to dryness,
and freshwater inflow (Fig. 9).
ANNALS OF THE SOUTH AFRICAN MUSEUM
90
OSTRACODA
Aglaiella
Perissocytheridea
Loxoconcha
Saldanha
if
18 ¢
oO
££
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ae
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Fig. 8. Distribution of ostracode and foraminifer faunas.
PLEISTOCENE HISTORY AND COASTAL MORPHOLOGY
TABLE 2
Late Pleistocene Microfauna
FORAMINIFERA Sites
Quinqueloculina spp.
Sigmoilina sp.
Frondicularia sp..
Lagena hexagona (Williamson)
Lagena semistriata Williamson
Lagena cf. orbignyana (Seguenza) .
Lagena spp. ‘
Bolivina variably (Williamson)
Bulimina sp. ,
Rotalia beccarii (Linnaeus) . 2
Elphidium alvarezianum (wv Orbigny)
Elphidium macellum (Fichtel & Moll)
Elphidium spp.
Cibicides cf. pseudoungeriana (Cushman)
Cibicides spp. é
Virgulina cf. advena Cushman
OSTRACODA
Eucypris sp.
Paracypretta ampullacea Sars
Cypridopsis ochracea Sars
Pionocypris assimilis Sars
Candona sp.
Paracypris westfordensis Benson & Maddocks
Aglaiella railbridgensis Benson & Maddocks .
Cytheretta knysnaensis Benson & Maddocks .
Cyprideis cf. lmbocostata Hartmann.
Perissocytheridea estuaria Benson & Maddocks
Cytherura sp. :
Hemicytherura parvifossata Hartmann é
Bairdia cf. villosa Brady :
Aurila dayii Benson & Maddocks
Caudites eas Hartmann
Procythereis sp.
Urocythereis sp.
Loxoconcha parameridionalis
Benson & Maddocks
L. peterseni Hartmann
Cytheromorpha sp.
Bradleya sp.
Xestoleberis capensis G. W. Miiller
Cytherella punctata Brady
5
3
4
6
73
21
69
25
7
46
50
="
7
x
12
24
16
56
(Values expressed as percentages separately for Ostracoda and Foraminifera.
but not counted.)
91
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x
em ON
<x
7
6
1
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4
12
x 20925
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Krumbein & Sloss (1963) discuss the origins and associations of evaporite
deposits. Two conditions are necessary for the genesis of evaporites: a warm
and arid climate with little freshwater run-off, and a restricted body of sea-
water. Excessive evaporation at high temperature and little dilution by inflow
of freshwater raises salinity above that of the open sea, and ultimately leads
to precipitation of salt. Calcium carbonate is the first to precipitate when
92 ANNALS OF THE SOUTH AFRICAN MUSEUM
Bobbejaansberg
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Evaporite
4 Calcareous quartzose sand
Table Mountain sandstone
Fig. 9. Summary of the history of evaporite accumulation
at Cape Deseada.
evaporation halves the volume of water. Gypsum is next to precipitate at
about 25 °C, followed by halite after 85 per cent of the gypsum has precipitated.
At Cape Deseada a restricted lagoon has developed behind an emerged
breaker-bar in a warm and arid climate. The mollusc fauna of this transgression
suggests a hydroclimate warmer than today (Tankard 1975). Present-day
rainfall is less than 200 mm/year. Microscopic examination of evaporite speci-
mens shows that each cycle begins with carbonate precipitation, and is followed
by gypsum and frequently by halite. But the absence of halite in some of the
units suggests inflow of more sea-water, probably tidal, before precipitation
of gypsum from the previous brine had been completed. The precipitation
cycle would be terminated at that stage and a new cycle initiated. Since each
carbonate-gypsum-halite unit is, on average, about 2 mm thick, it would have
taken 300 to 400 tidal floodings of the salina (Fig. 9, A-B) to accumulate the
0,75 m of evaporite. The broken foraminifer tests suggest that they have come
from the neighbouring high-energy beaches. Complete evaporation of the brine
was occasionally followed by inflow of freshwater, runoff from the sandstone
hills, to produce freshwater pans (Fig. 9, C). This is suggested by the ostracode
PLEISTOCENE HISTORY AND COASTAL MORPHOLOGY 93
fauna of freshwater affinity (salinity less than 2%,). The articulated valves and
completeness of the thin-shelled carapaces suggest little agitation and the
abundance of juvenile forms suggests that the population was probably sud-
denly exterminated, possibly by another tidal flooding of sea-water.
It is possible that the unique conditions of a saline water—freshwater
cycle could also be satisfied by movement of sea-water through the shelly
breaker-bar, rather than over the top during high tides as suggested above.
The sea-water would at some stage meet and dip beneath ground-water (density
control). Saline or freshwater conditions in the salina area would then depend
entirely upon advance or retreat of the mixing zone due to ground-water
fluctuation.
2. Barrier-beach coast
The entire length of St. Helena Bay coastline is marked by an emerged
breaker-bar which, at some stage, must have formed a barrier-beach. Dune
sands now accentuate the crest of this breaker-bar. But only south of Dwarskers-
bos are there any good exposures.
North-east of Dwarskersbos (2,5 km) surface exposures show the terrace
behind the bar to consist of shell beds with entire valves in a predominantly
detrital shell deposit with pebbles of Malmesbury rock. The shells are not
evenly distributed throughout the exposure. Frequently shells of one species,
e.g. Aulacomya ater, may occur together. The mollusc assemblage is charac-
teristic of an intertidal sandy shore.
The assemblage is dominated by Crepidula capensis (46,7°%), Tellina
trilatera (12%), and Venerupis senegalensis (12%). The predominance of
undamaged valves of 7. trilatera, a thin-shelled species, suggests relatively
low energy conditions such as are presently found on the adjacent shore.
5,5 km south-west of this site the mollusc assemblage is very different.
Crepidula capensis is less dominant, Tellina trilatera is totally absent, and
Lutraria lutraria and Argobuccinum argus make up nearly 70 per cent of the
assemblage. Many barnacles, patellids, and Choromytilus meridionalis suggest
a partly rocky shoreline. There are also many angular Malmesbury rock
fragments. The Malmesbury platform is at 4 m a.s.l.
The Malmesbury platform continues west of the Berg River, and is exposed
behind the Varkvlei homestead where it is associated with rounded boulders
at 6,7-7 m a.s.l. (Visser & Schoch 1973). A deep excavation west of the home-
stead penetrated 4 m of horizontally bedded shell deposits with little detrital
quartz before reaching the Malmesbury platform at | m a.s.l. The platform was
mantled with wave-generated boulders. The shell bed rises gently inland to an
elevation of 6 m a.s.l.
Two breaker-bars are well developed in the Velddrif area (Fig. 10). They
are composed mainly of shell material with little detrital quartz, suggesting
low sedimentation rates. Longitudinal excavations show subhorizontal bedding
with alternating beds of whole shells and fragmented shells.
ANNALS OF THE SOUTH AFRICAN MUSEUM
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PLEISTOCENE HISTORY AND COASTAL MORPHOLOGY 95
The mollusc assemblage of the outer bar north of Laaiplek is dominated
by Crepidula capensis, and Venerupis senegalensis. The top 0,5 m of the bar
has a maximum of whole shells, and Venerupis senegalensis has a high
articulation ratio (6,2). This horizon is underlain by 1,1 m of comminuted
shell. Scissodesma spengleri is common. Its present geographic range is False
Bay to Algoa Bay (Barnard 1964), suggesting slightly warmer conditions in
the Late Pleistocene. The assemblage suggests an intertidal sandy beach.
Figure 11 shows the concentration of shell debris and articulated shells con-
stituting the bar.
Fig. 11. Shell accumulation typical of the bar deposits (north of Laaiplek).
The seaward side of the second bar at Velddrif (site 9) is constructed of
four basic units (Figs. 12-13). The lowest unit is at least 0,5 m thick (base
concealed) and consists of horizontally bedded sand and shell debris. From
2,5 m a.s.l. to 3,6 m a.s.l. whole bivalves increase in proportion. The dis-
articulated valves are convex-up and have a preferred long-axis direction
220°. Then follows 1,4 m of fine quartzose sand with several shell bands con-
taining articulated Perna perna. Other bivalves are convex-up and have a
preferred long-axis orientation 130°. The fourth unit from 5 m-7 m a.s.l.,
consists of a cross-bedded, partially cemented shell deposit, with cross-bed
azimuths 180°-220°.
Wave-energy appears to have been at a maximum in the lower unit where
shell debris is broken down and finer quartz sand winnowed out. Incipient bar
96 ANNALS OF THE SOUTH AFRICAN MUSEUM
Azimuths
Molluscs
long-axes
130°
Height (m.a.s. |.)
concealed
Fig. 12. Measured section through the inner bar and washover-fan
at Velddrif. 1 = Venerupis senegalensis; 2 = Crepidula capensis;
3 = Bullia laevissima; 4 = Lutraria lutraria; 5 = Perna perna;
6 = Other.
development to the west reduced the wave-energy and resulted in a greater
amount of unfragmented shell. But the very thick shells of Venerupis sene-
galensis reflect the high energy conditions near by. Further growth of the
breaker-bar afforded a greater degree of shelter. Energy on the leeward side
was insufficient to winnow away the fine quartz sand. Fragile Perna perna and
Phaxas pellucidus lived in the sand. There are generally more juveniles in these
sediments, and Venerupis senegalensis is thinner shelled. There was only enough
energy to turn the shells over, but not enough energy to fragment them (Fig. 14).
Finally, the bar migrated shoreward over the lower units. Cross-bedding
frequently dips 10°, but dips in the uppermost part of the bar are shallow and
characteristic of a washover fan.
PLEISTOCENE HISTORY AND COASTAL MORPHOLOGY
Fig. 14. Convex-up bivalves in the quartzose sand unit of the inner bar. Note articulated
Perna perna (lower left) (Velddrif).
98 ANNALS OF THE SOUTH AFRICAN MUSEUM
The fine quartz sand reflecting sheltered conditions was not necessarily
brought across the top of the bar, but could have arisen from longitudinal
transport on the leeward side and could have derived from tidal channels.
Another excavation in the bar south of Velddrif shows small-scale cross-beds
suggestive of a tidal inlet.
Figure 11 shows the typical composition of the mollusc assemblage.
Sand-dwelling forms predominate, and of these Venerupis senegalensis always
exceeds 50 per cent, and in the upper cross-bedded unit 81 per cent. Thin-
shelled Perna perna are commonest in the sandy unit. There are also burrows
in the sandy unit, possibly Callianassa.
The crest of the bar (7 m a.s.I.) is a washover fan which relates to MHWS
and can be correlated to a last interglacial sea level of 6,25 m a.s.].
Estuarine-lagoonal facies
Shorelines of the open-coast facies parallel the present shoreline very
closely. Besides Saldanha Bay and its southerly offshoot Langebaan Lagoon,
the Late Pleistocene sea extended up the Berg River valley and along Verlore-
vlei to form two extensive estuaries. Whereas the open-coast facies is continu-
ous, the estuarine-lagoonal facies is discontinuous.
Verlorevlei
Verlorevlei is situated in a north-west trending drowned valley which
was formed along a fault-plane by the ancestral Papkuils, Antonies, and Kruis
Rivers which now feed into the head of the vlei. Fossiliferous sediments are
exposed on the steeper southern bank. The present vlei is 16 km long, and a
maximum of 1,5 km wide. The present mouth of the vlei is marked by a bar
or platform of Table Mountain sandstone at | m a.s.l. A slight rise of sea
level, as in the last interglacial, would effect considerable environmental changes,
although the low bar at the mouth would still ensure a low energy regime.
Recent bridge foundations showed that the deepest channel, and hence the
original mouth, was further to the north.
Geological sheet 3118 C/3218 A shows a sand-covered terrace on the
north bank to 60 m a.s.l. A thick sand cover buries any evidence that may
indicate a marine origin.
Late Pleistocene fossiliferous deposits (sites 1-3 in Fig. 1) are typical of
a salt-water estuarine environment. Maximum thickness (site 3) is 2,2 m where
shelly sands lie unconformably upon Palaeozoic siltstones. The highest contact
is at 5 m a.s.l. where Ostrea stentina was found still adhering to the rock. The
lowest 0,5 m of the shelly sand consists of complete shells and valves, frag-
mented shell, and fine-grained quartzose sand. From 1,5 m to 2,35 m a.s.l. is
a reworked horizon where the original deposit is diluted with coarse, iron-
stained sand. From 2,35 m to 3,8 m a.s.]. the deposit consists of poorly sorted
colluvium with scattered shell debris.
PLEISTOCENE HISTORY AND COASTAL MORPHOLOGY 99
The mollusc assemblage (Tankard 1975, table 1) consists of an infaunal
bivalve element and a minor epifaunal gastropod element. The assemblage is
dominated by Dosinia lupinus (50%), and contains several thermophilic species
such as Tellina madagascariensis, Loripes liratula, Macoma tricostata, Venerupis
dura, etc. The assemblage suggests an estuarine sandy substrate. But Argo-
buccinum argus, Burnupena papyracea, Oxystele variegata, Patella spp., Ostrea
stentina indicate that the shore was in part rocky. The low number of rock-
dwelling forms shows that they were derived from a neighbouring environment
which may not have been extensive.
The ratio of foraminifers to ostracodes (Fig. 8, Table 2) is 9 to 1, indicating
an open exit to the sea. Cytheromorpha sp., which constitutes 49 per cent of
the ostracode assemblage, can tolerate brackish water. Aurila dayii, Loxoconcha
parameridionalis, Xestoleberis capensis, and Cytherella punctata are today
found in estuarine environments (Benson & Maddocks 1964; Hartmann 1974).
They indicate saline water and a sandy substrate. XYestoleberis and Loxoconcha
indicate the presence of marine grasses such as Zostera. The Foraminifera are
dominated by Rotalia beccarii (69°%) and Elphidium alvarezianum (17°%).
Rotalia beccarii thrives in brackish water (Loeblich & Tappan 1964).
Estuarine animals generally colonize the area from mid-tide to low tide
(Emery & Stevenson 1957). Ostrea stentina, which encrusts the rock at 5 m
a.s.l. is always found infratidally. These deposits would thus indicate a mean
sea level in the region of 6 m a.s.l. or more.
The molluscs constitute a mixed life and indigenous death assemblage.
Although some winnowing of juveniles may have taken place, the organisms
appear to have lived in one broad environment reflected in the sediments
(Tankard 1975).
Berg River
The Berg River has not developed a net of tributaries over the sandveld
where infiltration capacity is high. In its lower reaches it meanders and gives
rise to the usual features associated with meandering streams, e.g. oxbow
lakes, meander scrolls. The last interglacial climatic peak was probably asso-
ciated with aridity so that the Berg River did not flow as strongly as today,
and a last interglacial transgression to 6-7 m a.s.l. would have extended saline
conditions far up the valley.
Fossiliferous deposits containing marine molluscs are encountered 15 km
up the Berg River on the farm Kruispad (Fig. 10). The terrace contains evidence
for still-stands of the sea at about 6 m a.s.l. and 3,5 m a.s.l. The furthest inland
extent of the terrace (Fig. 10) has been drawn from aerial photographs. The
inner edge of the terrace is covered with dune sand and calcrete so that it is
impossible to evaluate the significance of this shoreline. Mr D. S. Melck of
Kruispad has determined the furthest inland extent of the shell deposits (Fig. 10)
by hand-augering.
A shallow excavation southeast of the homestead and 15 km from the
100 ANNALS OF THE SOUTH AFRICAN MUSEUM
Berg River mouth, revealed a shell bed beneath 1 m of dune sand. The surface
of the shell bed is 5,1 m a.s.l. The mollusc assemblage is typically estuarine,
and contains several tropical west African species: Nuculana_ bicuspidata,
Loripes liratula, Leporimetis hanleyi, Venerupis dura, and Panopea glycymeris.
Mr S. Kannemeyer of the South African Museum identified the otolith of the
sea eel Tachysurus fossor. Excavation shows Panopea glycymeris to be always
articulated and in life orientation, and on the same plane 0,6—0,7 m from the
surface of the shell deposit, i.e. a siphon-length from the surface. Living Panopea
glycymeris in the Mediterranean is always infratidal, and never extends into
the intertidal zone. This suggests a sea level in the vicinity of 6 m a.s.l. or higher.
The microfauna (Fig. 8, Table 2) consists almost entirely of ostracodes,
and the dominant species are Aurila dayii (27%), Loxoconcha peterseni (16%),
and Xestoleberis capensis (56%). They confirm a shallow estuarine-type environ-
ment. Loxoconcha and Xestoleberis suggest an abundance of Zostera.
The sediment is fine-grained quartzose sand with 8-10 per cent shell
debris. Except for Panopea glycymeris few of the bivalves are articulated.
Panopea is a deep burrower and would have been protected from disturbance.
A dark layer (hydrotroilite) with corroded pollens is situated 0,25 m from the
surface of the shell bed. The pollens include many Gramineae, Chenopodiaceae,
Cyperaceae, Compositae, (Professor E. M. van Zinderen Bakker, pers. comm.).
The Gramineae (grasses) and the Compositae are very varied. Chenopodiaceae
is a small plant that thrives in a salt marsh environment in relatively dry climates.
Cyperaceae (sedge) indicates the presence of water. These pollens suggest that
for a time the site was isolated from open circulation. Hydrotroilite suggests
reducing conditions created by restricted circulation.
An excavation to a depth of 10 m just inland of this site shows that the
shell bed does not continue landward.
There is evidence in the river bank a few hundred metres west of the home-
stead of two separate transgressions (sites 13 & 14). The oldest unit consists
of a semi-consolidated shelly sand with rounded quartz pebbles, 1 m thick,
with a surface 4,5 m a.s.l. The mollusc assemblage which is leached, is charac-
terized by abundance of Mactra glabrata. Solen capensis occurs articulated.
Loripes liratula is the only thermophilic mollusc. This bed grades upwards
through silty loam to a dune sand. The shelly deposit is most likely a continua-
tion of the 6 m transgression complex.
A shell bed dominated by ‘fresh’ Dosinia lupinus is banked against this
older unit up to 3,2 m a.s.l., but extending below river level. At river level
there is an extensive oyster reef 50 m in length consisting of Ostrea algoensis.
Nearer the homestead gypsum deposits indicate former saline conditions with
little dilution by fresh water. The younger shell bed reflects a sea level in the
vicinity of 3,5 m a.s.l.
Other fossiliferous deposits are exposed on the south bank of the Berg
River at Bloemendal (site 11) at 3-4 m a.s.l. and near the bridge at Velddrif.
The Bloemendal assemblage, which contains the thermophilic species Loripes
PLEISTOCENE HISTORY AND COASTAL MORPHOLOGY 10]
liratula and Leporimetis hanleyi, is suggestive of a shallow, sheltered environ-
ment (Tankard 1975, table 1). The bridge assemblage is a mixture of sand-
dwelling and rock-dwelling forms: Patella sp., Choromytilus meridionalis,
Dosinia lupinus, and Venerupis senegalensis. Well-rounded cobbles are sugges-
tive of a surf beach.
Withdrawal of the 6,25 m sea left the barrier-beach and bar complex at
Velddrif emerged, while a lagoon developed in the area between the bars. Aerial
photography (Fig. 15) shows that this area is an old lagoon floor with micro-
relief of shallow channels and sand-bars. Several pits into these fine-grained
lagoonal sediments enabled the collecting of mollusc fossils (site 8). All the
forms were small. ‘Rissoa’ constitutes about 90 per cent of the assembiage.
The assemblage is typical of a shallow, sheltered environment. Nassarius
all
1 Km be 4 %
ee
Fig. 15. Aerial photograph showing relict Late Pleistocene lagoon floor between emerged
bars at Velddrif. (Aerial photograph reproduced under Government Printer’s Copyright
Authority 4603 of 2-11-1971.)
102 ANNALS OF THE SOUTH AFRICAN MUSEUM
kraussianus is abundant in weed-beds at low tide on muddy sands (Day 1969).
Other fossils include crab (Brachyura) chelae, echinoid spines, bryozoan
remains, sponge spicules, ostracodes, and foraminifers.
The ratio of foraminifers to ostracodes is 34 to 1, indicating free access
to the sea. The foraminifer assemblage (Fig. 8, Table 2) is dominated by Elphi-
dium sp. (50%) and Rotalia beccarii (46°). Snider & Curran (1974) have found
large numbers of Rotalia beccarii and Elphidium spp. to characterize the lagoonal
environment. The large numbers of tests per unit volume of sediment indicate
slow sedimentation rates. Carter (1951) has found that sorting governs the size
of the specimens, so that the size of the foraminifer tests would be similar to the
sediment grain size. According to Krasheninnikov (1960 quoted in Loeblich
& Tappan 1964) large numbers of Elphidium indicate mobile water. The tests
of the Velddrif (site 8) specimens show no wear or evidence of transportation.
In comparison with other west coast sites the tests of Rotalia and Elphidium
are small, and are present in large numbers. Phleger (1960) has found that
under optimum conditions, and in large living populations, small size may be
taken to indicate unusually favourable conditions and rapid reproduction.
Cytheromorpha sp. constitutes 49 per cent of the ostracodes, Loxoconcha
peterseni 21 per cent, and Perissocytheridea estuaria 6 per cent. But 8 per cent
of the assemblage comprises freshwater forms: Eucypris sp. (1%), and Para-
cypretta ampullacea (7%), suggesting periodic influxes of fresh water.
Cytheromorpha can tolerate brackish water. Loxoconcha and Xestoleberis prefer
saline water (Benson 1961) and suggest the presence of marine grasses.
These fossiliferous deposits between the bars are at 1,5 m a.s.l. and must
relate to a transgression to about 2 m a.s.l.
Saldanha Bay
Saldanha Bay, the only major inlet on this coast today, has a deep entrance
(42 m) which shoals to 2 m at the mouth of Langebaan Lagoon. The bay
probably originated by differential erosion of the different granite types and
Malmesbury shale. (Drilling operations show the presence of extensive Malmes-
bury shale north and north-east of the shoreline.) There is no evidence of
faulting or of an old river valley.
On the landward side of the Hoedjiespunt peninsula a low plain below
10 m connects Noordbaai with Smitswinkelbaai. Fossiliferous deposits show
that this area was probably flooded by the last interglacial sea. The surface of
the shelly sands is only | m a.s.l. (site 18). The dominant molluscs are Oxystele
variegata (15,7%), Clionella sinuata (19,0%), and Tellimya trigona (26,1 %).
The assemblage is composed of rock-dwelling and sand-dwelling forms, and
indicates a shallow environment.
Limestones form a low cliff along the Blouwaterbaai shore (Fig. 16). The
oldest unit is an indurated aeolianite which is capped with calcrete. At 2,8 m
a.s.l. the calcrete is overlain by 0,6 m of shelly marine limestone which con-
tains angular cobbles of Malmesbury hornfels. The molluscs from this marine
PLEISTOCENE HISTORY AND COASTAL MORPHOLOGY 103
Marine limestone
Calcrete
Aeolianite
Fig. 16. Typical section through Blouwaterbaai limestones.
unit include rock-dwelling forms such as Patella spp., Burnupena papyracea,
etc., and sand-dwelling forms such as Turritella capensis, Bullia digitalis, etc.
They suggest an intertidal deposit. The youngest unit is a thin pavement of
beachrock in the present intertidal zone, and reaches nearly to the foot of the
cliff. Beachrock is formed in the intertidal zone.
Langebaan Lagoon
Extensive shell beds along the southern and western shores of the lagoon
show that the lagoon is a feature which existed at least by the Late Pleistocene.
The lagoon is a north-westerly trending body 15 km long and a maximum of
4 km wide whose present channels are maintained by strong currents generated
by the summer south-easterly winds. Where the eastern shore is controlled by
granite topography the last interglacial and present shorelines probably coincide.
But extensive shell beds on the southern shore on the farms Geelbek, Abrahams-
kraal, and Skrywershoek, demonstrate a more southerly extension in the Eem.
The lagoon is separated from the Atlantic Ocean by a long ridge that
connects Ysterfontein with the South Head granites. One can only surmise
that it developed as a prograding spit and tombolo or a barrier-beach complex.
Whatever its origin, it was reinforced by dune sand accumulation. The present
lagoon is the result of Flandrian flooding of a pre-existing dune landscape.
The Geelbek deposits are a lagoon floor shelly limestone accumulation
reaching an elevation of 2 m a.s.l., and overlain by calcrete. The coquina
includes quartz porphyry cobbles. Many of the bivalves are still articulated.
The mollusc assemblage is indicative of a shallow, sandy substrate, and con-
tains several thermophilic molluscs (Tankard 1975, table 1), for example
Loripes liratula, Tellina madagascariensis, and Venerupis dura.
104 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 17. Planation platform on marine limestone, Churchhaven.
At Churchhaven a shelly limestone has been locally planed at 1,2 m a.s.].
(Fig. 17). Thin-section examination shows the limestone to consist of 90 per
cent quartz and 10 per cent detrital shell with drusy calcite cement. All the
detritus is fine-grained and well rounded, and both quartz and shell are exten-
sively corroded. Most of the shell grains are molluscan, with only occasional
echinoid spines and foraminifer tests. Figure 18, which shows the growth of
calcite scalenohedra on the inside of an ostracode carapace (Urocythereis sp.),
demonstrates the extensiveness of diagenesis in these deposits.
A layer of mollusc shells contains many articulated shells and unbroken
valves. They are predominantly shallow water, sand-dwelling forms. Aurila
dayii and Aglaiella railbridgensis are common ostracodes (Fig. 8, Table 2).
Ichnofossils include crab-burrows (Fig. 19), and some 2-3 cm diameter vertical
burrows, probably Ophiomorpha which is attributed to the marine decapod
Callianassa.
According to Mr B. W. Flemming of the University of Cape Town (pers.
comm.) the erosional platform at Churchhaven is part of an extensive platform
which he has traced for 100 m off Kraalbaai. At Kraalbaai there is a contact
with the marine limestone and overlying aeolianite at 2,5 m a.s.l. (Fig. 20)
with bivalve remains at the contact. Ophiomorpha are common in the marine
limestones at this site. A very interesting trace-fossil that indicates that the
top of the marine limestone at Kraalbaai is partly non-erosional is the series of
PLEISTOCENE HISTORY AND COASTAL MORPHOLOGY 105
st
x
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1 Fa
2;
at
154
Fig. 18. Calcite scalenohedra on inner surface of ostracode carapace demonstrates extensive
diagenesis at Churchhaven.
. 19. Crab-burrows associated with marine limestone at Churchhaven.
106 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 20. Contact of marine limestone and aeolianite at Kraalbaai. Note typical beach low-
angle cross-stratification below the contact.
footprints of a terrestrial mammal, probably the strandwolf Hyaena brunnea
(Fig. 21). The preservation of these footprints, the depth of the print, and the
detail of the print suggest a wet surface. Ridges of sand on one side of each
foot-print demonstrate that the animal was walking on a slope. Together
with ripple-marks on the same plane this suggests a wet beach.
That the platform was eroded by another transgression is shown by the
sedimentary succession at Churchhaven (Fig. 22). The lower marine limestone
platform is overlain by dune sands in which at about 4 m a.s.]. there is an
outwash deposit. The outwash consists of cobbles derived from the lower
limestone along with mollusc shells. Excavation showed that the platform
extends beneath the outwash. The outwash deposit is composed of 90 per cent
cobbles and 10 per cent shells (Parker 1968). The shells are mainly disarticulated.
Parker (1968) recognized two distinct units at Churchhaven which he
separated accordingly to lithology and the mollusc fossils. He recognized that
the lower unit was not a shoreline feature, but a lagoon floor deposit. He
believed the outwash deposit to be a 4,3 m shoreline. Davies (1973), on the
other hand, recognized only one bed into which a 1,5 m transgression incised.
The outwash deposit occurs at varying altitudes in this region, that at
4 m a.s.l. being the highest exposure. Pie diagrams comparing the mollusc
and ostracode assemblages from the two horizons (Figs. 8, 22) confirm Parker’s
interpretation. But the field evidence shows that the shelly conglomerate is
not an in situ marine unit, but rather a colluvial deposit that has gravitated
PLEISTOCENE HISTORY AND COASTAL MORPHOLOGY
Fig. 21. Mammalian footprints at top of marine limestone Kraalbaai, and detail. Note claw
marks in photograph on right.
Dune sand
i a a
Gastrana matadoa
Fig. 22. Sedimentary succession at Churchhaven.
108 ANNALS OF THE SOUTH AFRICAN MUSEUM
TABLE 3
Radiocarbon Determinations on Samples from the Area between False Bay and the
Berg River.
Sample Altitude Material Date
No. Locality (m) dated (B.P.)
Pta—094 Kreeftebaai 3-5 shell 40 200+ 130
—095 Luisterhoek 3-5 shell 48 200 -+- 2 600
— 2200
—096 Churchhaven is shell 48 500 + 3 600
— 2900
—097 Elandspunt 6,5 shell 41 100 + 1 200
—098 Elandspunt 6,5 shell > 49 500
—461 Jutten Point 6-7 shell 2070+ 50
—794 Kruispad SKI shell 43 700 + 4 300
— 2700
—7196 Kruispad Sal shell 39 000 + 1 860
GrN —5803 Melkbos 4 shell 43 200 + 2000
— 1500
—5878 Langebaan Lagoon —1(?) shell 6410+ 45
I—8372 Milnerton 1,5 shell 33 750 + 1 780
-- Muizenberg Inter-tidal beachrock 25 860 + 1040
— 1190
= Muizenberg Inter-tidal beachrock 25 430 + 1050
1210
(References: Parker 1968; Vogel 1970; Vogel & Marais 1971; Davies 1973; Pta—794
and Pta—796, Dr Vogel pers. comm. 30.1.73; I—8372, Dr B. Kensley pers. comm.; Muizen-
berg beachrock, Siesser 1974).
down a dune surface. It must therefore reflect a shoreline higher than 4 m a.s.l.
Although the higher beach material (outwash) may well contain fossils derived
from the lower limestone unit, its mollusc assemblage does appear to be distinct.
Furthermore, the shells are fresher.
The conglomeratic colluvium at Churchhaven implies a more vigorous
abrasion than occurs along the shores of the present lagoon. Two explanations
can be suggested. The first assumes that the peninsula separating the lagoon
from the ocean owes its present extent to dune accumulation during the last
glacial lowering of sea level. (Evidence will be cited shortly to substantiate this
claim.) The width of the tombolo or barrier-beach would have been con-
siderably less than at present. The ratio between the present lagoon width to
barrier width is 2:1, whereas the world average is 6:1 (Tanner 1960). A sudden
rise of sea level, as suggested at Churchhaven, could have partially breached
the barrier, or pushed it back, and reworked the lower limestone. The mollusc
and ostracode fauna in the colluvium demonstrates that sheltered lagoonal
conditions did exist at some stage during this sea level rise.
An alternative explanation could be that the breach of the barrier took
place further north where Kraalbaai is connected with the ocean by a low area.
But it is doubtful that this would have created the vigorous abrasion at Church-
haven. Besides, this gap may be the last remaining evidence of more extensive
breaching or flooding of the original barrier.
PLEISTOCENE HISTORY AND COASTAL MORPHOLOGY 109
For several years oyster deposits (Ostrea atherstonei) have been com-
mercially dredged from the floor of the lagoon where there are reputed to be
reserves of 3 million metric tons. East of Kraalbaai they form a layer 1-3 m
thick 5,5 m beneath the lagoon surface. Oysters generally settle on a hard
substrate and it is possible that initial colonization took place on the limestone
platform. Ostrea atherstonei valves are common in the Geelbek and Skrywers-
hoek deposits as well as among the colluvium at Churchhaven, but rare in
the older limestone. The lagoon floor oyster deposits would most likely correlate
with these last interglacial deposits which in turn are correlated with a climatic
peak at that time (Tankard 1975).
Age of the Deposits
Assessing the age of these deposits depends upon the assumption that they
are primarily of glacio-eustatic origin, and that they can be regionally corre-
lated. Despite the evidence for substantial down-warping in the Pleistocene,
there is good reason to believe that tectonism has been minimal in the last
100 ka or so, and that shorelines below + 7 m can be correlated.
Table 3 summarizes the available “C dates pertaining to former sea levels
between present sea level and + 7 m between Muizenberg (False Bay) and
the Berg River. All the dates except Pta—461 and GrN—5878 are greater
than 20 ka.
MoOrner (1971) discusses some of the difficulties of dating shell material
older than 15 ka. Miniscule contamination of shells older than 20 ka will
give completely spurious dates (Fairbridge 1971). Glacio-climatic evidence
shows that any interstadial glacio-eustatic sea ievels close to present sea level
or higher are unlikely (Fairbridge 1971; Morner 1971; Thom 1973). All of
the dates shown in Table 3 greater than 20 ka would be pre-last glacial.
Only two dates (Pta—461 and GrN—5878) are comparatively young.
The date of 2070 B.P. (Pta—461) was derived from Patella argenvillei which
appears to have been surface collected from a heavily vegetated rampart at
6-7 maz.s.]. (Davies 1973). Even if reliable this date would hardly date the rampart;
Davies mentions the ‘high throw’ of waves along this coast in connection with
the ramparts. The other date (6410 B.P.) was derived from an Ostrea atherstonei
valve dredged from the southern end of Langebaan Lagoon. This date would
appear to be too young because sea level would only have recovered by 5,5 ka
(Godwin et al. 1958). Furthermore, 3 million metric tons of shells implies a
lengthy period of accumulation and optimum conditions for reproduction
which would be better satisfied in the last interglacial. Flandrian transgression
sediments and their mollusc fossils at an equivalent depth below present sea
level show that water temperatures in Saldanha Bay in the mid-Holocene
were no warmer than at present. The Bomgat bed is probably a Holocene
feature (date 3,5 ka, Mr B. Flemming, pers. comm. January 1976).
Available '*C measurements indicate that the marine deposits are mostly
110 ANNALS OF THE SOUTH AFRICAN MUSEUM
older than the range of this technique, and suggest a pre-Weichselian age. The
mollusc fauna indicates an Eem age. Composition of the fauna compared with
older Pleistocene faunas suggests a comparatively young age. Abundance of ther-
mophilic forms correlates best with the warm isotopic substage Se peak of deep-
sea cores (Shackleton 1969) which occurred at 100-120 ka. Chappell (1974a)
points out that the widely recognized shoreline between 2 and 7 m az.s.l. in
areas remote from plate boundaries, and dated as 120-130 ka, is becoming
widely used as a Late Pleistocene sea level datum.
The higher shorelines up to 6,25 m a.s.]. would thus date to about 120 ka,
although the lower shorelines nearer present sea level may be younger. In
Die Kelders cave on the south coast wave generated boulders are preserved up
to 2 m a.s.]. in an erosional strike-passage in steeply dipping Table Mountain
sandstone. Occupation of the cave by Middle Stone Age people followed soon
after withdrawal of the sea and serves to date the boulder bed as pre-Weich-
selian, but not much older. Since the boulder bed would have arisen from wave
surges through the passage it probably reflects a sea level close to the present
and would correlate with the intertidal beachrock at Muizenberg described by
Siesser (1974). An immediately pre-Weichselian age, about 80 ka, is suggested
for these lower units.
RECENT
Figure 23 summarizes the stratigraphy revealed in an excavation 100 m
from the Saldanha Bay shoreline at 18°E. Cross-bedded units of medium and
fine-grained sands with articulated Perna perna, and suggestive of shoreface
sedimentation, form the lowest bed from —4 m to —3,4 m. This passes upwards
into horizontally laminated fine sand and coarse sand with Perna perna, a
typical fining-upward beach sequence. There is a gradual coarsening of the
sediment to —1,4 m suggestive of a temporary regression. Shells are more
fragmented, and the bivalves disarticulated. The assemblage is an intertidal
sand-dwelling one. A period of pedogenesis resulted in calcrete formation.
The calcrete is again overlain by a fining-upward shelly sand due to a final
transgression.
The molluscs include: Turritella capensis, Burnupena papyracea, Bullia
digitalis, B. laevissima, Clionella sinuata, Petricola bicolor, Perna perna, Scis-
sodesma spengleri, Lutraria lutraria, Donax serra, and Venerupis senegalensis.
Drilling operations on the coastal flats adjacent to Saldanha Bay have
shown the widespread occurrence of shelly sands and shelly limestones, with
intercalated calcrete horizons, below sea level datum. Furthermore, several
boreholes into the bay floor north of Hoedjiespunt penetrated a peat at —22 m
which was preserved beneath calcrete (Mr A. De La Cruz, pers. comm.).
FLUVIAL SEDIMENTS
At several localities from Cape Town to Verlorevlei there is evidence of
considerable fluvial activity in the past. These sediments are presumably of
PLEISTOCENE HISTORY AND COASTAL MORPHOLOGY 111
Pleistocene age, although extensive deltaic sediments on the continental shelf
between Dassen and Robben Islands have been assigned to the Lower Tertiary
(Dingle 19716).
0
5
Fine sand a
a
ra
©
14:20 Be e8sty)| Shelly sand
Calcrete
Cc
oO
w
Shelly sand @
oO)
| w
24
a Fine to medium
£
= sand
c
a)
o
a
@ 34264: re rse shelly vy
oO LAG ‘ Aad sand ©
£ Fine horizontally a
S laminated sand o
3 =
Coarse shelly sand
Shoreface
Cross bedding
4
Perna perna in coarse sand
Turritella , Burnupena, Bullia, Clionella
Venerupis, Petricola, Scissodesma, Donax.
Fig. 23. Flandrian sedimeniary succession at Saldanha.
112 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 24. Relict fluvial sediments at head of Verlorevlei.
The planed surface of the Malmesbury Group at 30 m a.s.l. east of the
Sout River is an old river terrace and is mantled with angular gravel (the
farms Maatjesfontein and Hamburg in the Hopefield area).
Tankard (1974) has discussed in detail fluvial sediments at Langebaan-
weg, the Baard’s Quarry deposits. The deposits of the northward draining
channels were mineralized by phosphate derived from the Varswater Formation.
The mammal fossils suggest an Early Pleistocene age.
At Ysterplaat (Cape Town) an exposure of cross-bedded, rounded, quartz
gravel is suggestive of a point-bar deposit, while opposing dips laterally
separated through the section suggest a meandering stream. Another exposure
through relict fluvial sediments occurs at Redelinghuis (Fig. 24) at the eastern
extremity of Verlorevlei (26 km from the mouth). Here at least 18 m of river
sediment occupy an old channel with a thalweg less than 6 m (concealed)
above vlei level. The sediment is predominantly fine-grained, iron-stained,
quartzose sand, with lenticular, gravelly, channel lag deposits which show
some imbrication of the pebbles.
Extensive fossiliferous deposits demonstrate that the ancestral Berg River
valley cutting was pre-Eem. Consolidated fluvial gravels and bog-iron-ore are
exposed in the bank at Swartjiesbaai. Weichselian lowering of sea level to
—130 m lowered the controlling base level of the Berg River. Visser & Schoch
(1973) mention four boreholes for the Velddrif bridge foundations which
PLEISTOCENE HISTORY AND COASTAL MORPHOLOGY 113
intersected at least 4,3 m of conglomerate beneath a 5 m cover of shelly sands
which probably originated during the Flandrian rise of sea level.
AEOLIANITES
The coastal plain is covered with a veneer of dune sands and aeolianites.
Du Toit (1917) referred to the aeolianites as Dorcasia limestone, while Visser
& Schoch prefer the name Langebaan limestone since the common fossil
land-snail is Trigonephrus globulus and not Dorcasia. These authors present a
lengthy discussion of the aeolianites.
Several exposures suggest that the aeolianite may range in age back to
the Neogene. In a quarry 4 km north of Saldanha high-angle cross-bedded
aeolianite overlies marine Bredasdorp Formation rock of probable Miocene
age. Behind the President Jetty on Saldanha Bay a well-lithified, fine-grained
aeolianite underlies younger aeolianites.
Most of the aeolianites appear to be much younger, and probably accumu-
lated during the last glacial lowering of sea level when vast tracts of unvegetated
sand lay exposed on the emerging sea floor. Several occurrences of interbedded
outwash suggest that the climate was not necessarily arid. Cross-bedding
azimuths at Kraalbaai (mean 360°) show that a southerly wind was prevalent.
Rogers & Du Toit (1909) describe a well at Paternoster which penetrated
sandy limestone containing land-snail shells and tortoise bones to 21 m below
sea level. Tankard & Schweitzer (1974) have described the former seaward
extension of dunes in the Die Kelders area, and also mention in situ Middle
Stone Age artefacts in the outwash deposits which shows that there the aeoli-
anites must be Weichselian or younger. Furthermore, limestone blocks in the
cave show that lithification of the aeolianites occurred prior to about 6 000 B.P.
Similar inter-bedded outwash deposits containing Middle Stone Age artefacts
occur as far north as Saldanha. A single “C assay on ostrich eggshell from a
midden in the limestones behind the Sea Harvest factory gave an age greater
than 40 000 years (UW 282).
An interesting feature of the coastal limestones are the solution-pipe
cavities which have formed by karst weathering. At Churchhaven and Velddrif
they penetrate the last interglacial marine limestones and shell beds. The pipes
are 0,3-0,8 m in diameter and vertical. At Churchhaven they are marked by a
3 cm thick lithified crust. All the pipes are filled with non-calcareous terrestrial
terra rosa type sediment. Blackburn et a/. (1965) described an identical situation
from Australia and suggest that they arose from solutional weathering and the
filling of the voids with sediment by simple gravitation. The terra rosa sediment
together with the karst weathering and colluvial deposits suggest a wetter
climate in the south-western Cape in the last glacial.
Extensive Late Stone Age shell middens overlie the Langebaan limestone
on the North and South Heads. On the South Head the midden is in part
calcretized.
114 ANNALS OF THE SOUTH AFRICAN MUSEUM
PROPOSED NEW LITHOSTRATIGRAPHIC NAMES
The Bredasdorp Formation comprises a variety of limestone types of
marine and aeolian origin. It includes marine limestone with conglomerate
and coquina horizons, and beds consisting mainly of whole shells. The major
part of the formation consists of dune sand and ‘shell grit and locally grades
into pure shell-beds; it has been calcified into rocks varying from crumbly
calcareous sandstone to hard, crystalline limestone. . . . Solution channels
are often encountered in the limestone’ (Spies et a/. 1963). Lithologically this
description describes the marine and aeolian limestones and shell beds of the
Saldanha area. Although locally the limestones may be indistinguishable
lithologically, it is usually possible to separate the aeolian and marine com-
ponents. Two members of the Bredasdorp Formation are here defined:
1. Velddrif Member: the type-section is the bar exposure at Velddrif
(Figs 12-13). Lithologically it consists of unconsolidated beds of shell and
comminuted shell, locally cemented, and with little quartz. But it ranges through
limestones with shell layers to coquina, for example at Churchhaven.
2. Langebaan Limestone Member: type-section on Hoedjiespunt behind
the Sea Harvest factory where it is over 20 m thick. It consists of fine to medium-
grained calcareous sand and limestone with fine shell detritus and complete
shells of the land-snail Trigonephrus globulus. This land-snail is distinctive
enough to feature as a lithological component. The member includes outwash
horizons and Middle Stone Age middens.
CONCLUSIONS AND SYNTHESIS
Tilting and warping since the Miocene have displaced marine units from
their original elevations, so that the equivalent of the Namaqualand 45-50 m
transgression complex (Carrington & Kensley 1969) is situated at only 10 m
a.s.l. in the Saldanha area. Sediments equivalent to the Namaqualand 17-21 m
transgression complex are possibly submerged in the Saldanha area. But there
is good reason to believe that warping has been minimal in the Late Pleistocene.
Evidence for stillstands of the sea in the last interglacial are not always
unequivocal. There is no difficulty in recognizing shorelines, but it is frequently
difficult to discriminate between major stillstands and lesser features. In sum-
mary, data from the south-western Cape identifies three stillstands of the sea
in the last interglacial, with the shoreline at 6,3 m a.s.l. being a major feature.
These stillstands are at 6,3 m a.s.l. (C I), 2-3,5 m a.s.l. (C ID), and sea level
(C Til).
The platform on the seaward edge of the Vredenburg pluton was carved
partly in the Neogene, remodelled in the Early Pleistocene, but owes its present
relief to Late Pleistocene abrasion. Relief of the platform is such that it could
have been carved by a single stationary sea level. The outer ramparts on the
North Head and South Head platforms were formed by a sea level in the
PLEISTOCENE HISTORY AND COASTAL MORPHOLOGY 115
region of 6,3 m a.s.l. Shell beds on the platform agree with this interpretation.
Similarly shell beds on the Malmesbury platform west of the Berg River mouth
are related to a major stillstand at 6,3 m a.s.l., as are the extensive barrier-beach
and breaker-bar complex of the St. Helena Bay coastline. That there are few
indications of lower last interglacial shorelines along the open coast suggests
that this stillstand was a major one.
Shell deposits in the former Verlorevlei and Berg River estuaries have
maximum elevations of 5 m a.s.l. Infratidal molluscs still in their living positions
suggest a mean sea level at least at 6 m a.s.l. But the fauna in general, which
in estuaries would inhabit the mid-tide to low-tide zone (Emery & Stevenson
1957), shows that this sea level could not have been much above 6 m, and
thus agrees with the C I transgression.
Evidence for a transgression to 2-3,5 m (C II) consists of a shell bed
which is banked against older C I sediments on Kruispad, the lagoon sediments
between the emerged bars at Velddrif, Blouwaterbaai sediments, Geelbek tidal
flat sediments at 2 m a.s.]., and the Kraalbaai aeolianite-marine limestone
contact at 2,5 m a.s.l.
A boulder bed in the Bomgat on the Hoedjiespunt peninsula obviously
owes much of its present elevation to waves surging up the passage through the
granite in front of the cave. Considering that the elevation of this boulder bed
is only 3,5 m a.s.]. it could be attributed to very heavy surf during high tides.
The ‘fresh’ appearance of the shells which still have their colour preserved would
suggest a Recent age, and the elevation is certainly not excessive given the
physical setting of the cave. Samples were submitted for radiocarbon dating
even though groundwater contamination seems likely. A date of 3,5 ka suggests
a Holocene rather than a last interglacial origin. This site illustrates some of
the difficulties encountered in raised shoreline studies.
At four sites there is evidence for a sea level in the last interglacial coin-
ciding with the present beach (C III). This includes beachrock at Muizenberg
(Siesser 1974), Blouwaterbaai, and Jutbaai. Wave-generated boulders beneath
Middle Stone Age sediments in Die Kelders cave are attributed to waves surging
into the cave from about the present position of sea level.
These results are summarized in a graph of Late Pleistocene eustatic sea
level changes (Fig. 25). That portion of the curve from about 47 ka to the
present is based on “C dates of submerged material, and will be fully discussed
elsewhere. The last interglacial part of the curve agrees well with the shorelines
on Mallorca (Butzer & Cuerda 1962). It is suggested that the C I shoreline
correlates with their Tyrrhenian (T) Ila, C II with T IIb, and C III with T III.
According to Butzer & Cuerda it is the T II shoreline (5-10 m) which contains
thermophilic molluscs, and Tankard (1975) has suggested correlation of the
C I thermophilic fauna with that sea level. Broecker ef al. (1968) have dated
three Eem transgressions on Barbados: B III (6 m a.s.l.) at 122 ka, B II at
103 ka, and B I at 82 ka, and Chappell (1974) has dated three sea-level peaks
on New Guinea at 120 ka, 100 ka, and 80 ka. Alternatively, the thermophilic
116 ANNALS OF THE SOUTH AFRICAN MUSEUM
8 10 12 14
6
Age ka
Fig. 25. History of sea level oscillation over last 120 ka shown in graph form. CI = 6,3 ma.s.l.;
Gil 2-3 5 mia. Cie —Oinnras
mollusc content of the C I and C II shorelines suggests the distinct possibility
that the three last interglacial peaks of the south-western Cape may all correlate
with the warm peak at 120 ka, and that the two lower shorelines would then
represent only temporary halts in a regression from the 6,3 m level.
It is not known whether the C II and C III shorelines represent temporary
stillstands in regression from the major C I stillstand, or whether they are in
fact transgressive and separated by regressions.
Withdrawal of the sea with the build-up of high latitude glaciers left vast,
unvegetated tracts of sand exposed on the emerging shelves, and these were
blown into dune fields by wind systems which may have increased in intensity
with the onset of glaciation. But in the south-western Cape they do not signify
an arid climate. On the contrary, there is evidence that precipitation was in
fact higher than today at times.
There is little evidence to suggest Holocene sea levels much higher than
the present.
ACKNOWLEDGEMENTS
The writer wishes to acknowledge with thanks the hospitality of Mr &
Mrs D. Melck of the farm Kruispad, and Captain W. G. van der Merwe,
Officer Commanding SAS Saldanha, for permission to survey the Naval
Academy property. Professor E. M. van Zinderen Bakker kindly undertook
pollen analyses. The writer has benefited greatly from discussions with Drs D. K.
Hobday and W. G. Siesser. Mr V. Branco drew the illustrations. This project
was supported by a grant-in-aid from the Council for Scientific and Industrial
Research.
PLEISTOCENE HISTORY AND COASTAL MORPHOLOGY 117
REFERENCES
BARNARD, K. H. 1964. Contributions to the knowledge of South African marine Mollusca.
Part V. Lamellibranchiata.— Ann. S. Afr. Mus. 47: 361-593.
BENSON, R. H. 1961. Ecology of ostracode assemblages. Jn: Moore, R. C., ed. Treatise on
invertebrate paleontology. (Q) Arthropoda (3): Q56—Q63.
BENSON, R. H. & Mappocks, R. L. 1964. Recent ostracodes of Knysna Estuary, Cape Pro-
vince, Union of South Africa.—Paleont. Contr. Univ. Kans. 34: 1-39.
BLACKBURN, G., Bonp, R. D. & CLARKE, A. R. P. 1965. Soil development associated with
stranded beach ridges in south-east South Australia. — Soil Publ. C.S.I.R.O. Aust. 22: 1-65.
BRANCH, G. M. 1971. The ecology of Patel/a Linnaeus from the Cape Peninsula, South Africa.
1. Zonation, movements and feeding. — Zoologica afr. 6: 1-38.
BROECKER, W. S., THURBER, D. L., GODDARD, J., Ku, T—L., MATTHEWS, R. K. & MESOLELLA,
K. J. 1968. Milankovitch hypothesis supported by precise dating of coral reefs and deep-
sea sediments.— Science 159: 297-300.
Butzer, K. W. & CUERDA, J. 1962. Coastal stratigraphy of southern Mallorca and its implica-
tions for the Pleistocene chronology of the Mediterranean sea.—J. Geol. 70: 398-416.
CARRINGTON, A. J. & KENSLEY, B. F. 1969. Pleistocene molluscs from the Namaqualand coast.
—Ann. S. Afr. Mus. 52: 189-223.
Carter, D. J. 1951. Indigenous and exotic Foraminifera in the Coralline Crag of Sutton,
Suffolk.— Geol. Mag. 88: 236-248.
CHAPPELL, J. 1974a. Late Quaternary glacio- and hydroisostasy, on a layered earth. — Quatern.
Res. 4: 405-428.
CHAPPELL, J. 19746. Relationships between sea levels, 0'8 variations and orbital perturbations
during the past 250 000 years. — Nature, Lond. 252: 199-202.
Davies, O. 1973. Pleistocene shorelines in the western Cape and South-West Africa.— Ann.
Ntl Mus, 21: 719-765.
Day, J. H. 1969. A guide to the marine life of South African shores. Cape Town: A. A. Balkema.
Dietz, R. S. 1963. Wave-base, marine profile of equilibrium, and wave-built terraces: a
critical appraisal. — Bull. geol. Soc. Am. 74: 971-990.
DINGLE, R. V. 1971la. Tertiary sedimentary history of the continental shelf off southern Cape
Province, South Africa.— Trans. geol. Soc. S. Afr. 74: 173-186.
DINGLE, R. V. 19716. Mesozoic and Tertiary sedimentary history of the continental shelf
between Cape Columbine and Cape Seal.—Tech. Rep. SANCOR mar. geol. Progm.
3: 28-37.
DINGLE, R. V. 1973a. Post-Palaeozoic stratigraphy of the eastern Agulhas Bank, South
African continental margin.— Mar. Geol. 15: 1-23.
DINGLE, R. V. 19735. The geology of the continental shelf between Luderitz and Cape Town
(southwest Africa), with special reference to Tertiary strata.—J. geol. Soc. Lond. 129:
337-363.
DINGLE, R. V. & GENTLE, R. I. 1972. Early Tertiary volcanic rocks on the Agulhas Bank,
South African continental shelf.— Geol. Mag. 109: 127-136.
DINGLE, R. V. & ScruTTON, R. A. 1974. Continental breakup and the development of post-
Paleozoic sedimentary basins around southern Africa.— Bull. geol. Soc. Am. 85: 1467—
1474.
Du Torr, A. L. 1917. Report on the phosphates of Saldanha Bay.— Mem. geol. Sury. S. Afr.
10: 1-38.
Emery, K. O. & STEVENSON, R. E. 1957. Estuaries and Lagoons. Jn: HEDGPETH, J. W., ed.
Treatise on marine ecology and palaeoecology. Vol. 1. Ecology. Mem. geol. Soc. Am. 67:
673-750.
FAIRBRIDGE, R. W. 1971. Quaternary shoreline problems at Inqua, 1969.—Quarternaria 15:
1-18.
FLEMMING, B. W. In press. Rocky Bank—evidence for a relict wave-cut platform.— Ann.
S. Afr. Mus. 71.
FLEMMING, N. C. 1965. Form and relation to present sea level of Pleistocene marine erosion
features. —J. Geol. 73: 799-811.
Gopwin, H., SuGGATE, R. P. & WiL.is, E. H. 1958. Radiocarbon dating of the eustatic rise
in ocean-level.— Nature Lond. 181: 1518-1519.
118 ANNALS OF THE SOUTH AFRICAN MUSEUM
HARTMANN, G. 1974. Die Ostracoden des Untersuchungsgebiets.— Mitt. Hamburg. Zool.
Mus. Inst. 69: 229-520.
HAUGHTON, S. H. 1931. The late Tertiary and Recent deposits of the west coast of South
Africa. — Trans. geol. Soc. S. Afr. 34: 19-57.
Homes, A. 1965. Principles of Physical Geology. London: Nelson.
Hsu, K. J. 1972. Origin of saline giants: a critical review after the discovery of the Mediter-
ranean evaporite. — Earth-Sci. Rev. 8: 371-396.
KiLBuRN, R. N. & TANKARD, A. J. 1975. Pleistocene molluscs from the west and south coasts
of the Cape Province, South Africa.— Ann. S. Afr. Mus. 67: 183-226.
Krice, A. V. 1927. An examination of the Tertiary and Quaternary changes of sea-level in
South Africa, with special stress on the evidence in favour of a recent world-wide sinking
ocean level.— Annals Univ. Stell. V, Sectn. A: 1-81.
Kroner, A. 1973. Comments on ‘Is the African Plate stationary ?’ — Nature, Lond. 243: 29-30.
KRUMBEIN, W. C. & SLoss, L. L. 1963. Stratigraphy and Sedimentation. 2nd ed. San Francisco:
Freeman.
Loes.icu, A. R. & TAPPAN, H. 1964. Foraminifera. Jn: Moore, R. C., ed. Treatise on inverte-
brate Paleontology (C) Protista 2(1): C 55-C164.
MassuttT, J. A. 1956. The physiography and surface geology of the Hopefield fossil site.
—Trans. R. Soc. S. Afr. 35: 21-58.
Massutt, J. A. 1957. Some Quaternary events in the winter rainfall area of the Cape Province.
In: CLARK J. D. & CoLe S. eds. Third Pan-African congress on prehistory, Livingstone,
1955: 6-13. London: Chatto & Windus.
Morner, N.-A. 1971. The position of the ocean level during the interstadial at about 30 000
B.P.—a discussion from a climatic-glaciologic point of view.— Canadian J. Earth Sci. 8:
132-43.
ParRKeER, R. J. 1968. Eustatic shorelines of Saldanha Bay. Unpublished B.Sc. (Honours) thesis,
Cape Town University.
PHLEGER, F. B. 1960. Ecology and Distribution of Recent Foraminifera. Baltimore: Johns
Hopkins Press.
Pomeroy, A. S. 1965. Notes on the physical oceanographic environment of the Republic of
South Africa. Cape Town: Jagger Library, University of Cape Town.
Rocers, A. W. & Du Tort, A. L. 1909. An introduction to the geology of the Cape Colony.
London: Longmans, Green & Co.
RuSSELL, R. J. 1964. Techniques of eustasy studies. —Z. Geomorph. 8: 25-42.
SHACKLETON, N. J. 1969. The last interglacial in the marine and terrestrial records. —Proc.
R. Soc. Lond. B. 174: 135-154.
SHANNON, L. V. 1966. Hydrology of the south and west coasts of South Africa.—Jnvestl.
Rep. Div. Sea Fish., S. Afr. 58: 1-22.
SHEPARD, F. P. 1963. Submarine geology. 3rd ed. N.Y.: Harper & Row.
SressER, W. G. 1974. Relict and Recent beachrock from Southern Africa.— Bull. geol. Soc.
Am. 85: 1849-1854.
Simpson, E. S. W. 1971. The geology of the south-west African continental margin: a review.
— Rep. Inst. geol. Sci. 70/16: 153-170.
Simpson, E. S. W. & Forpber, E. 1968. The Cape submarine canyon. Part 1. Bathymetry. —
Bull. Div. Sea Fish. S. Afr. 5: 35-38.
Sniper, E. B. & CurRAN, H. A. 1974. Distribution and ecology of estuarine and salt marsh
Foraminifera.— A. Mtg. geol. Soc. Am. 8 (N.E. Sectn., Abstr.): 75.
Spies, J. J., ENGELBRECHT, L. N. J., MALHERBE, S. J. & VILJOEN, J. J. 1963. Die geologie van
die gebied tussen Bredasdorp en Gansbaai. An explanation of sheets 3419 C and 3419 D
(Gansbaai) and 3420 C (Bredasdorp).— Pretoria: Geological Survey.
TALBoT, W. J. 1947. Swartland and Sandveld. Cape Town: Oxford Univ. Press.
TANKARD, A. J. 1974. Varswater Formation of the Langebaanweg-Saldanha area, Cape
Province. —Trans. geol. Soc. S. Afr. 77: 265-283.
TANKARD, A. J. 1975. Thermally anomalous Late Pleistocene molluscs from the south-western
Cape Province, South Africa.— Ann. S. Afr. Mus. 69: 17-45.
TANKARD, A. J. In press. The marine Neogene Saldanha Formation.— Trans. geol. Soc. S. Afr.
TANKARD, A. J. & SCHWEITZER, F. R. 1974. The geology of Die Kelders cave and environs:
a palaeoenvironmental study.—S. Afr. J. Sci. 70: 365-369.
PLEISTOCENE HISTORY AND COASTAL MORPHOLOGY 119
TANNER, W. F. 1960. ‘Perched’ barrier islands, east Florida coast.—Southeastern Geol. 2:
133-135.
THom, B. G. 1973. The dilemma of high interstadial sea levels during the last glaciation. —
Progress in Geography 5: 167-246.
TRUSWELL, J. F. 1970. Introduction to the historical geology of South Africa. Cape Town:
Purnell.
VissER, H. N. & ScHocu, A. E. 1973. The geology and mineral resources of the Saldanha Bay
area.— Mem. geol. Surv. S. Afr. 63: 1-150.
VoGEL, J. C. 1970. Groningen radiocarbon dates IX.— Radiocarbon 12: 444471.
VoGEL, J. C. & Marais, M. 1971. Pretoria radiocarbon dates I.— Radiocarbon 13: 378-394.
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6. SYSTEMATIC papers must conform with the International code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. n., sp. n., comb. n.,
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Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (figs 8a—b).
Nucula largillierti Philippi, 1861: 87
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
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ANTHONY J. TANKARD
PLEISTOCENE HISTORY AND COASTAL
MORPHOLOGY OF THE YSTERFONTEIN-—
ELANDS BAY AREA, CAPE PROVINCE
VOLUME 69 PART 6 APRIL 1976 ISSN 0303-2515
aS /f- C aper a
L Bee
pe 24 1 1976
OF THE SOUTH AFRICAN
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BULLouGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FiscHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. —J. Conch., Paris 88: 100-140.
FISCHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines.— Archs
Zool. exp. gén. 74: 627-634.
Konan, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. —
Ann. Mag. nat. Hist. (13) 2: 309-320.
Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. —
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
THEE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. — Denkschr. med.-naturw. Ges. Jena 16: 269-270. :
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 69 Band
April 1976 April
Part 6 Deel
ANCHISAURUS CAPENSIS (BROOM)
AND A REVISION OF THE ANCHISAURIDAE
(REPTILIA, SAURISCHIA)
by
P. M. GALTON & M. A. CLUVER
Cape Town Kaapstad
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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap
ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE
ANCHISAURIDAE (REPTILIA, SAURISCHIA)
By
P. M. GALTON
University of Bridgeport, Bridgeport, U.S.A.
&
M. A. CLUVER
South African Museum, Cape Town
(With 13 figures and 3 tables)
LMS accepted 22 October 1975]
ABSTRACT
A complete description of the skeleton of the type specimen (SAM-990) of the pro-
sauropod dinosaur Gyposaurus capensis Broom is given. Comparison with other prosauropods
indicates that this dinosaur represents a valid South African species of Anchisaurus Marsh,
1885 from the Upper Triassic of North America. It is considered that the infra-order Pro-
sauropoda should be divided into three families, viz. Anchisauridae, Plateosauridae and
Melanorosauridae. It is proposed that the family Anchisauridae be restricted to prosauropods
with relatively slender feet, and that broad-footed forms previously assigned to the Anchi-
sauridae be transferred to the Plateosauridae. The Family Anchisauridae is therefore con-
sidered to include the genera Anchisaurus, Efraasia, Thecodontosaurus and several indetermi-
nate species. The genera Ammosaurus, Aristosaurus, Massospondylus (including Gryponyx,
Aetonyx and Dromicosaurus), and Lufengosaurus (including Yunnanosaurus) are included in
the Plateosauridae, while Arctosaurus and Ischisaurus are referred to the suborder Theropoda.
‘Thecodontosaurus’ gibbidens, Spondylosoma absconditum and Teleocrater alphos are placed
within the Thecodontia, and Tanystropheus primus and T. latespinatus in the order Lacertilia.
Thus constituted, the family Plateosauridae becomes the dominant and most widespread
prosauropod family, while the Anchisauridae, in contrast, is known from a geographically
and numerically restricted fossil record.
CONTENTS
PAGE
Introduction : : : 2 122
Redescription of Anchisaurus capensis (Broom) : : «; 2124
Vertebral column . : ‘ : : : : ‘ 124
Pelvic girdle . ; : - ‘ : ‘ : : 126
Hind limb ‘ : : : : 129
Slender and broad- footed Prosauropods : ; ~ : 131
Systematic discussion . : E ; f : : 132
The Family Anchisauridae 3 ; ; ; ; 3 132
Anchisauridae nomina dubia. : : 141
Prosauropod species incorrectly assigned to Anchisauridae . 143
Prosauropoda nomina dubia : : : z 150
Non-prosauropod species incorrectly assigned to.
Anchisauridae . : : 150
Notes on the Families Anchisauridae and Plateosauridae F 153
Summary . ‘ F : ; : ; ‘ 5 155
Acknowledgements : : : : : ; : : 155
References ; : ¢ F : : ‘ : ; 156
Abbreviations. i : : F ' , : i LSS
121
Ann. S. Afr. Mus. 69 (6), 1976: 121-159, 13 figs, 3 tables.
122 ANNALS OF THE SOUTH AFRICAN MUSEUM
INTRODUCTION
Broom (1906) described a specimen (SAM-990) from the Cave Sandstone,
Stormberg Series (Upper Triassic) of Ladybrand, Orange Free State, South
Africa, and referred it to the prosauropod taxon Hortalotarsus skirtopodus
Seeley, 1894. The holotype of Hortalotarsus skirtopodus was referred to the
genus Thecodontosaurus Riley & Stutchbury, 1836, by Huene (1906) as 7.
skirtopodus and, because SAM-990 differed in several aspects from Theco-
dontosaurus, Broom (1911) made it the holotype of Gyposaurus capensis.
Broom (1906) noted several resemblances between SAM-990 and Anchisaurus
Marsh, 1885, from the Upper Triassic of North America, so he referred Gypo-
saurus capensis (SAM-990) to the Family Anchisauridae Marsh, 1885. Galton
(1973, in press) provisionally accepted the validity of Gyposaurus capensis
but noted that the holotype should be carefully compared with Anchisaurus
polyzelus (Hitchcock) to determine whether or not these species are generically
distinct. A comparison of photographs of SAM-990 with a specimen of Anchi-
saurus polyzelus (YPM 1883) showed that Gyposaurus is a junior synonym of
Anchisaurus and that SAM-990 should be redescribed.
Huene in several papers between 1906 and 1932 made important contri-
butions to an understanding of the Family Anchisauridae (as Thecodonto-
sauridae Lydekker, 1890). Charig et al. (1965) considerably enlarged the family
by referring to it genera of Triassic theropods (Family Gryponychidae =
‘Palaeosauridae’) based solely on postcranial material which was _indis-
tinguishable from that of prosauropods. The generic list of Romer (1966: 370)
includes these changes and the comprehensive list given by Steel (1970) faithfully
but rather uncritically records all the genera and species referred to the family.
A taxonomic revision of the Family Anchisauridae is necessary for several
reasons:
1. Several of the suggested and generally accepted synonymies are probably
incorrect. In assessing these the recognition of slender- and broad-footed
types (Galton 1971, 1973, in press) is useful.
2. The skeletal anatomy of prosauropods is remarkably uniform, so taxa
should be based on specimens which include either most of the specimen or
bones which are diagnostically different from those of other prosauropods.
Because of the limited number of skeletal variations it is necessary to diagnose
taxa on the basis of a combination of characters which, if each was taken in
isolation, would not be diagnostic. Unfortunately several of the taxa listed by
Steel (1970) are based on specimens which are generically and specifically
indeterminate.
3. The group has been over-classified and insufficient account has been
taken of the range of individual variation possible within a dinosaurian species;
this can be quite extensive, as shown by the prosauropod Lufengosaurus huenei
Young (see Rozhdestvensky 1966) and the ornithopod Hypsilophodon foxti
Huxley (see Galton 1974).
4. A few of the taxa included within the family are not prosauropods.
ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 123
Fig. 1. Anchisaurus capensis (Broom). Type specimen (SAM-990) showing vertebral column
(top), pubis, ischia and pes (middle) and ilium, femur and fibula (bottom).
124 ANNALS OF THE SOUTH AFRICAN MUSEUM
Summaries of certain aspects of the revision of the family Anchisauridae
have been published earlier (Galton 1971, 1973, in press). The first author
(P. M. G.) is responsible for all sections except the description and illustration
of SAM-990, which is the work of the second author (M. A. C.).
REDESCRIPTION OF ANCHISAURUS CAPENSIS (BROOM)
The specimen (SAM-990) is preserved in a soft sandstone matrix (Fig. 1).
The bone is generally not well preserved and is inclined to crumble during
preparation. Much of the specimen (including most of the vertebral column) is
preserved as bone impression, and details of certain of these portions were
obtained from silicone rubber positives. Chief portions of the preserved skeleton
are 17 vertebrae (including, probably, dorsals, sacrals and caudals), an incom-
plete left pubis, the right ilium and pubis and both ischia, and the right femur,
fibula and pes. Several other fragmentary bones, mostly seen as impressions,
are scattered through the block: anteriorly a portion of the right scapula blade
can be made out, while impressions of 13 ribs lie ventral to the dorsal vertebrae.
VERTEBRAL COLUMN (Fig. 2)
Altogether 17 vertebrae are preserved, some very incompletely. Eleven of
these are in articulation and consist of a number of dorsals and possibly two
sacrals. A space separates the last of these from the first of the posterior group,
which have been displaced to the right of the anterior series. The space is
sufficient to accommodate three vertebrae of the size of those on each side of
it, and there is thus the possibility that this gap was originally filled by three
sacral vertebrae. However, as will be shown below, it is more likely that a
parting of the vertebral column between two sacral vertebrae occurred prior
to fossilization of the specimen.
The anterior three vertebrae, imperfectly seen, are not in natural articula-
tion, although still in relatively close association with each other. The centrum
of the second vertebra is opisthocoelous and slightly convex anteriorly, while
the third centrum is concave anteriorly and posteriorly. The fourth vertebra has
a procoelous centrum and is markedly convex posteriorly. This convex rear
meets the apparently convex anterior surface of the fifth vertebra’s centrum,
which also appears to be opisthocoelous. The sixth vertebra is provided with
a biconcave centrum, the neural spine is broad in lateral view, and its postero-
dorsal edge overhangs the postzygapophysis so that a posterior embayment is
formed above the postzygapophysis. The seventh vertebra resembles the sixth
and the succeeding eighth in the shape of the neural spine, and both the seventh
and eighth vertebrae nave biconcave centra, similar to that of the sixth vertebra.
The eighth vertebra shows the neural spine clearly, and probably represents
the condition which existed in the less complete sixth and seventh vertebrae.
The spine, posteriorly situated, is antero-posteriorly lengthened and fairly low.
The neural spine of the ninth vertebra is shorter antero-posteriorly. The
ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 125
Fig. 2. Anchisaurus capensis (Broom). SAM-990. A. Vertebral column in lateral view. Broken
edges shown in dashed outline, hatched areas seen in section only. > 0,5. B. Mould of centra
of dorsal vertebrae.
126 ANNALS OF THE SOUTH AFRICAN MUSEUM
tenth vertebra has the only complete neural spine and there is a shallow notch
above the postzygapophysis. The ninth and tenth vertebrae are possibly pro-
coelous, with a convexity on the posterior articular surface.
The posterior group of six vertebrae lies slightly to the right of the anterior
row, and appears to be in near-natural association with the right ilium. The
spine of the most anterior vertebra (the twelfth in the column as preserved)
is fairly high and slightly rounded anteriorly; there is only a slight posterior
notch above the postzygapophysis. The spine resembles that of the tenth (and
eleventh?) vertebra of the anterior row fairly closely. The thirteenth vertebra
has a high and narrower spine, posteriorly inclined, and a chevron can be seen
extending back and down from below its centrum. The fourteenth and fifteenth
vertebrae are similar, as far as can be seen. Remnants of chevrons are seen
between the centra of vertebrae fourteen to seventeen.
Identification of vertebral types
Caudal vertebrae are fairly clearly represented by nos. 13 to 17 in the rear
series, with narrower, obliquely inclined spines and a series of chevrons. The
last two vertebrae (10 and 11) of the anterior row have narrower and possibly
higher spines than the preceding ones, and they resemble the first member of
the posterior group. From this it can be argued that the tenth, eleventh and
twelfth vertebrae of the column as a whole are sacrals, separated by the dis-
integration of the pelvic girdle. Vertebra 12 is in fairly natural association
with the right ilium, and 10 and 11 are close to the ilium of the left side. The
left ilium has been displaced sideways and forward relative to the right side,
and this could account for the gap in the series. Moreover, no recognizable
loose vertebral portions can be identified in the surrounding matrix.
In Efraasia (Galton 1973) and Plateosaurus (Huene 1926) there are fifteen
dorsal vertebrae so, if the above interpretation is correct, the specimen as
preserved probably includes dorsal vertebrae 7 to 15, sacral vertebrae 1 to 3,
and caudal vertebrae | to 5.
Dimensions of the vertebrae, numbered according to their above identi-
fication, are given in Table 1.
PELVIC GIRDLE
The pelvic girdle is represented by the right ilium, both ischia, and an
incomplete left and almost complete right pubis. The right side of the pelvis
(Figs 3, 5) is preserved in almost natural association. The ilium is well pre-
served, but both pubis and ischium are incomplete distally, and do not make any
clear contact with each other. The areas of articulation between ischium, pubis
and ilium are imperfectly preserved and could not be determined.
Ilium
The ilium (Figs 3A, 5) of the right side is uncrushed and complete except
for a portion of the dorsal crest. It is characterized by long anterior and posterior
ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 127
TABLE 1.
Dimensions of Vertebrae (mm). Anchisaurus capensis (Broom) SAM-990
Length Height of
Length of between pre- spine above
centrum Maximum and post- post- Length of
Vertebra ventrally height zygapophysis zygapophysis spine
7th dorsal . ' 27 — — — ==
8th dorsal . : 29 — — = =
9th dorsal . : +30 — — = —
10th dorsal . : 30 os — — _—
11th dorsal . ; 31 — — — —_
12th dorsal . : 31 40 — 13 30
13th dorsal . : aS 42 40 16 32
14th dorsal . 3 33 40 48 14 30
15th dorsal . i= 32 38 44 13 20
Ist sacral . : 31 43 39 19 18
2nd sacral . ; — — — == ae
3rd sacral . ; _ = a5 18 21
Ist caudal . 18 51 34 22 13
2nd—Sth caudals ; +18 — = ae ==
processes, and a pre-acetabular process considerably longer than the post-
acetabular process. The anterior process lies slightly external to the more
posterior surface of the bone, and extends as far forwards as the anterior
edge of the pre-acetabular process. The slender finger-like form of the anterior
process (Fig. 10B) is similar to that of Anchisaurus polyzelus (Fig. 10A) and
Ammosaurus (Galton 1971) and in contrast to the small triangle of other pro-
sauropods (Fig. 10C—D).
A prominent pre-acetabular buttress is developed, arising from close
above the tip of the pre-acetabular process and flaring out laterally before
merging with the body of the ilium at the base of the postacetabular process.
The body of the ilium is expanded to a certain extent above the buttress. The
postacetabular process is considerably shorter than the pre-acetabular, and takes
no part in the formation of the buttress. Above the postacetabular process
the illum is continued posteriorly as a short crest, medial to the base of the
posterior process. The maximum length of the ilium is 130 mm.
Pubis
The proximal part of the pubis (Fig. 3A) is incomplete, and the areas of
articulation with the ilium and, to a lesser extent, the ischium are not fully
preserved. The ventral edge of the bone is deeply notched below the proximal
end, and this appears to be a natural condition. Below this embayment, which
represents an open obturator foramen, the pubis curves medially and forwards
to terminate as a horizontal plate with an average width of 24 mm. The length
of the pubis, as preserved, is 145 mm.
For prosauropods an open obturator foramen is described to date only
128 ANNALS OF THE SOUTH AFRICAN MUSEUM
ant. proc.
A
pub. ped
Fig. 3. Anchisaurus capensis (Broom), SAM-990. A. Pelvic girdle in right lateral view.
B. Ischia in ventral view. x 0,5.
ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 129
in SAM-990 (Fig. 10H) and Anchisaurus polyzelus (Fig. 10G, K); in all other
prosauropods it is enclosed ventrally as in Efraasia (Fig. 10J, N). A similar
open obturator foramen is present in most theropods (for Al/osaurus and
Ceratosaurus see Gilmore 1920). Romer (1923) noted that as a result of the
more vertical orientation of the archosaurian femur there is a trend amongst
archosaurs to reduce that part of the pelvis equivalent to the central portion
of the pubo-ischiadic plate of primitive reptiles. The loss of the ventral border
of the obturator foramen in Anchisaurus (Fig. 10G, H, K) and most therapods
probably represents the loss of that portion of the m. pubo-ischio-femoralis
externus 2 which originated ventral to the acetabulum in most prosauropods
and in all sauropods (see Romer 1923: fig. 2, Camarasaurus).
Ischium
Both ischia (Fig. 3), with a preserved length of 136 mm, are present,
but are incomplete posteriorly. In each the widened proximal portion is curved
outwards and carries two embayments, one above a ventral hook-shaped
keel and the other, less clearly defined, lying more dorsally. The proximal
portions of the ischia are separated by an ovoid space, but the shafts are closely
appressed and form a dorsally open trough.
HIND LIMB
Femur
The femur (Fig. 4A), which is seen in dorsal (anterior) view, is broad
and fairly powerful but, with a preserved length of 194 mm, it is incomplete
proximally and distally. The proximal end, as shown by what is still preserved,
was inclined fairly sharply inwards. Below the proximal end the femur is
strongly built and raised to a smooth crest, which runs from proximo-laterally
to disto-medially where it merges into the flat distal end. No condyles are
preserved, and only an indication of the base of the fourth trochanter can
be made out, high in the upper half of the bone.
Fibula
The right fibula (Fig. 4B) lies in its natural position between the femur
and the pes and is seen in lateral (external) view. The proximal half is stoutly
built and leads to a crest formed in the middle of the bone, directed sideways
and forwards. The distal part of the bone is slender and tapers off to the articu-
lation with the pes. Both proximal and distal ends are incomplete and the
preserved length of the bone is 176 mm.
Pes
The pes (Figs 4C, 6, 11K) is seen in ventral view, the bones being either
eroded or indicated by bone impressions. The calcaneum is incomplete laterally,
and probably extended out slightly farther than shown. The astragalus is
transversely elongated, with a rounded anterior surface curving back to the
130 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 4. Anchisaurus capensis (Broom), SAM-990. A. Right femur in dorsal (anterior) view.
B. Right fibula in lateral view. C. Right pes in ventral view. 0,5.
narrow lateral corner of the bone. Both calcaneum and astragalus are pre-
served as impressions, both approximately 15 mm long.
Two small distal tarsal elements are preserved, probably nos 3 and 4.
No. 3 is no more than a bony nodule, while 4 is more robust with posterior
and medial surfaces at right angles to each other and a convex anterior surface
facing metatarsals IV and V. The considerable space between the astragalus
and the proximal ends of metatarsals I and II suggests that the two distal
tarsals are laterally displaced.
All five metatarsals are preserved, in several cases as somewhat imperfect
impressions. There is the usual overlapping of the proximal ends with each
metatarsal slightly underlying its medial fellow. Metatarsal V is short and
narrow but provided with a strong and wide base. Metatarsal IV is about
twice as long and of more or less equal width over its entire length. Meta-
ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 13]
tarsal III is the longest and most robust of the series, while II is slightly shorter
than IV. The distal ends of metatarsals III and IV are squarely truncated but
metatarsal II terminates in an oblique surface, so that the medial inclination
of the row of phalanges seems to be a natural one. This could be true, too,
of the short metatarsal I, although this element is incomplete distally.
A small fragment of bone in front of the fifth metatarsal probably repre-
sents a vestigial phalanx. The form and degree of preservation of the phalanges
are apparent from Figures 4C and 6, and from Table 2.
TABLE 2.
Anchisaurus capensis (Broom), SAM-990. Dimensions of pes (in mm).
Meratarsal Phalanx length
length (proximal to distal)
Metatarsal I 44 Digit I 30 34
II 76 II 30 22 31
Ill 86 Ill 30 Lift 14 30
IV a IV 24 16 12 12 21
Vv 48 Vv =
SLENDER- AND BROAD-FOOTED PROSAUROPODS
When the feet of anchisaurids and plateosaurids are drawn so that digit II
of the manus (Fig. 7) or digit III of the pes (Fig. 8) are reduced to unit length
then two groups are distinguishable, those with slender feet (Figs 7A, C—D, I,
8D-G, 81) and those with broad feet (Figs 7B, E-H, J-Q, 8A-—C, H, J-S).
Fig. 5. Anchisaurus capensis (Broom), SAM-990. Stereophotograph of right ilium.
132 ANNALS OF THE SOUTH AFRICAN MUSEUM
The difference is clearest for the manus (digits II to IV) and, where both fore-
and hind feet are known, the manus and pes are both slender (Figs 7C, E, I,
8D) or broad (Figs 7B, 8H; 7E, 8K; 7G, 8L; 7J, 8N; 7K, 80; 7Q, 8Q). The
difference between slender and broad feet is not growth related since there
are small prosauropods with broad feet (Figs 7B, 8A—C, H). These appear to
be juveniles of species that grew much larger (Figs 7B, H, K, 8H, O-P—all
Ammosaurus major, see Galton 1971) and all the larger prosauropods are
broad-footed. The prosauropod families Plateosauridae (Figs 7-8) and Melano-
rosauridae (see Bonaparte 1972a: figs 62, 70; Raath 1972: figs 9f, 10a—b) are
all broad-footed, whereas the Anchisauridae as currently classified include
both slender and broad-footed forms (Figs 7-8). “Gyposaurus’ capensis (Figs 3C,
5, 7G, 10K) and Thecodontosaurus antiquus (Figs 8F, 11G) are both slender-
footed species but broad-footed species have been incorrectly referred to both
genera.
SYSTEMATIC DISCUSSION
Order SAURISCHIA
Suborder SAUROPODOMORPHA
Infra-order PROSAUROPODA
Family Anchisauridae Marsh, 1885
Diagnosis
Smaller forms, skull lightly built, shallow posterior half of lower jaw with
articulation in line with tooth row, manus and pes slender.
Genus Anchisaurus Marsh, 1885
(includes Megadactylus Hitchcock, Amphisaurus Marsh, Gyposaurus Broom)
Diagnosis
Centra of dorsal vertebrae low, broad bases to neural spines of anterior
caudal vertebrae, ilium with long anterior process, pubis with open obturator
foramen and a relatively narrow distal part that is not apron-like.
The characters of the skull, neck and manus listed below (p. 133) for
Anchisaurus polyzelus may also be diagnostic of the genus, but these regions
are not known in A. capensis.
Anchisaurus polyzelus (Hitchcock, 1865)
Megadactylus polyzelus Hitchcock, 1865: 40, pl. 9 (fig. 6). Cope, 1870: 122A-G, pl. 13 (preocc.).
Amphisaurus polyzelus Marsh, 1882: 84 (preocc.).
Anchisaurus polyzelus Marsh, 1885: 169; 1892: pl. 16 (fig. 3) pl. 17 (fig. 6); 1896: 147, pl. 3
(figs 4-5). Lull, 1915: 119, figs 14-17; 1953: 99, figs 12-14a. Galton, 1971: 782, fig. 7C;
(97/32 sfie- EP, EME Or PSS:
Anchisaurus colurus Marsh, 1891: 267; 1892: 543, pl. 15, 16 (figs 1, 2); 1893: 169, pl. 6; 1896:
148, pl. 2 (figs 1-3), pl. 3 (figs 1, 2), pl. 4. Huene, 1906: 6, figs 1-6, pls 1-3; 1914b: 69,
figs 1-11; Lull, 1912: 414, figs 2-3; 1915: 130, figs 18-21, pls 4, 10.
Thecodontosaurus polyzelus Huene, 1906: 19, figs 10, 10a; 19146: 75, figs 23-24; 1932: 116.
Yaleosaurus colurus Huene, 1932: 119, pl. 14 (fig. 1), pl. 54 (fig. 3). Lull, 1953: 107, figs 15-18,
pl. 4.
ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 133
Types
Incomplete skeleton (AM 41/109) from Longmeadow Sandstone, upper
part of Newark Series (Upper Triassic) of Springfield, Massachusetts, U.S.A.
(A. polyzelus). Almost complete skeleton (YPM 1883) from Portland beds,
upper part of Newark Series near Manchester, Conn., U.S.A. (A. colurus).
Diagnosis
+9 maxillary teeth, 16 dentary teeth, basipterygoid processes very small,
cervical vertebrae elongate, metacarpal I broad, digits If and III of manus
subequal in length, ungual I of pes smaller than ungual II.
Discussion
Megadactylus Hitchcock being preoccupied, Marsh (1882) replaced it with
Amphisaurus (also preoccupied) and then Anchisaurus (Family Anchisauridae
also proposed). Huene (1906) referred the material of Megadactylus polyzelus
Hitchcock to Thecodontosaurus as T. polyzelus (Hitchcock) and used the Family
Thecodontosauridae (originally proposed by Lydekker, 1890: 246) to replace
Anchisauridae. Marsh (1891) made YPM 1883 the holotype of a new species
of Anchisaurus, A. colurus, but he did not indicate how it differed from A. poly-
zelus (AM 41/109). Huene (1906) suggested that Anchisaurus polyzelus resembled
Thecodontosaurus and differed from Anchisaurus colurus in several features,
discussed here together with others noted later by Huene (1907-08, 1932) (for
full discussion see Galton, in press):
1. Shortness of cervical vertebrae. Huene (1932) noted that AM 41/109
resembled Thecodontosaurus (Fig. 11A) in the shortness of the anterior
cervical vertebrae, which are elongate in YPM 1883 (Figs 9H, 11B). How-
ever, this comparison was based on misidentification of part of the neural
arch of a dorsal vertebra (Galton in press, fig. 3a—c) as a cervical vertebra
(Huene 19145: fig. 23a).
2. Shortness of dorsal vertebrae. Huene (1906) originally noted that AM
41/109 differed from Thecodontosaurus and resembled YPM 1883 in having
very elongate dorsal vertebrae. Later Huene (1914) figured an extremely
short centrum of AM 41/109 as that of a dorsal vertebra. Although not
stated, this implied that the dorsal vertebrae of AM 41/109 are extremely
Short as in Thecodontosaurus. The isolated centrum figured by Huene
(19146) could not be located but judging from the proportions it was
probably part of an anterior caudal vertebra. The proportions of an
isolated neural arch (Galton in press, fig. 3a—c) and of a centrum (Galton
in press, fig. 5c) show that the dorsal vertebrae of AM 41/109 were probably
elongate (i.e. the centra were low) as in YPM 1883 (Fig. 9K).
3. Slenderness of neural spines of anterior caudal vertebrae. This comparison
by Huene (1906) was based on misleading figures given by Cope (1870)
and Marsh (1893, 1895, 1896). Cope (1870: pl. 8 (fig. 7): see Lull 1953:
134 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 6. Anchisaurus capensis (Broom), SAM-990. Stereophotograph of right pes in
ventral view.
fig. 12a) did not indicate that the neural spines of the anterior caudal
vertebrae of AM 41/109 were damaged and incomplete (Fig. 9P); originally
the neural spines were broader and not as narrow as in the vertebrae
referred to Thecodontosaurus (Fig. 9S) by Huene (1907-08: pl. 77 (fig. 4);
19146: fig. 40).
Most of the supposed differences between AM 41/109 and YPM 1883 were
either the result of misinterpretation (1-3 above, different position of fourth
trochanter of femur) or the result of differences in preservation (form of radius,
metacarpals, tibia, fibula: see Galton in press). YPM 1883 does differ from
AM 41/109 in having a proportionally long centrum to the last dorsal vertebra
(but this might be sacral vertebra 3) and a proportionally smaller manus with
a less trenchant first ungual phalanx (Fig. 7A, C). However, these differences
probably represent individual variations within a species because individuals
of the ornithopod dinosaur Hypsilophodon foxii show a much wider range of
morphological variation (see Galton 1974). YPM 1883 should be referred to
Anchisaurus polyzelus because, on the basis of available material, it cannot be
distinguished from AM 41/109 by any characters of taxonomic significance
and, in addition, AM 41/109 does not show any unique resemblance to Theco-
dontosaurus. Consequently, Anchisaurus colurus Marsh is a junior synonym of
Anchisaurus polyzelus (Hitchcock) and, as A. colurus is the type species of the
135
ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE
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136 ANNALS OF THE SOUTH AFRICAN MUSEUM
genus Yaleosaurus (Huene 1932: 122), Yaleosaurus is a junior synonym o!
Anchisaurus Marsh, 1885.
Anchisaurus capensis (Broom, 1911)
Hortalotarsus skirtopodus (non Seeley, 1894) Broom, 1906: 201, pl. 3.
Gyposaurus capensis Broom, 1911: 293.
Type
Partial skeleton (SAM-990) from the Cave Sandstone, Stormberg Series
(Upper Triassic) of Ladybrand, Orange Free State, South Africa.
Diagnosis
Ungual | largest on pes.
Discussion
Because of its nature, SAM-990 can be distinguished from Anchisaurus
polyzelus (YPM 1883) oniy by the relative size of ungual | of the pes—large
in SAM-990 and small in YPM 1883. Anchisaurus capensis (Broom) is dis-
tinguishable from all prosauropods other than A. polyzelus (Hitchcock) by
the following combination of characters:
1. Centra of posterior dorsal vertebrae (about the tenth) are proportionally
L :
low so that the ratio of central length to height Cy) is 2,1 (Figs 2, 9L).
2. Anterior process of ilium is elongate (Figs 3, 5, 10B).
3. Subacetabular part of pubis is emarginated ventrally so that the obturator
foramen is open (Figs 3, 10H).
4. The pes is slender (Figs 4C, 6C, 11K).
5. Broad bases to neural spines of anterior caudal vertebrae (Figs 2, 9Q).
The form of these elements in the species of Anchisauridae is summarized
in Table 3 to facilitate comparisons. The systematic position of other species
incorrectly referred to ‘Gyposaurus’ are discussed below (pp. 141, 143, 147).
TABLE 3.
Comparison of species attributed to ‘Gyposaurus’ with other anchisaurids.
Dorsal Neural Proximal
Centra: spine of Anterior pubis|
length] anterior — process of obturator
height caudal ilium foramen Pes
“Gyposaurus’ capensis . 2,1 wide long shallow/open slender
*‘Gyposaurus’ erectus 1,4 wide short deep/closed broad
“Gyposaurus’ sinensis ; if | — short deep/closed broad
Anchisaurus polyzelus ; pi | wide long shallow/open slender
Thecodontosaurus antiquus 1B 9/ narrow short —_— slender
Efraasia diagnostica DS narrow short deep/closed slender
ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 137
Genus Efraasia Galton, 1973
Diagnosis
Basipterygoid processes of medium length, cervical vertebrae elongate,
centra of dorsal vertebrae low, narrow bases to neural spines of anterior caudal
vertebrae, slender metacarpal I, digit II of the manus robust and appreciably
longer than digit III, ilium with short triangular anterior process, pubis with
closed obturator foramen with an apron-like distal part, ungual | largest in pes.
Efraasia diagnostica (Huene)
Thecodontosaurus diagnosticus Fraas, 1913: 1098 (nomen nudum).
Palaeosaurus (?) diagnosticus Huene, 1932: 52, 73, figs 1, 2, 7-8, pls 4-6.
Palaeosauriscus diagnosticus Charig, 1967: 712.
Efraasia diagnostica Galton, 1973: 247, figs 1A-E, 2-15, 16A, 17C_-D.
Syntypes
An almost complete skeleton (SMNS 12667) (Berckhemer 1938) together
with additional material (SMNS 12668) from the Stubensandstein (Upper
Triassic) of Pfaffenhofen, Wiirttemberg, West Germany (see Galton 1973).
Diagnosis
As for genus.
Discussion
As shown by the manus (Fig. 71) and the pes (Figs 8D, 11M), Efraasia
diagnostica undoubtedly represents a slender-footed prosauropod. Apart from
that of Anchisaurus polyzelus (YPM 1883, Fig. 12B; Huene 1906: pl. 1), this
is the only reasonably complete and well-preserved skeleton (Fig. 12A; Berck-
hemer 1938) of a slender-footed prosauropod described to date. Efraasia
diagnostica (Figs 71, 8D, 91, M, 10D, J, N, 11M) resembles Anchisaurus capensis
in several features (Table 3) but differs in three important respects, viz. the
ilium has a short triangular anterior process (Fig. 10D), the subacetabular
part of the pubis is deep with a complete obturator foramen (Fig. 10J, N), and
the bases of the neural spines of the anterior caudal vertebrae are narrow
(Fig. 9R).
Genus Thecodontosaurus Riley & Stutchbury, 1836
Diagnosis
At least 21 dentary teeth (in holotype, Fig. 9B); from referred specimens
without teeth diagnosis tentatively expanded as follows: elongate basiptery-
goid processes, cervical vertebrae proportionally short compared with other
anchisaurids, high centra to dorsal vertebrae, narrow base to neural spines of
anterior and caudal vertebrae, high placed deltopectoral crest on proximal
third of humerus, manus with slender metacarpal I and digits II and III subequal
in length, short triangular anterior process to ilium.
ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 139
Thecodontosaurus antiquus Morris
Thecodontosaurus Riley & Stutchbury, 1836: 398; 1840: 352, pl. 29 (figs 1-2).
Thecodontosaurus antiquus Morris, 1843: 211.
Type
Incomplete dentary with teeth from the Magnesian Conglomerate (Upper
Triassic) near Bristol, England.
Diagnosis
As for genus.
Discussion
Riley & Stutchbury (1836, 1840) did not give a specific name for Theco-
dontosaurus and this oversight was rectified by the proposal of 7. antiquus
Morris, 1843.
Thecodontosaurus was the first genus of prosauropod to be described
so it is unfortunate that there is no articulated association between teeth of
the type and the postcranial material referred to the genus by Seeley (1895a)
and Huene (1907-8, 19146). Indeed, the only articulated bones referred to
Thecodontosaurus are a few short sequences of vertebrae (Huene 1907-8:
figs 214, 218-220) and a fore limb with scapula, cervical vertebra and dorsal
ribs (Fig. 11A—G). The description and skeletal reconstruction of Thecodonto-
saurus antiquus given by Huene (1932: 116, pl. 54 (fig. 1)) are based on many
specimens. However, the postcranial remains from Bristol indicate the presence
of a slender-footed prosauropod (Fig. 11A—G) and it is reasonable to refer
this material to Thecodontosaurus antiquus. Species of Thecodontosaurus from
other parts of the world (see next section and pp. 145, 147, 152, 153) are incor-
rectly referred to this genus.
Fig. 8. Comparison of the pes in various prosauropods, either right in dorsal view or left in
ventral view (A, C, E, G, I,), all drawn to digit III unit length, scale lines represent 5 cm. These
genera are divided by Romer (1966) between the families Anchisauridae (B, C, E-I, L, N, Q)
and Plateosauridae (M, O, P, R, S). A. Ammosaurus major, YPM 209. B. Aristosaurus erectus,
from Van Hoepen (1920a). C. Thecodontosaurus browni, from Huene (1932). D. Efraasia
diagnostica, SMNS 12668. E. Anchisaurus polyzelus, YPM 1883. F. Thecodontosaurus antiquus,
metatarsal III, from Huene (1907-08). G. Anchisaurus capensis, SAM-990. H. Lufengosaurus
huenei, figured as Gyposaurus sinensis by Young (1941). 1. Hortalotarsus skirtopodus, figured
as Thecodontosaurus skirtopodus by Huene (1906). J. Ammosaurus major, YPM 208. K. Ammo-
saurus cf. major, from Galton (1971). L. Massospondylus harriesi, figured as M. browni by
Van Hoepen (19205). M. Plateosaurus gracilis, from Berckhemer (1938). N. Massospondylus
harriesi, from Broom (1911). O. Lufengosaurus huenei, figured as Yunnanosaurus magnus by
Young (1947). P. Lufengosaurus huenei, figured as Yunnanosaurus robustus by Young (1951).
Q. Gryponyx africanus, from Broom (1911). R. Plateosaurus robustus, from Huene (1932).
S. Plateosaurus sp., figured as Pachysaurus wetzelianus by Huene (1932).
140 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 9. Comparison of skulls and vertebrae of prosauropods, scale = 2,5 cm, A, B, D-F
x 0,5. A. Skull of Anchisaurus polyzelus, YPM 1883, lateral view. B. Thecodontosaurus anti-
quus, left partial dentary in lateral view, from Riley & Stutchbury (1840). C. Skull of Plateo-
saurus in lateral view, from Romer (1966). D. Massospondylus harriesi, left partial lower jaw
in lateral view, from Haughton (1924). E-G. Braincases in ventral view: E. Anchisaurus
polyzelus, YPM 1883. F. Thecodontosaurus antiquus, YPM 2192. G. Efraasia diagnostica
basisphenoid, SMNS 12667. H-J. Third cervical vertebra in lateral view: H. Anchisaurus
polyzelus, YPM 1883. I. Efraasia diagnostica, SMNS 12667. J. Lufengosaurus huenei, figured
as Gyposaurus sinensis by Young (1941). K—O. dorsal vertebrae (tenth to twelfth) in lateral
view: K. Anchisaurus polyzelus, YPM 1883. L. Anchisaurus capensis, SAM-990. M. Efraasia
diagnostica, SMNS 12667. S. Thecodontosaurus antiquus, from Huene (1907-08).
ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 141
Anchisauridae nomina dubia
Hortalotarsus skirtopodus Seeley, 1894
Hortalotarsus skirtopodus Seeley, 1894: 411, figs 1-3.
Thecodontosaurus skirtopodus Huene, 1906: 44, figs 72-78, pls 13, 14. Haughton, 1924: 370.
Huene, 1932: 117. Haughton & Brink, 1954: 35.
Gyposaurus skirtopodus Charig, 1967: 712.
Type
Incomplete hind limb in the Albany Museum, from the Cave Sandstone,
Stormberg Series, of Barkly East Division, Cape Province, South Africa.
Discussion
The assignment of Hortalotarsus skirtopodus to Thecodontosaurus by
Huene (1906) was based in part on the characters of isolated bones found at
localities different from that of the type specimen. The pes of the type (Fig. 81)
is obviously that of a slender-footed prosauropod, and Huene (1906) could
not distinguish it from the pes of Thecodontosaurus. However, this pes (Fig. 81)
is also indistinguishable from those of Anchisaurus capensis (Fig. 11K), Anchi-
saurus polyzelus (Fig. 11L) and Efraasia diagnostica (Fig. 11M). This specimen
is generically and specifically indeterminate so Hortalotarsus skirtopodus
Seeley is a nomen dubium.
Thecodontosaurus browni (Seeley, 1895b)
Massospondylus browni Seeley, 18956: 118, figs 13-14.
Thecodontosaurus browni Huene, 1906: 141, pl. 12 (figs 7-8); 1932: 118. Broom, 1911: 293.
Haughton, 1924: 370.
Type
Limb bones (BMNH R3302) from the Red Beds, Stormberg Series of Telle
River, Herschel, Cape Province, South Africa.
Discussion
Seeley (18955) noted that the proportions of the phalanges of the pes are
very similar to those of Hortalotarsus (Fig. 8H) so this is probably another
generically and specifically indeterminate specimen of a_ slender-footed
prosauropod.
Thecodontosaurus minor Haughton, 1918
Thecodontosaurus minor Haughton, 1918: 468; 1924: 376, fig. 21.
Thecodontosaurus browni: Huene, 1932: 118.
Type
Left tibia, a cervical vertebra and a portion of a left ilium (SAM-3451)
from the Red Beds, Stormberg Series, from road-cutting at Naude’s Nek,
Pitsing, Maclear District, Cape Province, South Africa.
Discussion
The tibia is slender so this may be a slender-footed prosauropod but, on
142 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 10. Comparison of ilium and pubis of prosauropods. Scale = 2,5 cm. A—F. Lateral view
of left ilium: A. Anchisaurus polyzelus, YPM 1883. B. Anchisaurus capensis, SAM-990. C.
Lufengosaurus huenei, figured as Gyposaurus sinensis by Young (1941). D. Efraasia diagnostica,
SMNS 12667. E. Aristosaurus erectus, fromVan Hoepen (1920a). F. Thecodontosaurus antiquus,
from Huene (1907-08). G—J. lateral view of left pubis: G. as A. H as B. Las C. Jas D. K-N.
Ventral view of right pubis (L, M) or dorsal view of left pubis (K, N): K as A, G. Las C, I.
M as E. N as D, J.
the basis of available material, this specimen is a generically and specifically
indeterminate prosauropod and Thecodontosaurus minor is a nomen dubium.
Thecodontosaurus macgilivrayi (Seeley)
Agrosaurus macgilivrayi Seeley, 1891: 161, figs 1-6.
Thecodontosaurus macgilivrayi Huene, 1906: 147, figs 86-90; 1932: 52.
Type
Tibiae, radius, an ungual and tooth (BMNH 49984) from York Peninsula,
Queensland, Australia.
Discussion
Huene (1932: 52) subsequently referred Agrosaurus macgilivrayi to the
Theropoda (Coelurosauria) as do Romer (1956, 1966), Steel (1970) and White
(1973). However, the latero-distal surface of the tibia is notched (Fig. 11H—J)
(Huene 1906: fig. 86a, d-e) to receive the central ascending process of the
astragalus in typical prosauropod fashion. This material undoubtedly represents
ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 143
a prosauropod but is generically and specifically indeterminate, so Agrosaurus
macgilivrayi is a nomen dubium; it may represent a slender-footed prosauropod.
PROSAUROPOD SPECIES INCORRECTLY ASSIGNED TO ANCHISAURIDAE
Family Plateosauridae Marsh, 1895
Diagnosis
Larger forms, skull massively built, deep posterior half to lower jaw with
articulation offset ventral to line of tooth row, manus and pes broad.
Ammosaurus major Marsh, 1889
Anchisaurus major Marsh, 1889: 331, fig. 1.
Ammosaurus major Marsh, 1891: 267; 1892: 545, pl. 16 (fig. 4), pl. 17 (fig. 3); 1896: 150,
pl. 3 (figs 3, 6). Huene, 1906: 15, pls 5-9; 1907-08: 303-04, figs 297-298; 1914a: 13;
19145: 74, figs 20-22; 1932: 26. Lull, 1915: 148, figs 24-25; 1953: 123, figs 19-20. Galton,
1971: 786, figs-9, 11A.
Anchisaurus solus Marsh, 1892: 545; 1896: 149. Huene, 19146: 72, figs 12-19. Lull, 1915:
144, figs 22-23; 1953: 120.
Anchisaurus (?) solus Huene, 1906: 14, pl. 4.
Ammosaurus solus Huene, 1932: 27, pl. 49 (fig. 1).
Type
Pelvis and hind limbs (YPM 208) from the Portland Beds, upper part of
Newark Series near Manchester, Connecticut, U.S.A. (Ammosaurus major).
Almost complete skeleton (YPM 209) from the same locality and horizon
(Anchisaurus solus).
Discussion
Ammosaurus has long been regarded as a primitive theropod dinosaur
but it is considered as an anchisaurid by Steel (1970) and by Galton (1971),
who provides a detailed discussion of the taxonomic position of this genus
(Galton in press). The pes of the holotype (Fig. 8J) and of the referred specimens
(Fig. 8A, K) plus a referred manus (Fig. 7E) are of the broad type. Ammo-
Saurus is a broad-footed prosauropod characterized by the following combina-
tion of characters: centra of dorsal vertebrae low, slender sacral rib 3, elongate
anterior process to ilium, subacetabular part of the ischium emarginated
ventrally (Galton, in press).
Aristosaurus erectus van Hoepen, 1920a
Aristosaurus erectus van Hoepen, 1920a: 82, figs 1-6, pls 9-10. Haughton 1924: 379. Haughton
& Brink, 1954: 33.
Gyposaurus capensis Huene, 1932: 123, pl. 54 (fig. 2).
Gyposaurus erectus Charig, 1967: 712. Steel, 1970: 49.
Type
An almost complete skeleton as slab and counterpart (TM 130) from the
Cave Sandstone, Stormberg Series, near Roosendal, Senekal District, Orange
Free State, South Africa.
144 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 11. Thecodontosaurus, Agrosaurus and comparisons of the anchisaurid pes. Scale = 5 cm,
A-J = x 0,45. A-G. Thecodontosaurus antiquus, YPM 2195, from Durdham Down, Bristol,
England. Matrix indicated by stipple, broken bone by diagonal shading: A. Right side of
anterior cervical vertebra. B. Right scapula in lateral view. C. Left humerus in medial view.
D. As C, in anterior view. E. Left ulna in proximal, lateral and distal views. F. Left radius
in lateral view. G. Left manus in lateral or dorsal view. H-J. Agrosaurus macgillivrayi, distal
end of left tibia in lateral view (H), anterior view (1), and distal view (J), all from Huene (1906).
K-M. Anchisaurid pes, drawn to digit III unit length: K. Anchisaurus capensis, SAM-990,
compare with Fig. 5. L. Anchisaurus polyzelus, YPM 1883. M. Efraasia diagnostica,
SMNS 12668.
ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 145
Discussion
Romer (1956, 1966), Charig (1967), Steel (1970) and White (1973) follow
Huene (1932, 1956) in regarding Arisftosaurus as a junior synonym for Gypo-
saurus but this is unlikely because, in contrast to the situation in Gyposaurus
capensis (Table 3):
1. In photographs of the skeleton of Aristosaurus erectus in Van Hoepen
(1920a: pls 9-10) and in the reconstruction (Fig. 12D) given by Huene
(1932) the dorsal vertebrae are proportionally higher with a central
length to height ratio of about 1,4 for dorsal 10.
2. The anterior process of the ilium is short (Fig. 10E).
3. The obturator foramen of the pubis is closed ventrally (Fig 10M).
4. The hind feet of Aristosaurus erectus appear to be of the broad type.
There is no reason why Aristosaurus erectus should be referred to the
genus Anchisaurus. Aristosaurus erectus appears to be a valid taxon of broad-
footed prosauropod, but further preparation and illustration of the holotype
is needed.
Genus Massospondylus Owen, 1854
(includes Leptospondylus Owen, Pachyspondylus Owen, Aetonyx Broom,
Gryponyx Broom, Dromicosaurus Van Hoepen)
Massospondylus carinatus Owen, 1854
Massospondylus carinatus Owen, 1854: 97. Seeley, 18955: 102, figs 1-12. Huene, 1906: 36,
figs 43-70, pls 13-16. Broom, 1911: 241. Haughton, 1924: 383. Huene, 1932: 124.
Leptospondylus capensis Owen, 1854: 97.
Pachyspondylus orpenii Owen, 1854: 97.
Type
Isolated bones from the Red Beds, Stormberg Series of Beaucherf, Harri-
smith, Orange Free State, South Africa. The holotype in the Museum of the
Royal College of Surgeons in London was destroyed during World War II
but casts of this material are in the National Museum of Southern Rhodesia,
Bulawayo (J. Attridge, pers. comm.).
Massospondylus harriesi Broom, 1911
Massospondylus harriesi Broom, 1911: 299, pls 15-17. Haughton, 1924: 384, figs 21-29.
Huene, 1932: 125.
Massospondylus browni (non Seeley, 18956): Van Hoepen 1920h: 118, pls 17-22.
Aetonyx palustris Broom, 1911; 304, figs 20-23. Haughton, 1924: 404, fig. 30. Huene, 1932: 91.
Gryponyx africanus Broom, 1911: 294, figs 1-9. Haughton, 1924: 417, figs 36-38. Huene,
1932: 88, pl. 7 (figs 1-4).
Gryponyx taylori Haughton, 1924: 420, fig. 39. Huene, 1932: 90.
Dromicosaurus gracilis Van Hoepen, 1920b: 103, figs 8-21, pls 13-16. Haughton, 1924: 405.
Thecodontosaurus dubius Haughton, 1924: 377.
146 ANNALS OF THE SOUTH AFRICAN MUSEUM
Types
Massospondylus harriesi: Bones of the fore limb (SAM-3394) from the top
of the Red Beds, Stormberg Series of Foutanie, Fouriesburg, Orange Free
State, South Africa.
Aetonyx palustris: Partial skeleton (SAM-2768, 2769, 2770) from the top
of the Red Beds (but listed as from Cave Sandstone by C. E. Gow in Anderson
& Anderson 1970), Stormberg Series from Foutanie, Fouriesburg, Orange
Free State, South Africa.
Gryponyx africanus: Pelvis and hind limb, right and left manus, vertebrae
(SAM-3357-9) from the top of the Red Beds (but listed as from the Cave
Sandstone by C. E. Gow in Anderson & Anderson 1970), Stormberg Series of
Foutanie, Fouriesburg, Orange Free State, South Africa.
Gryponyx taylori: Pelvic girdle and sacral vertebrae (SAM-3453) from the
top of the Red Beds (but listed as from Cave Sandstone by C. E. Gow in Ander-
son & Anderson 1970), Stormberg Series of Fouriesburg, Orange Free State,
South Africa.
Dromicosaurus gracilis: Partial skeleton (TM 123) from Red Beds of
Naaupoort Nek, Bethlehem, Orange Free State, South Africa.
Thecodontosaurus dubius: Larger portion of a skeleton (SAM-3712) from
the Cave Sandstone, Stormberg Series of Ladybrand, Orange Free State,
South Africa.
Discussion
On the basis of the phalanges Seeley (18955) stated that Massospondylus
carinatus had a broad hind foot and Huene (1906) separated this genus from
Plateosaurus mainly because of its Thecodontosaurus-like tibia. Massospondylus
was the first genus of broad-footed prosauropod to be described from South
Africa.
The manus (Fig. 7F—-G, J) and the pes (Fig. 8L, N) of Massospondylus
harriesi (SAM-3394) are obviously of the broad type. The material (Figs 7G,
8L, 12E) described by Van Hoepen (19205) as Massospondylus browni should
be referred to this species (Haughton 1924). The manus of Aetonyx palustris
Broom (Fig. 7O) is of the broad type and the pes is similar, as indicated by
the measurements given by Huene (1932: 92). J. Attridge (pers. comm.) regards
Aetonyx palustris as a junior synonym for Massospondylus harriesi. The manus
(Fig. 7Q) and pes (Fig. 8Q) of Gryponyx africanus are of the broad type and
Gryponyx africanus is probably a junior synonym for Massospondylus harriesi.
Gryponyx taylori Haughton is a nomen dubium because the material is
generically and specifically indeterminate; it probably represents another
specimen of Massospondylus harriesi.
The manus and pes of Dromicosaurus gracilis Van Hoepen are not pre-
served but were probably of the broad type because Dromicosaurus was regarded
as being closely allied to Aetonyx and Massospondylus by Van Hoepen (19206),
Haughton (1924) and Huene (1932). J. Attridge (pers. comm.) regards Dromico-
ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 147
saurus gracilis as a junior synonym for Massospondylus harriesi. The type
specimen of Thecodontosaurus dubius Haughton has never been figured, but
Huene (1932: 92) referred it to Aetonyx palustris so it is presumably a broad-
footed form; J. Attridge (pers. comm.) refers this specimen to Massospondylus
harriesi.
Stratigraphically and geographically, Massospondylus harriesi is the most
ubiquitous prosauropod in southern Africa.
Plateosaurus gracilis (Huene, 1907)
Thecodontosaurus (?) hermannianus Huene, 1907-08: 216, fig. 236, pl. 144 (fig. 1).
Plateosaurus gracilis Huene, 1932: 303.
Type
Right maxilla with teeth from the Stubensandstein (Upper Triassic) of
Heslach, in Stuttgart, West Germany.
Lufengosaurus huenei Young, 1941la
Lufengosaurus hueni Young, 1941a: 1, figs 1-25, pls 1-6; 1947: 41; 1951: 50, fig. 11, pl. 12.
Rozhdestvensky, 1965.
Gyposaurus sinensis Young, 1941b: 205, pls 1-9; 1948: 91, pls 1-5; 1951: 49.
Yunnanosaurus huangi Young, 1942: 64, figs 1-17; 1951: 56.
Lufengosaurus magnus Young, 1947: 2, figs 1-14.
Yunnanosaurus robustus Young, 1951: 58, figs 12-14, pls 7-10.
Types
Several incomplete skeletons from the lower Lufeng Series (Upper Triassic,
Rhaetic) of Lufeng, Yunnan, China.
Discussion
Rozhdestvensky (1966) restudied 70 specimens from the Lufeng Series
and decided that Lufengosaurus huenei, L. magnus, Yunnanosaurus huangi,
Y. robustus and Gyposaurus sinensis of Young were all differentiated only on
size-related characters and are conspecific (as Lufengosaurus huenei Young,
1941a). Rozhdestvensky (1966) noted that, judging from the original diagnosis
of Young (19416), Gyposaurus sinensis is hardly distinguishable from G. capensis,
but that without visual comparisons or more detailed descriptions it is impos-
sible to decide the relationship between these two species. Galton (1973, in
press) noted that ‘“Gyposaurus’ sinensis was incorrectly referred to the genus
Gyposaurus, and may represent a new genus or be based on juveniles of either
Lufengosaurus or Yunnanosaurus. ‘Gyposaurus’ sinensis should not be referred
to the genus Gyposaurus (or Anchisaurus) because of the following anatomical
features (Table 3):
1. Centra of posterior dorsal vertebrae are proportionally high with a
central length to height ratio of 1,1 (Fig. 90).
2. Anterior process of ilium is short and triangular (Fig. 10C).
3. Subacetabular part of pubis is broken but originally this region was
148 ANNALS OF THE SOUTH AFRICAN MUSEUM
re
- eS
- <
=
47% =
Fig. 12. Skeletal reconstructions of prosauropods. Scale = 20 cm. Tails of A and B diagram-
matically folded over. A. Efraasia diagnostica, SMNS 12667, 12668, from Galton (1973).
B. Anchisaurus polyzelus, YPM 1883, AM 41/109, from Galton (1973). C. Lufengosaurus huenei,
based on figures of individual bones given by Young (19415) as Gyposaurus sinensis.
ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 149
x
Lys 94
by lee,
SS
YY
LL
Fig. 12. (cont.)
D. Aristosaurus erectus, modified from Gyposaurus capensis of Huene (1932). E. Massospondylus
harriesi, modified from Thecodontosaurus browni of Huene (1932), based on specimen described
as Massospondylus browni by Van Hoepen (1920a).
deep with a complete obturator foramen (Young 1941b: 222; 1948: 96)
(Figs 10I, L).
4. Pes (Fig. 8H) and manus (Fig. 7B) are broad.
Young (1941b) did not cite the papers of Broom (1906, 1911) so the assign-
ment as Gyposaurus sinensis was probably based on the skeletal reconstruction
of Aristosaurus erectus given by Huene (1932) as Gyposaurus capensis. However,
the skeleton of ‘Gyposaurus’ sinensis (Fig. 12C) differs greatly in several aspects
from that of Aristosaurus erectus (Fig. 12D) and this is especially true for the
form of the neck vertebrae (Fig. 9J) and fore limb. It should be noted that the
skeletal reconstruction of ‘Gyposaurus’ sinensis given by Young (19418, pl. 9)
bears practically no resemblance to a reconstruction (Fig. 12C) based on
figures of the bones of the same specimen given by Young (19410). The correct-
ness of the contention of Rozhdestvensky (1966) that Gyposaurus sinensis
150 ANNALS OF THE SOUTH AFRICAN MUSEUM
Young, 19415 is a junior synonym of Lufengosaurus huenei Young, 1941a is
shown by the proportionally short neck and proportionally massive manus
of the two types.
Prosauropoda nomina dubia
Gryponyx transvaalensis Broom, 1912
Gryponyx transvaalensis Broom, 1912: 82, figs 3-4. Van Hoepen, 1920b: 102. Haughton,
1924: 420. Huene, 1932: 91, pl. 7, fig. 5. j
Type
Ungual | of the manus and a metatarsal (in the Transvaal Museum) from
the Bushveld Sandstone (Cave Sandstone), Stormberg Series of Wiepe 1258,
northern Transvaal, South Africa.
Discussion
This material is probably prosauropod but is generically and specifically
indeterminate.
NON-PROSAUROPOD SPECIES INCORRECTLY ASSIGNED TO ANCHISAURIDAE
Order SAURISCHIA
Suborder THEROPODA
Arctosaurus osborni Adams, 1875
Arctosaurus osborni Adams, 1875: 177. Lydekker, 1889: 352.
Type
Isolated cervical vertebra (NMI 62 1971) from Heiberg Formation (Upper
Triassic) of north-west extremity of Cameron Island, Bathurst Group, Arctic
Archipelago, Canada.
Discussion
Arctosaurus was described as reptilian by Adams (1875) but subsequently
Lydekker (1889) referred it to the family Anchisauridae. Regarded as a turtle
by Huene (1906) and White (1973) but referred to the prosauropod family
Melanorosauridae (as Plateosauravidae) by Huene (1956) and to the Anchi-
sauridae (as Thecodontosauridae) by Romer (1966). The region of the dia-
pophysis is slightly damaged (Fig. 13B, F) but from the adjacent curves of the
neural arch (Fig. 13E) it is obvious that the diapophysis was very small and,
as a result, this vertebra is from the anterior part of the series and is probably
either the third or fourth cervical vertebra. It is proportionally very much
shorter than the equivalent vertebrae of Anchisaurus (Figs 9H, 12B), Efraasia
(Figs 91, 12A) and Plateosaurus (Huene 1926). The only prosauropods with
cervical vertebrae proportionally as short are Thecodontosaurus (Fig 11A) and
Lufengosaurus (Figs 9J, 12C) but in both cases the vertebrae are proportionally
much lower, the neural spines are not so well developed and there is no pleuro-
coel (exaggerated in Arctosaurus because of crushing). Arctosaurus osborni is
ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 15]
|
|
|
Fig. 13. Arctosaurus osborni Adams, holotype NMI G2 1971, anterior cervical vertebra in:
A. Anterior view. B. Left lateral view. C. Posterior view. D. Ventral view. E. Dorsal view.
F. Stereo photograph of left side, compare with B. Scale = 2,5 cm. Broken bone indicated
by diagonal shading.
not a prosauropod and, on the basis of the general form-of the vertebra and
the presumed presence of a pleurocoel, this specimen is tentatively regarded as
Theropoda incertae sedis as listed by Steel (1970).
Ischisaurus cattoi Reig, 1963
Ischisaurus cattoi Reig, 1963: 10, figs 4B, 5. Colbert, 1970: 27. Bonaparte, 19724: 673, fig. 22.
Type
Two incomplete skeletons from the Ischigualasto Formation (lower Upper
Triassic) of Argentina.
Discussion
Ischisaurus is listed as an anchisaurid by Steel (1970) and as Saurischia
incertae by Bonaparte (1972b: 674), who notes that ‘the suggested affinities
152 ANNALS OF THE SOUTH AFRICAN MUSEUM
with the Coelurosauria appears as one of the possible relationships’. Romer
(1966) and Colbert (1970) list /schisaurus as a coelurosaurian theropod.
Order THECODONTIA
‘Thecodontosaurus’ gibbidens Cope, 1878
Thecodontosaurus gibbidens Cope, 1878: 177. Huene, 1921: 571, figs 14-15.
Type
Isolated teeth from the Upper Triassic of Pennsylvania, U.S.A.
Discussion
The isolated teeth are almost circular rather than oval in cross-section as
in prosauropods (Thecodontosaurus antiquus Riley & Stutchbury, 1840; Anchi-
saurus, YPM 1883; Plateosaurus. AMNH 6810), so these teeth are provisionally
referred to the Ornithischia (Galton, in press).
Spondylosoma absconditum Huene, 1935
Spondylosoma absconditum Huene, 1935: 247, pl. 30, figs 1-13. Charig, 1967: 712. Colbert,
1970: 19. Bonaparte, 19725: 674.
Type
Scapula, humerus, femur, tibia (all incomplete) and eight vertebrae from
the Santa Maria Formation (Upper Triassic) of Brazil.
Discussion
Huene (1935, 1942) regarded Spondylosoma as a saurischian but did not
make a more specific assignment for this genus. Romer (1956, 1966) referred
Spondylosoma to the Anchisauridae (as Thecodontosauridae) as did Colbert
(1970) and Charig (1967), who noted at the same time the possibility of its
being a prestosuchid pseudosuchian. Bonaparte (19725: 674) notes that ‘there
are doubts regarding its assignment to the Saurischia, or even to Prosauropoda.
Unfortunately there are not sufficient diagnostic pieces to define better its
taxonomic position’ and Spondylosoma is listed as Saurischia incertae.
Teleocrater alphos (Haughton, 1932)
Thecodontosaurus (?) alphos Haughton, 1932: 662, fig. 19.
Teleocrater alphos Charig, 1967: 712.
Type
Two cervical vertebrae (SAM-10654) from the Manda Formation (Upper
Triassic, Anisian) of Tanzania.
Discussion
Charig refers this material to the pseudosuchian Teleocrater.
ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 153
Subclass LEPIDOSAURIA
Order LACERTILIA
Huene (1932, not 1931 as cited by Steel 1970) considered that the follow-
ing Middle Triassic (Muschelkalk) species were based on specimens which
should be referred to the eosuchian Tanystropheus, a lacertilian according to
Wild (1974); the non-prosauropod nature of these species is also discussed by
Colbert (1970):
Tanystropheus primus (Huene, 1907-08)
Thecodontosaurus primus Huene, 1907-08: pl. 42 (figs 8-9).
Tanystropheus primus Huene, 1932: 6.
Tanystropheus latespinatus (Huene, 1907-08)
Thecodontosaurus (?) latespinatus Huene, 1907-08: figs 237-245.
Tanystropheus latespinatus Huene, 1932: 6.
NOTES ON THE FAMILIES ANCHISAURIDAE AND PLATEOSAURIDAE
The infra-order Prosauropoda is currently divided into three families:
Anchisauridae (= Thecodontosauridae), Plateosauridae and Melanorosauridae
(see Romer 1956; Colbert 1964; Charig et a/. 1965; Bonaparte 1972a). Post-
cranially the separation is clearest between melanorosaurids and non-
melanorosaurids (Romer 1956: 617; Bonaparte 1972a: 160). Galton (1971,
1973) suggests that the range of morphological variation is insufficient to
warrant the retention of two families of non-melanorosaurid prosauropods.
However, the skulls of Anchisaurus (Fig. 9A) and Plateosaurus (Fig. 9C) are
very different and, because of this, Galton (in press) now considers that they
should not be included in the same family. Fortunately the genera concerned
are the basis for the first two valid prosauropod family names to be proposed:
Anchisauridae Marsh, 1885, and Plateosauridae Marsh, 1895.
In only one case (Anchisaurus polyzelus, YPM 1883) is a well-preserved
skull found in natural association with a skeleton of a slender-footed pro-
sauropod. Consequently the referral of Efraasia and Thecodontosaurus to the
Family Anchisauridae is tentative. Contrary to the impression given by Huene
(1932: fig. 7; 1956: fig. 10), the skull of Efraasia is very incomplete but, as
noted by Galton (1973), Efraasia is an ideal ancestor for the more recent
Anchisaurus. The holotype of Thecodontosaurus antiquus is an incomplete
dentary but, judging from what is preserved (Fig. 9B), the complete lower
jaw was probably more like that of Anchisaurus (Fig. 9A) than that of Plateo-
Saurus (Fig. 9C). Although considered unlikely, the discovery of additional
material may show that the restriction of the family Anchisauridae to slender-
footed forms is artificial. However, the criterion is practical and with it most
taxa and specimens of non-melanorosaurid prosauropods are readily referable
to either the Family Anchisauridae or the Family Plateosauridae.
Plateosaurus engelhardti Meyer, 1837 from the Keuper (Upper Triassic)
154 ANNALS OF THE SOUTH AFRICAN MUSEUM
of Germany is the earliest taxon of plateosaurid to be described, but since
then a multitude of taxa have been erected for material from the Triassic of
Germany (Steel (1970) lists 9 species as Plateosaurus, 8 species as Gresslyo-
saurus). However, this material is probably extremely overclassified and all
the taxa of European plateosaurids listed by Steel (1970: 53-56) should be
provisionally regarded as junior synonyms of Plateosaurus engelhardti Meyer,
1837. All the European plateosaurid material can probably be referred to (at
the most) three species of Plateosaurus, but a restudy of all the holotypes is
needed to determine the other valid species.
Lufengosaurus (see Young 194la, b, 1942, 1947, 1951) and Plateosaurus
(see Huene 1907-8, 1926, 1932) possess the features listed above as charac-
teristic of plateosaurids. The skulls of Ammosaurus (see Galton, in press) and
Aristosaurus (see Van Hoepen 1920a) are not well enough preserved to tell
anything about the form of the skull. However, the holotype of Massospondylus
harriesi includes a lower jaw (Fig. 9D), the posterior part of which is deep
with the articulation offset ventral to the line of the tooth row. J. Attridge is
studying two skulls of Massospondylus harriesi (SAM- K388 and K1314) and
has found that both skulls show the features listed above (pers. comm.).
Massospondylus is the most ubiquitous prosauropod in southern Africa
and its previous classification as an anchisaurid made the prosauropod fauna
of Africa unique, because in other areas with abundant, well-preserved pro-
sauropod skeletons, plateosaurids are the most common form. However,
Cox (1973: 213) notes that ‘it is clear that land connections between all the
continents existed for much, at least, of the Triassic’. As regards prosauropods
the presence of Anchisaurus in North America and South Africa and the presence
of Plateosaurus in Germany and South America (Casamiquela 1964; Bonaparte
19725) indicated that this was the case for the continents on either side of the
Atlantic. With the transfer of broad-footed forms, previously listed under the
Anchisauridae, to the Plateosauridae, this family becomes the dominant and
cosmopolitan prosauropod family of the world. In marked contrast, the Anchi-
sauridae have an extremely restricted fossil record (total of about 10 articulated
specimens for North America, Europe and South Africa) with no remains
discovered to date from Asia (Young 1951; Rozhdestvensky 1966) or South
America (Bonaparte, pers. comm.). Haughton (1924) noted that the Stormberg
Series of South Africa was deposited under conditions of progressively increasing
aridity, and it is interesting that skeletal remains of melanorosaurids occur in
the lowermost levels (Passage Beds, Charig et a/. 1965; basal Red Beds,
Haughton 1924), most plateosaurid skeletons occur higher in the Red Beds,
and those of anchisaurids are found in the overlying Cave Sandstone (Charig
et al. 1965; Haughton 1924; Haughton & Brink 1956). Charig et al. (1965)
report the presence of small tridactyl footprints in the Passage Beds and in the
lower Red Beds and note (p. 204) that these ‘*. . . may indicate the movement
of thecodontosaurids from one upland region to another via a lowland area’.
The world-wide rarity of anchisaurid skeletal remains is presumably because
ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 155
these species occupied the drier upland areas, which are rarely represented in
the fossil record.
SUMMARY
The holotype (SAM-990) of the prosauropod dinosaur Anchisaurus
capensis (Broom, 1911) is characterized by the following combination of
characters: low centra to posterior dorsal vertebrae, broad bases to neural
spines of anterior caudal vertebrae, ilium with long anterior process, pubis
with open obturator foramen, and with slender pes with first ungual phalanx
the largest. SAM-990 differs from the North American Anchisaurus polyzelus
(Hitchcock) only in the large size of the first ungual of the pes, and SAM-990
is referred to Anchisaurus Marsh, 1885 as Anchisaurus capensis (Broom). Taxa
which have been incorrectly referred to the genus Gyposaurus (= Anchisaurus)
are Gyposaurus erectus (Van Hoepen) (= Aristosaurus erectus Van Hoepen),
Gyposaurus sinensis Young (= Lufengosaurus huenei Young) and Gyposaurus
skirtopodus (Seeley) (= Hortalotarsus skirtopodus Seeley, nomen dubium). The
Family Anchisauridae is restricted to those species with slender feet, viz. Anchi-
saurus polyzelus, A. capensis, Efraasia diagnostica, and Thecodontosaurus
antiquus. Taxa with broad feet previously classified as anchisaurids (Aristo-
saurus, Ammosaurus, Gyposaurus sinensis (as Lufengosaurus huenei), Masso-
spondylus (including Aetonyx, Dromicosaurus, Gryponyx africanus, G. taylori,
Thecodontosaurus dubius), Yunnanosaurus (= Lufengosaurus) are transferred to
the family Plateosauridae, the dominant and cosmopolitan family of
prosauropods.
ACKNOWLEDGEMENTS
P. M. Galton is grateful to the following for the loan or use of specimens
from their respective institutions, listed in alphabetical order: Drs W. Coombs,
Jr and A. E. Wood, Amherst College Museum, Amherst, Massachusetts;
Drs E. H. Colbert and E. S. Gaffney, American Museum of Natural History,
New York; Dr A. J. Charig, British Museum (Natural History), London;
C. E. O’Riordan, National Museum of Ireland, Dublin; Dr K. D. Adam,
Staatlichen Museum fiir Naturkunde in Stuttgart; Drs J. H. Ostrom and E. L.
Simons, Peabody Museum, Yale University, New Haven, Connecticut. Galton
thanks Mr J. Attridge, Birkbeck College, University of London; Dr D. Baird,
Princeton University, New Jersey and Dr J. Bonaparte, National University
of Tucuman, Argentina for helpful discussions and personal communications
as cited above, and Drs J. H. Ostrom and E. L. Simons of Yale University
for the continued use of the research facilities of the Division of Vertebrate
Paleontology of the Peabody Museum of Natural History. The photographs
were taken by Mr N. J. Eden, Department of Palaeontology, South African
Museum (Figs 1, 5,6) and Mr Alan Coleman, Yale Peabody Museum (Fig. 13F).
156 ANNALS OF THE SOUTH AFRICAN MUSEUM
Support for this research was provided to the senior author by the Connecticut
Research Commission (Grant no. RSA 680) while at Yale University and by
a Faculty Research Grant from the University of Bridgeport.
REFERENCES
ApaMs, A. L. 1875. On a fossil saurian vertebra, Arctosaurus osborni, from the Arctic regions.
— Proc. R. Ir. Acad. 2: 177-179.
ANDERSON, H. M. & ANDERSON, J. M. 1970. A preliminary review of the biostratigraphy of
uppermost Permian, Triassic and lowermost Jurassic of Gondwanaland.— Palaeont. afr.
13 (Suppl.): 1-22.
BERCKHEMER, F. 1938. Wirbeltierfunde aus dem Stubensandstein des Strombergs.— Naturwiss.
Monatsh Stuttgart 51: 188-198.
BONAPARTE, J. 1972a. Los tetrapodos del sector superior de la Formacion Los Colorados,
La Rioja, Argentina. (Triasico Superior) I Parte.—Op lilloana 22: 1-183
BONAPARTE, J. 19725. Annotated list of the South American Triassic tetrapods. Proceedings,
Second Gondwana Symposium. I.U.G.S. Commission on Stratigraphy, South Africa, 1970
665-682.
Bonp, G. 1955. A note on dinosaur remains from the Forest Sandstone (Upper Karroo).—
Occ. Pap. Rhod. Mus. 20: 795-800.
Broom, R. 1906. On the South African dinosaur (Hortalotarsus).— Trans. S. Afr. phil. Soc.
16: 201-206.
Broom, R. 1911. On the dinosaurs of the Stormberg, South Africa.— Ann. S. Afr. Mus. 7:
291-308.
Broom, R. 1912. On the remains of the theropodous dinosaur from the Northern Transvaal.—
Trans. geol. Soc. S. Afr. 14: 82-83.
CASAMIQUELA, R. M. 1964. Sobre el hallazgo de dinosaurios tridsicos en la Provincia de Santa
Cruz. Argent. austral. 35: 10-11.
CuariGc, A. J. 1967. Archosauria. In: The fossil record. A symposium with documentation:
708-718. London: Geological Society of London.
CuariG, A. J., ATTRIDGE, J. & CRoMPTON, A. W. 1965. On the origin of the sauropods and
the classification of the Saurischia.— Proc. Linn. Soc. Lond. 176: 197-221.
CoLsBerT, E. H. 1964. Relationships of saurischian dinosaurs.— Am. Mus. Novit. (2181): 1-24.
CoLperT, E. H. 1970. A saurischian dinosaur from the Triassic of Brazil.—Am. Mus. Novit.
(2205): 1-39.
Core, E. D. 1870. Synopsis of the extinct Batrachia, Reptilia and Aves of North America.—
Trans. Am. phil. Soc. 14: 1-252.
Cope, E. D. 1878. On some saurians found in the Triassic of Pennsylvania, by C. M. Wheatley.
—Proc. Am. phil. Soc. 17: 177.
Cox, C. B. 1973. Triassic tetrapods.—Jn: HALLAM, A. ed. Atlas of biogeography: 213-223.
London: Elsevier Scientific Publishing Company.
Fraas, E. 1913. Die neusten Dinosaurierfunde der Schwabischen Trias.— Naturwissen-
schaften (1) 45: 1097-1100.
GALTON, P. M. 1971. The prosauropod dinosaur Ammosaurus, the crocodile Protosuchus,
and their bearing on the age of the Navajo Sandstone of northeastern Arizona.—J.
Paleont. 45: 781-795.
GALTON, P. M. 1973. On the anatomy and relationships of Efraasia diagnostica (Huene)
n. gen., a prosauropod dinosaur (Reptilia: Saurischia) from the upper Triassic of Ger-
many.— Paldont. Z. 47: 229-255.
GALTON, P. M. 1974. The ornithischian dinosaur Hypsilophodon from the Wealden of the
Isle of Wight.— Bull. Br. Mus. nat. Hist. (Geol.) 25: 1-125c.
GALTON, P. M. (in press). Prosauropod dinosaurs (Reptilia: Saurischia) of North America.—
Postilla.
Georrroy, E. F. S.-H. 1833. Divers mémoires sur les grands sauriens trouvés a l’etat fossile
vers les confins maritimes de la Basse Normandie, attribués d’abord au crocodile, puis
determines sous les noms de Teleosaurus et Steneosaurus. Mém. Acad. Sci. Inst. Fr. 12:
3-138.
ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 157
Gitmore, C. W. 1920. Osteology of the carnivorous Dinosauria in the United States National
Museum, with special reference to the genera Antrodemus (Allosaurus) and Ceratosaurus.
— Bull. U.S. natn. Mus. 110: 1-159.
HauGutTon, S. H. 1918. A new dinosaur from the Stormberg Beds of South Africa.— Ann.
Mag. nat. Hist. 2: 468-469.
HauGuHTon, S. H. 1924. The fauna and stratigraphy of the Stormberg series.— Ann. S. Afr.
Mus. 12: 323-497.
HauGutTon, S. H. 1932. On a collection of Karroo vertebrates from Tanganyika Territory.
Q. Jl. geol. Soc. Lond. 88: 634-671.
HaAuGHTON, S. H. & BRINK, A. S. 1954. A bibliographic list of Reptilia from the Karroo Beds
of Africa.—Palaeont. afr. 2: 1-187.
Hitcucock, E. 1865. A supplement to the ichnology of New England. Boston: Wright and Potter.
Huene, F. von. 1906. Uber die Dinosaurier der aussereuropaischen Trias.— Geol. paléont.
Abh. (N.F.) 8: 99-156.
HueEne, F. von. 1907-1908. Die Dinosaurier der Europdischen Triasformation mit Beruck-
sichtigung der aussereuropdischen Vorkomnisse.— Geol. paldont. Abh. (Suppl.) 1: 1-419.
Huene, F. von. 1914a. Saurischia et Ornithischia Triadica (‘Dinosauria’ Triadica). In: Fos-
silium Cat. (Animalia) 4: 1-21.
HueENE, F. von. 19145. Nachtrage zu meinen fruheren Beschreibungen triassischer Saurischia.
—Geol. paldont. Abh. (N.F.) 13: 69-82.
HUuENE, F. von. 1921. Reptilian and stegocephalian remains from the Triassic of Pennsylvania
in the Cope Collection.— Bull. Am. Mus. nat. Hist. 44: 561-574.
HUuENE, F. von. 1926. Vollstandige Osteologie seines Plateosauriden aus dem Schwabischen
Keuper.— Geol. paldont. Abh. (N.F.) 15: 134-180.
HUuENE, F. von. 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte.
— Monogrn Geol. Paldont. (1) 4: 1-361.
HUuENE, F. Von. 1935-1942. Die Fossilen Reptilien des Sudamerikanischen Gondwanalandes.
Munich: Beck’sche Verlagsbuchhandlund.
HUuENE, F. von. 1956. Palaontologie und Phylogenie der niederen Tetrapoden. Jena: Fischer.
LuLL, R. S. 1912. The life of the Connecticut Valley.— Am. J. Sci. 33: 397-422.
LuLL, R. S. 1915. Triassic life of the Connecticut Valley.— Bull. Conn. St. geol. nat. Hist. Sury.
24: 1-285.
LuLL, R. S. 1953. Triassic life of the Connecticut Valley. Revised edition.— Bull. Conn. St.
geol. nat. Hist. Surv. 81: 1-331.
LYDEKKER, R. 1890. Catalogue of fossil reptilia and amphibia. Part 4. London: British Museum
(Natural History).
Marsh, O. C. 1882. Classification of the Dinosauria.— Am. J. Sci. 23: 81-86.
Marsh, O. C. 1885. Names of extinct reptiles.— Am. J. Sci. 37: 331-336.
Marsu, O. C. 1889. Notice of new American Dinosauria.— Am. J. Sci. 37: 331-336.
Marsh, O. C. 1891. Notice of new vertebrate fossils.— Am. J. Sci. 42: 265-269.
Marsu, O. C. 1892. Notes on Triassic Dinosauria.— Am. J. Sci. 43: 543-546.
Marsu, O. C. 1893. Restoration of Anchisaurus.—Am. J. Sci. 45: 169-170.
Marsu, O. C. 1895. On the affinities and classification of the dinosaurian reptiles.— Am. J. Sci.
50: 483-498.
Marsh, O. C. 1896. The dinosaurs of North America.— Rep. U.S. Geol. Surv. (1): 133-244.
Meyer, H. von. 1837. Briefliche Mitteilung an Prof. Bronn iiber Plateosaurus engelhardti.—
Neues Jb. Miner. Geogr. Geol. Petrefakt. 1837: 316.
Morais, J. 1843. A catalogue of British Fossils. London: British Museum (Natural History).
Owen, R. 1854. Descriptive catalogue of the fossil organic remains of Reptilia contained in
the Museum of the Royal College of Surgeons of England. London: British Museum
(Natural History).
RAATH, M. A. 1972. Fossil vertebrate studies in Rhodesia: a new dinosaur (Reptilia, Saurischia)
from near the Trias-Jurassic boundary.— Arnoldia (Rhod.) 30: 1-37.
Relic, O. A. 1963. La presencia de dinosaurios saurisquios en los ‘Estratos de Ischigualasto’
(Mesotriasico Superior) de las Provincias de San Juan y la Rioja (Republica Argentina).
—Ameghiniania 3: 3-20.
158 ANNALS OF THE SOUTH AFRICAN MUSEUM
Ritey, H. & StrutcHBury, S. 1836. A description of various fossil remains of three distinct
saurian animals discovered in the autumn of 1834, in the Magnesian Conglomerate on
Durdham Down, near Bristol.— Proc. geol. Soc. 2: 397-399.
Ritey, H. & StutcHBurRY, S. 1840. A description of various fossil remains of three distinct
saurian animals, recently discovered in the Magnesian Conglomerate near Bristol.—
Trans. geol. Soc. Lond. 5: 349-357.
Romer, A. S. 1923. The pelvic musculature of saurischian dinosaurs.— Bull. Am. Mus. Nat.
Hist. 48: 605-617.
Romer, A. S. 1956. Osteology of the Reptiles. Chicago: University of Chicago Press.
Romer, A. S. 1966. Vertebrate paleontology. (3rd ed.) Chicago: University of Chicago Press.
ROZHDESTVENSKY, A. K. 1966. Growth changes in Asian dinosaurs and some problems of
their taxonomy (translation of 1965).—Jnt. Geol. Rev. 8: 782-793.
SEELEY, H. G. 1891. On Agrosaurus mcgillivrayi, a saurischian reptile from the north east
coast of Australia.— Q. Jl. geol. Soc. Lond. 47: 164-165.
SEELEY, H. G. 1894. On Hortalotarsus skirtopodus, a new saurischian fossil from Barkly East,
Cape Colony.— Ann. Mag. nat. Hist. 14: 411-419.
SEELEY, H. G. 1895a. On Thecodontosaurus and Palaeosaurus.—Ann. Mag. nat. Hist. 15:
144-153.
SEELEY, H. G. 1895b. On the type of the genus Massospondylus, and on some vertebrae and
limb-bones of M. (?) browni.— Ann. Mag. nat. Hist. 15: 102-125.
STEEL, R. 1970. Handbook of Palaeontology. Part 14. Saurischia. Portland: G. Fischer.
VAN Hoepen, E. C. N. 1920a. Contributions to the knowledge of the reptiles of the Karroo
Formation. 5. A new dinosaur from the Stormberg Beds.— Ann. Transy. Mus. 7: 77-92.
VAN Hoepen, E. C. N. 19205. Contributions to the knowledge of the reptiles of the Karroo
Formation. 6. Further dinosaurian material in the Transvaal Museum.— Ann. Transy.
Mus. 7: 93-141.
Waite, T. E. 1973. Catalogue of the genera of dinosaurs.— Ann. Carneg. Mus. 44: 117-155.
WILb, R. 1974. Die Triasfauna der Tessiner Kalkalpen. XXIII. Tanystropheus longobardicus
(Bassani).— Schweiz. palaeont. Abh. 95: 1-162.
YOuNG, C.-C. 1941a. A complete osteology of Lufengosaurus huenei Y oung (gen. et sp. nov.).
Palaeont. sin. (C) 7: 1-53.
YOUNG, C.-C. 19415. Gyposaurus sinensis (sp. noy.), a new Prosauropoda from the Upper
Triassic Beds at Lufeng, Yunnan.— Bull. geol. Soc. China 21: 205-253.
YounG, C.-C. 1942. Yunnanosaurus huangi (gen. et sp. noy.), a new Prosauropoda from the
Red Beds at Lufeng, Yunnan.— Bull. geol. Soc. China 22: 63-104.
Youna, C.-C. 1947. On Lufengosaurus magnus (sp. nov.) and additional finds of Lufengosaurus
huenei Young.— Palaeont. sin. 12: 1-53.
YouncG, C.-C. 1948. Further notes on Gyposaurus sinensis Young.— Bull. geol. Soc. China 28:
91-103.
YOUNG, C.-C. 1951. The Lufeng saurischian fauna in China.—Palaeont. sin. 13: 1-96.
ABBREVIATIONS
acetabulum acet
anterior process of ilium ant proc
astragalus as
centrum c
caudal vertebra ca
calcaneum cal
chevron ch
dorsal vertebra dor
neural spine ns
pleurocoel pl
postzygapophysis poz
prezygapophysis prz
ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 159
pubic peduncle
sacral vertebra
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Staatlichen Museum fiir Naturkunde in Stuttgart
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P. M. GALTON & M. A. CLUVER
ANCHISAURUS CAPENSIS (BROOM) -
AND A REVISION OF THE ANCHISAURIDAE
(REPTILIA, SAURISCHIA)
VOLUME 69 PART 7 MARCH 1976 ISSN 0303-2515
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war 26 1976
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Zool. exp. gén. 74: 627-634.
Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. —
Ann. Mag. nat. Hist. (13) 2: 309-320.
Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. —
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. — Denkschr. med.-naturw. Ges. Jena 16: 269-270.
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 69 Band
March 1976 Maart
Pact | Deel
YABEICERAS (CONIACIAN AMMONITE) FROM THE
ALPHARD GROUP OFF THE SOUTHERN
CAPE COAST
By
H. C. KLINGER, W. J. KENNEDY & W. G. SIESSER
Cape Town Kaapstad
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are issued in parts at irregular intervals as material
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YABEICERAS (CONIACIAN AMMONITE) FROM THE ALPHARD
GROUP OFF THE SOUTHERN CAPE COAST
By
H. C. KLINGER
South African Museum, Cape Town
W. J. KENNEDY
Department of Geology and Mineralogy, Oxford University
&
W. G. SIESSER
Marine Geoscience Unit, University of Cape Town
(With 4 figures)
[MS accepted 30 October 1975]
ABSTRACT
During dredging operations off the southern Cape coast a bored concretion containing
a specimen of the ammonite Yabeiceras manasoaense Collignon was recovered. The species
had been previously recorded only from the Coniacian of Madagascar and Japan; the occur-
rence extends the geographic range of the species and provides reliable dating for the out-
cropping offshore Mesozoic Alphard Group of sediments of the area.
CONTENTS
PAGE
Introduction . ; : = 6
Material ; é : = L6l
Systematic Palaeontology = 62
Acknowledgements : z 67
References . , : elon
INTRODUCTION
During dredging operations off the southern Cape coast, undertaken by
the Marine Geoscience Unit of the University of Cape Town, a concretion
containing an ammonite was brought to the surface at 35.06S, 20.32E from a
depth of 110 metres. The specimen was identified as belonging to the genus
Yabeiceras, thus far only described from the Coniacian stage of the Upper
Cretaceous of Madagascar and Japan. Apart from adding to our knowledge
of the geographical distribution of the genus, this record permits a precise
dating of the offshore Alphard Group of sediments in the area. The only other
recorded Mesozoic ammonite from the South African offshore is an Eubaculites
sp. recorded by Dingle (1973: 10), although occurrences of Tertiary nautiloids
were reported by Cayeux (1934) and Miller & Furnish (1956).
MATERIAL
The concretion containing the ammonite consists of a dark greyish-green,
fine-grained quartz siltstone, with a calcite cement. The ammonite itself is
161
Ann. S. Afr. Mus. 69 (7), 1976: 161-168, 4 figs.
162 ANNALS OF THE SOUTH AFRICAN MUSEUM
partially filled with sparry calcite, but the greater part is preserved as an internal
mould, consisting of the same material as the concretion. Part of the original
aragonitic shell has been preserved.
That part of the concretion and the ammonite which was exposed at the
sediment/water interface is pitted with two types of borings of Recent orga-
nisms. The first consists of large (up to 1 cm in diameter) flask-like crypts,
internally smooth, with a constricted aperture. The second is much smaller,
2 to 3 mm across and irregular in habit. Both types of borings show cross-
cutting relationships. The larger may be ascribed to lithodomous bivalves,
the smaller to polychaete worms, and perhaps other organisms. Apart from
a few encrusting ectoprocts (bryozoans) and serpulids, most of which occur
within the borings, no other epizoans are present.
SYSTEMATIC PALAEONTOLOGY
Family Collignoniceratidae Wright & Wright, 1951
Subfamily Barroisiceratinae Basse, 1947
Genus Yabeiceras Tokunaga & Shimizu, 1926
(= Eboroceras Basse, 1947)
Type species
Yabeiceras orientale Tokunaga & Shimizu, 1926 by original designation.
Discussion
Matsumoto et al. (1964; Matsumoto 1969) have provided recent reviews
of Yabeiceras and demonstrated that it should be referred to the Barroisi-
ceratinae rather than to the Peroniceratinae, as in the current Treatise (Wright
1957: L429).
Yabeiceras is an uncommon genus represented by four species in Japan:
Y. orientale Tokunaga & Shimizu, Y. kotoi Tokunaga & Shimizu, Y. himuroi
Tokunaga & Shimizu, and Y. manasoaense Collignon, whilst Basse (1946) and
Collignon (1965) record six species from Madagascar: Y. magnumtuberculatum
Basse, Y. manasoaense Collignon, Y. menabense Collignon, Y. costatum Col-
lignon and Y. ankinatsyense Collignon. Undescribed species also occur in
Zululand (Kennedy & Klinger 1975). All records of Yabeiceras are of either
Lower or Middle Coniacian age.
Yabeiceras manasoaense Collignon, 1965
Figs 1-4
Yabeiceras manasoaense Collignon, 1965: 84, pl. 452 (fig. 1839).
Matsumoto 1971: 144, pl. 24 (55) (fig. 2), text-fig. 9 (110).
Holotype
The specimen figures by Collignon (1965, pl. 452 (fig. 1839)) from the
Coniacian of Manasoa (Betioky), Madagascar.
YABEICERAS (CONIACIAN AMMONITE) OFF THE SOUTHERN CAPE COAST 163
Fig. 1. Yabeiceras manasoaense specimen 4492 left lateral view x 1.
Material
Sample 4492 from the Alphard Group at 35.06S, 20.32E, and housed
with the Marine Geoscience Unit, University of Cape Town collections.
Description
The specimen comprises just over two whorls of phragmocone and an
incomplete body chamber of slightly more than a third of a whorl. The inner-
most whorls up to a diameter of 10 mm are not preserved. Coiling is very
evolute with an umbilical diameter of 54,6 per cent of the total diameter. The
outer whorls embrace only slightly, covering less than 10 per cent of the previous
164 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 2. Yabeiceras manasoaense specimen 4492 right lateral view x 1.
whorl. The whorl section of the body chamber is coronate, with maximum
breadth across the dorsal third of the flanks. The venter is ornamented by a
low broad keel, bounded on either side by two equally broad depressions, in
turn flanked by low lateral keels.
Flank ornament consists of a single row of tubercles numbering sixteen
on the outer whorl. On the inner whorls the tubercles are conical to pointed,
and are housed in notches in the umbilical wall of the succeeding whorl. With
increasing diameter the tubercles become more bullate and migrate progressively
from the umbilical suture towards the midflank, and eventually to the dorsal
third of the flanks. Ornament declines markedly on the body chamber.
YABEICERAS (CONIACIAN AMMONITE) OFF THE SOUTHERN CAPE COAST 165
Fig. 3. Yabeiceras manasoaense specimen 4492. A. Ventral view x 1.
B. Dorsal view x 1.
166 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 4. External suture line of Yabeiceras manasoaense Collignon, specimen 4492 at whorl
breadth of 30 mm x 3
Dimensions
Specimen D Wb Wh Wb/Wh U
4492 5 2 re. 92,5 34669) 215233) "8 50,5(54,6)
Holotype O81 Se ce pee 37(35) 24(23) 1,5 51(49)
(Collignon 1965)
Matsumoto 1971 . . = 149 38,6(26) 43,2(29) 0,9 72,4(48)
(Muramoto coll. Ob-S-6-p1)
Discussion
Differences between the nine described species of Yabeiceras are slight,
and well within the range of variation documented in related ammonite groups.
Study of large populations will probably show that most of the described
forms are variants of one or two variable species. Available material is inade-
quate for any constructive discussion of this point, however. The present
specimen compares most closely with Yabeiceras manasoaense, showing a
similar whorl section and ornament which declines on the outer whorls. Dimen-
sions are closely similar to that of the holotype. The larger specimen figured
and described by Matsumoto (1971: 144, pl. 24 (fig. 2)) is virtually identical
at similar diameters, and shows further development of the declining ornament
seen on the body chamber of our specimen.
Yabeiceras bituberculatum Collignon (1965: 821, pl. 451 (fig. 1836), pl. 452
(fig. 1838)) is readily distinguished on the basis of the presence of ventral
tubercles when young, whilst Collignon’s figure shows what appears to be a
YABEICERAS (CONIACIAN AMMONITE) OFF THE SOUTHERN CAPE COAST 167
siphonal row of elongate clavi rather than a continuous keel at this diameter,
whilst the adult body chamber is remarkably smooth and constricted.
Yabeiceras magnumbilicatum (Basse) (1946: 73, fig. 2, pl. 2 (figs 2a—b);
Collignon 1965: 82, pl. 451 (fig. 1836)) has larger, coarser umbilical nodes, a
contracted, virtually smooth body chamber, and far more involute coiling
(U = 35 per cent of diameter).
Y. costatum Collignon (1965: 87, pl. 454 (fig. 1841)) is a costate, rather
than tuberculate species, with 26-28 ribs per whorl, as is Y. ankinatsyense
Collignon (1965: 87, pl. 454 (fig. 1842)).
Y. menabense Collignon (1965: 86, pl. 453 (fig. 1840)) is characterized by
a very depressed whorl section, evolute coiling and 12-15 massive tubercles
per whorl.
Y. magnumbilicatum, Y. bituberculatum and Y. manasoaense are contempo-
raries, as are Y. menabense, Y. costatum and Y. ankinatsyense. It is difficult to
see these as more than one, or perhaps two species, whilst the Y. costatum
group is scarcely distinguishable from the type species, Y. orientale or the
costate Y. himuroi and Y. kotoi (Tokunaga & Shimizu 1926).
Occurrence:
Y. mansoaense is recorded from the Lower to Middle Coniacian of Japan
and the Middle Coniacian Kossmaticeras theobaldianum|Barroisiceras onila-
hyense Zone of Madagascar. In Zululand related, but as yet undescribed, forms
occur in the St. Lucia Formation in the second division of the Coniacian,
associated with Forresteria alluaudi (Boule, Lemoine & Thevenin), Proplacen-
ticeras spp. and other forms, again suggesting an early Coniacian age.
ACKNOWLEDGEMENTS
We should like to express our thanks to the Marine Geoscience Unit for
placing the material and data at our disposal. Thanks are due to Mr Neville
Eden for the photography.
REFERENCES
Basse, E. 1946. Sur deux ammonites nouvelles du Coniacien du Sud-ouest de Madagascar:
Subbarroisiceras n.g. mahafalense n. sp. et Eboroceras n.g. magnumbilicatum n. sp.—
Bull. Soc. géol. Fr., 5 ser, 16: 71-76, pl. 2.
Basse, E. 1947. Les peuplements malgachés de Barroisiceras.—Ann. Paléont. 33: 99-178,
pl. 1(7)-9(15).
Cayeux, L. 1934. The phosphatic nodules of the Agulhas Bank.—Ann. S. Afr. Mus. 31:
105-135, pls 32-35.
COLLIGNON, M. 1965. Atlas des fossiles caracteristiques de Madagascar (Ammonites). 13
(Coniacien). Tananarive: Service Geologique.
DinGc_e, R. V. 1973. Post-Palaeozoic stratigraphy of the eastern Agulhas Bank, South African
continental margin. — Mar. Geol. 15: 1-23.
KENNeDyY, W. J. & KLINGER, H. C. 1975. Cretaceous faunas from Zululand and Natal, South
Africa. Introduction, stratigraphy.— Bull. Br. Mus. nat. Hist. (Geol.) 25: 265-315.
168 ANNALS OF THE SOUTH AFRICAN MUSEUM
Matsumoto, T. 1969. A Monograph of the Collignoniceratidae from Hokkaido. Part III.
(Studies of the Cretaceous Ammonites from Hokkaido and Saghalien—xx.)— Mem. Fac.
Sci. Kyushu Univ. (D) 19: 297-330.
Matsumoto, T. 1971. A Monograph of the Collignoniceratidae from Japan. Part V. (Studies
of the Cretaceous Ammonites from Hokkaido and Saghalien—xxiii.)— Mem. Fac. Sci.
Kyushu Univ. (D) 21: 129-162.
Matsumoto, T., OBATA, I., MAEDA, S. & Sato, T. 1964. Yabeiceras (Cretaceous ammonites)
from Futaba, Northeast Japan.—Trans. Proc. palaeont. Soc. Japan (N.s.) 55: 322-331.
Miter, A. K. & FurRNISH, W. M. 1956. Tertiary Nautiloids dredged near Cape of Good
Hope. — Ann. S. Afr. Mus. 42: 327-328.
ToKUNAGA, S. & SuHimizu, S. 1926. The Cretaceous Formation of Futaba in Iwaki and its
fossils.—J. Fac. Sci. Tokyo Univ. (2) 1: 181-212.
WRIGHT, C. W. 1957. In R. C. Moore ed. Treatise on invertebrate paleontology Pt I, Mollusca,
Cephalopoda, Ammonoidea. Lawrence: University of Kansas Press.
WRIGHT, C. W. & WriGutT, E. V. 1951. A survey of the cephalopoda of the Chalk of Great
Britain. London: Palaeontographical Society.
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H. C. KLINGER, W. J. KENNEDY & W. G. SIESSER
YABEICERAS (CONIACIAN AMMONITE) FROM
THE ALPHARD GROUP OFF THE
SOUTHERN CAPE COAST
VOLUME 69 PART 8 APRIL 1976
ESF - CJaehur ]
_ COMP. ZOOL.
= RY
JUN 2 1 1976
OF THE SOUTH AFRIC
MUSEUM
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FIscHER, P.-H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. — Archs
Zool. exp. "gen. 74: 627-634.
Konn, A. J. 1960a. Ecological — on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. —
Ann. Mag. nat. Hist. (13) 2: 309-320.
Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. —
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
THEE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. — Denkschr. med.-naturw. Ges. Jena 16: 269-270.
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 69 Band
April 1976 April
Part 8 Deel
HE CRANIAL AND CERVICAL MUSCLES OF THE
SOUTH AFRICAN LIMBLESS LIZARD
TYPHLOSAURUS AURANTIACUS AURANTIACUS
PETERS (REPTILIA, SAURIA)
By
JURI A. VAN DEN HEEVER
Cape Town Kaapstad
The ANNALS OF THE SOUTH AFRICAN MUSEUM
are issued in parts at irregular intervals as material
becomes available
Obtainable from the South African Museum, P.O. Box 61, Cape Town
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OUT OF PRINT/UIT DRUK
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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap
THE CRANIAL AND CERVICAL MUSCLES OF THE SOUTH AFRICAN
LIMBLESS LIZARD TYPHLOSAURUS AURANTIACUS AURANTIACUS
PETERS (REPTILIA, SAURIA)
By
JuRI A. VAN DEN HEEVER
South African Museum, Cape Town
(With 18 figures)
[LMS accepted 25 November 1975]
ABSTRACT
The skull and cervical vertebrae of Typhlosaurus aurantiacus aurantiacus exhibit marked
adaptations to a fossorial mode of life which is reflected in the associated musculature. Neither
eye muscles nor supratemporal arches are present. The adductor musculature of the jaws,
compacted into the temporal region, are dominated by two tripartite tendons, the bodenapo-
neurosis and the quadrate tendon. The m. pseudotemporalis is single. The m. cervicoman-
dibularis is probably the major jaw opening muscle. Posttemporal fenestrae are absent and
the cervical musculature encroaches far anteriorly on to the bulbous occiput. Discussed in
terms of a lever of the third class the action of the jaw shows great similarity to that of a
non-fossorial skink like Mabuia, and probably functions in an identical manner. Primary
adaptations for a fossorial mode of life appear to be the strengthening and streamlining of the
skull, loss of limbs and limb girdles and general attenuation of the body.
CONTENTS
IRCLOMUCHOM es. 5) Rise? Sah rae evita coo
Materialand methods ....... . I7Il
Description
Osteology
Granmlosieclogy< 7) £9 0 oP
Geryical.osteologyey. “ai. sce yen eee 179
Myology
Muscle classification and nomenclature . 181
Constrictor dorsalis gooup. . . . . 182
Adductor mandibulae group . . . . 184
Intermandibular musculature. . . . 194
Tongue musculature . . . . . . 195
Depressor mandibulae group . . . . 197
Cervical musculature. <7 ~0058%) 16 202
Discussion and conclusions . . . . . . . 204
Acknowledgements. “7. . 3) . “aye = Seto
Reterenees a0)... ac ere oe, a ee 210
ADBEVIATIOOS: fren Y tuivees’ bel) bce Be Seek telS
INTRODUCTION
~The doubtful taxonomic position of Typhlosaurus has been dealt with by
various authors. Boulenger (1887) regarded the genus as related to Acontias but
placed it, like Gadow (1901), together with Anelytropsis and Feylinia in the
family Anelytropidae, close to the Scincidae. Camp (1923) places Typhlosaurus
169
Ann. S. Afr. Mus. 69 (8), 1976: 169-214, 18 figs
170 ANNALS OF THE SOUTH AFRICAN MUSEUM
in the Feyliniidae, within the superfamily Scincoidea, together with the Scin-
cidae, the Anelytropsidae and the Dibamidae.
Hewitt (1929), De Witte & Laurent (1943) and FitzSimons (1943) also
refer the genus to the Scincidae and according to Smit (in press) FitzSimons
regards Typhlosaurus as derivable, via Aconthophiops, from Acontias. Romer
(1956) rather doubtfully includes Typhlosaurus within the Scincidae. Greer
(1970), on the basis of the external morphology and cranial osteology especially
the relationship of the frontal bones and the bones of the secondary palate,
regards Acontias, Aconthophiops and Typhlosaurus as a subfamily of the Scinci-
dae, i.e. the Acontinae. Broadley (1968) agrees with FitzSimons and states:
‘The genus Typhlosaurus appears to have been derived from an ancestral form
of Acontias, after passing through an intermediate stage which is demonstrated
by the monotypic genus Aconthophiops.’ According to Smit (1964) the close
relationship between Typhlosaurus and Acontias is abundantly confirmed by
the cranial osteology of JT. caecus, indicating that the genus Typhlosaurus
undoubtedly belongs within the Scincidae.
The following classification is thus adopted.
Class: Reptilia
Order: Squamata
Suborder: Sauria
Family: Scincidae
Subfamily: Acontinae Greer, 1968
Genus: Typhlosaurus Wiegmann, 1834
Species: T. aurantiacus Peters, 1882
Subspecies: T. aurantiacus aurantiacus Broadley, 1968.
Limbless skinks of the genus Typhlosaurus are confined to southern Africa
(Broadley 1968). Eight species were recorded by FitzSimons (1943), and one
additional species, the greatly attenuated 7. braini from the Namib Desert,
was described by Haacke (1964). Subsequently Broadley (1968) revised the
genus, recognizing eight species placed into three species groups.
The genus as a whole is fossorial and according to Mertens (1955) only
appears on the surface towards evening. Their diet includes small insects and
myriapods of which small beetle latvae and termites form the most important
groups (Broadley 1968).
The cranial osteology of the fossorial Scincidae and forms with scincid
affinities, such as Dibamus, are well known from the work done on Acontias
(De Villiers 1939; Brock 1941; Van der Merwe 1944), Dibamus (Gasc 1968;
De Weerdt 1971), Typhlosaurus (Smit 1964), Feylinia (Du Toit 1971), Typhla-
contias (Cluver 1965), Melanoseps (Boyd 1969) and Scelotes (Leonard 1973).
The postcranial skeleton is less well known and except for the work of
Gasc (1967a, b, c; 1968) and Hofstetter & Gasc (1969) has attracted few
investigators,
CRANIAL AND CERVICAL MUSCLES OF TYPHLOSAURUS AURANTIACUS AURANTIACUS iW 8 |
Myological studies on limbless lizards are few and far between. According
to Auffenberg (1962) there is no record of the axial muscles of limbless lizards
up to that time. Except for the work of Gasc (1968), De Weerdt (1971) and
Leonard (1973) the cranial muscles of limbless lizards are largely unknown.
De Weerdt’s account differs from the more detailed description of Gasc. How-
ever, Gasc’s description is difficult to evaluate in the light of comments given
by Haas (1973), and Leonard discusses only the eye muscles in Scelotes.
Haas (1973) reviews the jaw muscles of the Rhynchocephalia and the
Squamata stating that: ‘*. . . detailed studies of the cranial muscles are lacking
for two families (or groups often considered to be families) of lizards, namely
the Anelytropsidae and the Feyliniidae’. Greer (1970) assigned these two groups
to the Scincidae as the subfamilies Acontinae and Feylininae. Together they
represent the fossorial Scincidae of which the above statement is certainly true.
The previously mentioned studies have shown that a fossorial habit pro-
duces distinctive skeletal changes. Change in skeletal proportions should
inevitably affect associated musculature and in view of this fact the acute lack
of literature on the myology of fossorial Scincidae is believed to sufficiently
justify this paper.
MATERIAL AND METHODS
Three alcohol-fixed specimens of Typhlosaurus aurantiacus were obtained
from the South African Museum. One specimen was dissected and the skull
and postcranial skeleton were used for comparative and photographic purposes.
Both the other specimens were decalcified for a period of seven days in 7,5 per
cent solution of nitric acid in 70 per cent alcohol. Subsequently the specimens
were separately dehydrated, cleared in terpineol and embedded in paraffin wax
(52-54°C). Sectioned at 20 microns, one specimen gave excellent results; the
other proved of no use and was discarded. Staining and counter-staining were
done with the azocarmine-azan method and the enlarged drawings of the
sections were made with the aid of a camera lucida microscope attachment.
Owing to the paucity of material several specimens of the related but
more abundant genus Acontias were dissected to elucidate gross topography.
DESCRIPTION
OSTEOLOGY
Cranial osteology
The skull of Typhiosaurus caecus, which closely resembles that of
T. aurantiacus, was described by Smit (1964), and the reader is referred to this
paper for a comprehensive account of the cranial osteology. However, prior
to embarking on a description of the myology, the cranial and the cervical
osteology merit a few additional remarks on certain areas important to muscle
attachments, where Smit’s description is inadequate for the purpose of this
paper
172 ANNALS OF THE SOUTH AFRICAN MUSEUM
T. aurantiacus has the elongate skull and reduced orbits common to
attenuate fossorial lizards (Figs 1|A—B, 2A). The eyes are visible as inconspicu-
ous black dots through the transparent integument and are devoid of associated
musculature. However, an optic nerve is present and the retinal pattern resembles
Fig. 1B. Stereophotographs of Typhlosaurus aurantiacus skull; lateral view.
CRANIAL AND CERVICAL MUSCLES OF TYPHLOSAURUS AURANTIACUS AURANTIACUS 173
that of amphisbaenids, in which light perception has been demonstrated by
Bonin (1965).
Fig. 2B. Stereophotographs of Typhlosaurus aurantiacus lower jaw;
lingual view.
174 ANNALS OF THE SOUTH AFRICAN MUSEUM
No remnants of lacrimal bones were found. The supratemporal arch is
absent but both the squamosal and supratemporal bones are present as small,
flattened, slightly overlapping elements dorsal to the quadrate.
The squamosal, lying anterolateral to the supratemporal, is tendinously
connected to the quadrate head (Figs 3, 13). The supratemporal lies postero-
medial to the squamosal and shares with the much reduced paroccipital process
of the otic capsule the articulation with the quadrate head by means of a pad
of fibrocartilage (Fig. 3).
PAR
Fig. 3. Posterolateral view of the skull.
The proportions of the posterior half of the skull are of importance. The
temporal region, where the adductor musculature is accommodated, is laterally
compressed and the otico-occipital region is much expanded both laterally
and posteriorly (Figs 1A—B, 2A). These relations create the impression that the
suspensorium is more anteriorly located than in a non-fossorial lizard such as
Mabuia. The entire posterior border of the parietal meets the supraoccipital
and the fused exoccipital-opisthotic complex in a dorsally lying suture. There
is thus no posttemporal fenestra and the back of the skull is smooth and bluntly
CRANIAL AND CERVICAL MUSCLES OF TYPHLOSAURUS AURANTIACUS AURANTIACUS 175
rounded (Fig. 1A). Immediately below the foramen magnum a kidney-shaped
condyle is present, composed laterally of the exoccipitals and medially of a
suturally distinct basioccipital bone. Ventrally the basioccipital forms the
posterior portion of the skull base, curving upwards to meet the exoccipital
ventrolaterally to the condyle.
A tympanum and middle ear cavity are absent. The columella is massive,
consisting of a large footplate and a short anterolaterally directed stapes
(Figs 3-4), which extends laterally beyond the quadrate as a rod-like carti-
laginous extracolumella, terminating a short distance anterior to the quadrate
Fig. 4. Posteroventral view of the skull.
and ventrolateral to the medius portion of the external adductor muscle (Figs 5,
7-8, 12-13). The tendinous sheath surrounding the extracolumella is joined to
the retroarticular process of the lower jaw as in Typhlosaurus caecus (Smit 1964)
and Acontias meleagris (De Villiers 1939; Brock 1941; Van der Merwe 1944),
and is suspended anteriorly by a ribbon of fascia overlying the tendinous
covering of the adductor musculature and attaching to the dorsolateral border
of the parietal (Figs 5, 7-8, 12). The same condition exists in T. caecus and
T. lineatus.
176 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 5. Lateral superficial view of musculature.
The quadrate ramus of the pterygoid is gutter-shaped, with the trough
directed medially in the region of the basipterygoid joint to receive the basi-
pterygoid process. Posteriorly the quadrate ramus is twisted through ninety
degrees so that the trough faces ventrally where the posterolateral extremity
of the bone is tendinously connected to the ventromedial surface of the quadrate
(Fig. 4).
The anterior border of the quadrate is gently rounded for the attachment
of the lateral lamina of the quadrate tendon, while the posterior border of the
bone is concave to accommodate the laterally protruding stapes (Fig. 3).
Ventrally the large quadrate condyle articulates synovially with the lower jaw.
Dorsal to the attachment of the quadrate ramus of the pterygoid the medial
surface of the quadrate is slightly concave to accommodate the origin of the
posterior adductor muscle (Fig. 13). Dorsally the posterior part of the quadrate
head articulates synovially with both the supratemporal and the otic capsule
(Figs 3, 14).
The anterior part of the quadrate head is separated from the cranium by
fibres of the medius portion of the external adductor muscle and serves for
the origin of the vertical lamina of the quadrate tendon. The central part of
the quadrate head is tendinously attached to the squamosal (Figs 3, 13).
The thin, rod-like epipterygoid fits ventrally into the columellar fossa on
the dorsal surface of the pterygoid, lateral to the basipterygoid joint. Both
condylar surfaces are capped by cartilage. The dorsal extremity of the bone
attaches tendinously to the lateral surface of the parietal downgrowth immedi-
ately in front of the anterior superior process of the pro-otic (Fig. 4).
Because of the importance of the mandibular muscle insertions it is neces-
sary to augment the brief description of Smit (1964) of the lower jaw of Typhlo-
saurus caecus. Each ramus consists of the normal six bones, viz. dentary,
coronoid, splenial, surangular, angular and a fused articular-prearticular, to
CRANIAL AND CERVICAL MUSCLES OF TYPHLOSAURUS AURANTIACUS AURANTIACUS 177
which the surangular is also partially fused (Fig. 17). Between the coronoid
process and the glenoid fossa both dentary and surangular bulge laterally,
forming a dorsolateral mandibular shelf on to which elements of the adductor
musculature insert (Fig. 11). Posteriorly, at the insertion of the pterygomandi-
bularis muscle, the ventral border of the jaw is concave (Fig. 17).
The dentary extends from the symphysis to the supra-angular foramen and
constitutes the anterior portion of the mandibular shelf (Figs 9-10). It carries
eight to nine pleurodont teeth, contributes the lateral half of the coronoid
process and is medially recessed along its posterior half to accommodate the
remaining lower jaw bones (Fig. 17). It is pierced laterally and antero-ventrally
below the tooth row by a line of four mental foramina, and postero-medially
by a large foramen transmitting the anterior mylohyoid nerve and the lingual
branch of the inferior alveolar nerve; anteromedially there are foramina for the
Meckelian cartilage and the anterior tip of the inferior alveolar nerve. The
position of the posteromedial foramen varied in the two Typhlosaurus auranti-
acus specimens investigated. In the serially sectioned skull the anterior process
of the coronoid bone and the anterior tip of the splenial form the posterior
border of the foramen, whereas, in the cleared specimen, the foramen lies well
within the boundaries of the dentary (Fig. 17). Behind the coronoid process,
fibres of the posterior adductor muscle and the medius portion of the external
adductor muscle insert along the dorsolateral border of the dentary.
The coronoid lies midway along the mandible, flattened medially against
the dentary, prearticular and surangular. Its coronoid process is a prominent
vertical sheet lying against the coronoid process of the dentary (Figs 2B, 9).
Ventrally the bone is braced against the action of the adductor muscles by an
anterior and a posterior process, the former bridging the dentary-prearticular
suture and the latter the surangular-prearticular suture (Fig. 17). Two parallel,
near-vertical grooves are present on the trailing edge of the coronoid process.
Separated by a ridge, they continue posteroventrally on to the medial side of
the posterior process; the more lateral of the two grooves receives the insertion
of the medius portion of the external adductor muscle while the pseudotem-
poralis muscle inserts into the medial groove (Figs 10, 17).
The surangular (Figs 9-12, 17) lies posteromedially to the dentary, postero-
laterally to the coronoid, dorsally to the prearticular and anteriorly to the
articular. Laterally it forms the posterior section of the mandibular shelf
(Fig. 11) and dorsally it bears a ridge extending between the coronoid and the
articular processes. Its anterior extremity underlies the coronoid process while
in addition to covering the Meckelian canal up to the adductor fossa, the
posterodorsal tip of the bone constitutes the anterior part of the articular
process. Posteromedially the bone forms the dorsal border of the adductor
fossa and is laterally pierced by the posterior supra-angular foramen, which
leads from the adductor fossa, and the anterior supra-angular foramen which
leads from the Meckelian canal. The posterior adductor muscle and medius
portion of the external adductor muscle insert on the dorsomedial and dorso-
178 ANNALS OF THE SOUTH AFRICAN MUSEUM
\_NI Imm
Fig. 6. Dorsal superficial view of the cervical musculature; spinalis capitis muscle removed
on the right side.
lateral surfaces of the surangular. Posteriorly the bone is fused to the articular-
prearticular complex.
The splenial (Figs 9-10) is a thin sliver of bone on the inner surface of
the jaw in line with the coronoid process. It lies posteromedially to the dentary,
medially to the prearticular, dorsally to the angular and ventrally to the coro-
noid. It is devoid of any muscle insertions and neither bears foramina nor
contributes to the inner wall of the Meckelian canal.
The angular (Figs 9-10, 11, 17), a narrow ventral element below the pre-
articular and splenial, lies with its anterior tip within the dental recess and
traverses the jaw posteroventrally between the prearticular and dentary to
terminate posteriorly on the lateral surface of the mandible, ventral to the
posterior tip of the surangular. Between the prearticular and dentary it forms a
narrow medial section of the floor of the Meckelian canal, from the adductor
fossa to immediately anterior to the coronoid process. At its midpoint it is
pierced ventrally by the posterior mylohyoid foramen, which transmits the
posterior mylohyoid nerve.
The prearticular (Figs 11-12, 17) lies ventrally to the surangular, postero-
medially to the dentary, dorsally to the angular and laterally to the coronoid
and the splenial. It forms most of the medial wall and part of the floor of the
Meckelian canal as well as the ventral border of the adductor fossa. Medially
it receives the insertion of the adductor musculature and posteriorly it is com-
pletely fused to the articular.
Behind the glenoid fossa the articular-prearticular (Figs 13-17) forms the
spoon-shaped retroarticular process, which receives the insertion of the ptery-
goideus muscle on its medial and lateral surfaces and that of the depressor
mandibulae muscle on the dorsal surface. The glenoid fossa, lined with cartilage,
lies in front of the insertion of the depressor mandibulae muscle and behind and
against the articular process. The foramen for the chorda tympani lies medially
on the retroarticular process, with its canal extending anteriorly through the
CRANIAL AND CERVICAL MUSCLES OF TYPHLOSAURUS AURANTIACUS AURANTIACUS 179
bone to open into the adductor fossa.
The hyoid apparatus in Typhlosaurus aurantiacus is a triradiate, carti-
laginous structure resembling a tuning-fork with posteriorly divergent prongs.
It lies ventral to the trachea and extends posteriorly from the glottis to a point
in line with the posterior border of the pterygoideus muscle. Anteriorly, in the
ventral midline, the lingual process (proc. entoglossus) supports the tongue
musculature (Figs 9-10).
No part of the hyoid apparatus is ossified and the structure could conse-
quently not be divided with certainty into the various components commonly
found in reptiles. From the work of Van der Merwe (1944), Langebartel (1968)
and De Weerdt (1971) it appears that the paired posterior prongs represent
the first ceratobranchials.
Cervical osteology
Vertebrae and ribs are highly variable structures and in the limbless
squamates the regional differentiation of the vertebral column has lead to
various interpretations of the cervical vertebrate. Zangerl (1945) recognizes
four vertebral divisions in the Amphisbaenidae, i.e. cervical, thoraco-lumbar,
cloacal and caudal. The cervical region includes all the anterior ribless verte-
brae, i.e. atlas, axis and one to two of the following vertebrae, whilst the thoraco-
lumbar region includes all vertebrae with movable unforked ribs. Sood (1948)
divides the ophidian vertebral column into a precaudal and a caudal region
of which the former is subdivided into cervical, thoracic and lumbar sub-
regions. The cervical sub-region consists only of the atlas and the axis whereas
the thoracic region includes all vertebrae following the axis and which bear
hypapophyses. List’s (1966) description of the burrowing snakes follows
Zangerl (1945) in defining the regions of the vertebral column. Consequently
he defines the cervical region in burrowing snakes as consisting only of the
atlas and the axis since the vertebrae following the axis bear unforked ribs
and are therefore included in the thoraco-lumbar region. List notes that although
this system appears satisfactory for the Typhlopidae and Leptotyphlopidae its
use is limited in that it cannot be directly applied to other vertebrates.
According to Gasc (1968) and Hofstetter & Gasc (1969) the cervical
vertebrae can only be defined as those vertebrae preceding the vertebrae carrying
the first rib attached to the sternum. These authors refer to the work of Stannius
(1849) and state that all other definitions of cervical vertebrae such as ribless
anterior vertebrae, vertebrae with hypapophyses or ribless vertebrae plus
vertebrae with short ribs are invalid because too many exceptions and contra-
dictions are involved.
Limb regression is usually accompanied by regression of the girdles and
in certain of the fossorial Scincidae, e.g. in the genus Typhlosaurus, this phe-
nomenon is rather pronounced. In Feylinia the ribs of the eighth vertebra are
still attached to a vestigial sternum (Gasc 1965), in Dibamus the pectoral girdle
is connected to the fifth vertebra (Gasc 1968) and in Acontias meleagris the
180 ANNALS OF THE SOUTH AFRICAN MUSEUM
\—EmM
Imm
Fig. 7. Ventrolateral view of the superficial neck musculature.
vestigial girdle is united by the serratus muscle to the second and the third ribs.
In 7. vermis the pectoral girdle is absent (Hofstetter & Gasc 1969). In T. auran-
tiacus the pectoral girdle is aiso absent, and the ribs consequently lack sternal
attachments. It is therefore not possible to define a specific cervical region
within the vertebral column of T. aurantiacus. In limbless squamates such as
ophidians (completely lacking a pectoral girdle) and amphisbaenids (lacking
sternal attachments of the ribs) Hofstetter & Gasc (1969) divide the vertebral
column into precloacal, cloacal and caudal regions.
In this paper the term ‘cervical’ does not define a region of the vertebral
column but refers only to that area on the precloacal region of the vertebral
column from which the musculature responsible for the movements of the head
arise.
Typhlosaurus aurantiacus has procoelous vertebrae as in all saurians
except the Gekkonidae. The broad elliptical condyles are slightly dorsally
orientated and are as wide as the centra of the vertebrae. The first pairs of ribs
are carried by the third vertebra, as in Acontias meleagris.
The ribs are holocephalous (unicipital) and each has two tuberculiform
processes close to the costal head; one anteroventrally and one postero-
dorsally for the attachment of the intercostal muscles. A similar condition
exists in T. vermis (Hofstetter & Gasc 1969). As a result of the increased func-
tional importance of the cervical musculature in a limbless burrower such as
T. aurantiacus, the synapophysis of the axis and the following three vertebrae
are laterally extended to enlarge the area of origin of the cervical musculature.
The atlas consists of paired neural arches and a ventral intercentrum.
A neural spine is absent and the two semilunate neural arches do not fuse
dorsally. No functional zygapophyses are present between the atlas and the
axis, although the atlas has a small process on the posterolateral margin of
the neural arch in a similar position to the postzygapophyses of the other
vertebrae.
The axial centrum bears two hypapophyses, of which the posterior one
is probably derived from the intercentrum of the third vertebra as Holder
(1960) found in gekkos. Anterolaterally the neural arch has a small process
CRANIAL AND CERVICAL MUSCLES OF TYPHLOSAURUS AURANTIACUS AURANTIACUS 181
coinciding with the position of the prezygapophyses on the vertebrae following
the axis. The neural spine resembles the blade of an axe and extends the full
length of the neural arch.
Midventrally on the centra of each of the seven vertebrae following the
axis a hypapophysis is present. The neural spines, posterodorsally situated
on the neural arches, extend obliquely caudally. The vertebrae articulate by
Imm
Fig. 8. Lateral view of the deep cervical musculature.
means of pre- and post-zygapophyseal processes and no zygosphene-zygantrum
type of articulation, as found in snakes and some saurian families, is present
in Typhlosaurus aurantiacus.
MYOLOGY
Muscle classification and nomenclature
The currently accepted classification of visceral cranial muscles is based
on Vetter’s (1874, 1878) and Ruge’s (1897) studies on selachians. The Constrictor
superficialis (Cs) is subdivided into segmentally innervated portions, i.e. the
trigeminus muscle complex as the Constrictor superficialis I (Cs,); the facial
muscles as the Constrictor superficialis II (Cs,); the glossopharyngeal muscles
as the Constrictor superficialis III (Cs); and the vagus muscles as the Constrictor
superficialis IV-VIII (Cs, ,). Luther (1914) extended this classification to
tetrapods and introduced a subdivision of the jaw adductors based on the
spatial relationships of the muscles with the three rami of the trigeminal nerve.
This system has been generally accepted for sauropsids by most authors includ-
ing Adams (1919), Lakjer (1926), Haas (1930, 1934, 1973), Lubosch (1933),
Edgeworth (1935), Brock (1941), Save-Sdéderbergh (1945), Ingeborg Poglayen-
Neuwall (1953, 1954), Ivo Poglayen-Neuwall (1953a, 1953b), Oelrich (1956),
Ostrom (1961), Gasc (1968) and Barghusen (1973). In sauropsids the Con-
strictor I is divided into the m. constrictor I dorsalis (M.C,d), the m. constrictor I
182 ANNALS OF THE SOUTH AFRICAN MUSEUM
lateralis (M.C,l) represented by the m. adductor mandibulae, and a m. con-
strictor I ventralis (M.C,v), the m. intermandibularis.
The constrictor I dorsalis (M.C,d) extends between the cranium and the
movable palatal complex. This group of muscles is involved with kinetic move-
ments of the skull and they are variable in their occurrence.
Basically, the constrictor I lateralis (M.C,l), the m. adductor mandibulae
of sauropsids, is divided into an external, and internal and a posterior muscle.
The external adductor is usually suodivided into three portions, i.e. super-
ficialis, medius and profundus. The internal adductor commonly consists of
two, well-separated muscles, the m. pseudotemporalis and the m. pterygoideus.
The posterior adductor is usually a single muscle.
The constrictor I ventralis (M.C,v), the m. intermandibularis, is situated
between the rami of the lower jaws, superficial to the throat musculature, and
is subdivided into an anterior and a posterior portion.
Lubosch (1933) recongizes three basic arrangements of jaw muscles, i.e.
selachian, amphibian and mammalian, of which the jaw muscles of sauropsids
belong to the amphibian type. Homologies between the three types are uncertain,
according to Haas (1973), and are further complicated by the varied nomen-
clature in existence for saurian jaw musculature. For a complete list of syno-
nyms see Lakjer (1926), Edgeworth (1935) and Haas (1973).
Nishi (1919) laid down the terminology for axial musculature and, together
with Vallois (1922), is amongst the few workers who have approached axial
musculature on a comparative basis. More recent accounts are those of Olson
(1936) and Evans (1939).
Reptilian epaxial musculature is divisible into three longitudinal systems.
Dorsomedially the transversospinalis system lies lateral to the spinous pro-
cesses of the vertebrae, the longissimus system lies lateral to the transverso-
spinalis system and dorsal to the heads of the ribs, and the iliocostalis system
lies on the ribs dorsal to the upper margin of the external oblique abdominal
muscles. Anteriorly, towards the occiput, the epaxial musculature breaks down
into various shorter groups of fibres, the cervical muscles, which insert pos-
teriorly on to the skull and are responsible for the movements of the head.
The hypaxial musculature does not fall within the scope of this paper.
Constrictor dorsalis group (C,d)
This group of muscles lies deep to the adductor musculature and is respon-
sible for the intercranial kinetic movements of the skull. The muscles arise,
in Versluys’ (1912) terminology, on the occipital segment of the skull and insert
on to the maxillary segment. In Typhlosaurus aurantiacus the group is repre-
sented by a minute m. levator pterygoidei and a large m. protractor ptery-
goidei. A levator bulbi muscle is absent.
The small m. levator pterygoidei (lp, Figs 9-10), roughly triangular in
transverse section, lies laterally to the basipterygoid joint, the opthalmic ramus
of the trigeminal nerve, the palatine ramus of the facial nerve, the internal
CRANIAL AND CERVICAL MUSCLES OF TYPHLOSAURUS AURANTIACUS AURANTIACUS 183
sl
Imm
PAR
PF
x
gy
oO
— il
)
(
7
HG — Saw ——s
\ wale
AE
ISN \ \ S050
IT SS
GGL SES
GHM
Fig. 9. Cross-section of the skull at the level of the coronoid process.
carotid artery and the origin of the protractor pterygoidei muscle; medially
to the pseudotemporalis muscle, the anterior part of the pterygoideus muscle
and the epipterygoid. It lies against the medial surface of the epipterygoid
with the posterior border of the muscle in line with that of the bone.
184 ANNALS OF THE SOUTH AFRICAN MUSEUM
The muscle arises as a ribbon of fascia from the lateral surface of the pro-
otic membrane, ventral to the lateral parietal downgrowth, medioventral to
the dorsal extremity of the epipterygoid bone and anteroventral to the anterior
superior process of the pro-otic.
The insertion is fleshy and bridges the palatine-pterygoid suture dorsally.
Anteriorly it attaches on to the posterolateral border of the palatine bone,
medially to the anterior extremity of the pterygoideus muscle, and posteriorly
it attaches on to the anterodorsal surface of the pterygoid bone medially to
the columellar fossa.
A nerve seen within the muscle in transverse section was too small for its
connections to be traced.
The large m. protractor pterygoidei (prp, Figs 11-14) lies behind the
the basipterygoid process, the individual fibres extending obliquely between
the lateral margin of the skull base and the full length of the quadrate ramus
of the pterygoid. The muscle is situated posteromedially to the levator ptery-
goidei muscle, medially to the mandibular ramus of V, dorsally to the ptery-
goideus muscle, ventrally to the Gasserian ganglion and the proximal part of
the opthalmic ramus, and laterally to the parasphenoid-basisphenoid complex,
the otic capsule, the palatine ramus of VII, and the internal carotid artery.
The muscle arises fleshily from the dorsal surface of the basipterygoid
process, the lateral surfaces of the parasphenoid-basisphenoid complex and
the anterior inferior process of the pro-otic, and anteroventrally from the
lateral surface of the pro-otic proper (Figs 9-11). Anteroventrally within the
muscle a flat tendon is present which arises ventrally on the basipterygoid
process. Fibres arise from both dorsal and ventral surfaces of the tendon.
The insertion is confined to the quadrate ramus of the pterygoid bone
posterior to the basipterygoid process. Fibres insert along the inner concave
surface and the dorsal surface of the ramus. An insertional tendon is present
within the muscle, posterodorsally to the tendon of origin, the fibres insert
on to its dorsal and ventral surfaces. This tendon attaches to the dorsal (inner)
rim of the quadrate ramus (Figs 11-12).
The muscle is innervated by a separate branch of V leaving the Gasserian
ganglion ventrally, piercing the muscle dorsally and coursing anteriorly a short
distance before ramifying.
Adductor mandibulae group
Compared with non-fossorial lizards, the adductor musculature as exemplified
by Typhlosaurus aurantiacus is modified to function as a compact unit within
the temporal indentation. The external adductors are dominated by two oblique,
parallel tendons, of which the anterodorsal one is a modified bodenaponeurosis.
It arises as a single tendon on the coronoid process of the lower jaw (Fig. 9),
posterior to which it fans out into the external adductor mass as three laminae.
In transverse section this unit appears as a tripartite structure resembling an
inverted Y (Fig. 11). The three laminae serve as areas of insertion to the external
———
CRANIAL AND CERVICAL MUSCLES OF TYPHLOSAURUS AURANTIACUS AURANTIACUS 185
and internal adductors and will be referred to in the text as the vertical, medial
and lateral laminae of the bodenaponeurosis.
The second tendon (Figs 11-12), located posteroventrally to the bodenapo-
neurosis, also shows a tripartite configuration. It arises on the quadrate and
extends anteroventrally to just posterior to the base of the coronoid process.
This tendon will be referred to as the quadrate tendon and its laminae as the
vertical, medial and lateral laminae of the quadrate tendon. Each lamina has a
separate origin from the quadrate. The vertical lamina arises from the dorsal
midline of the quadrate, piercing the medius portion of the external adductor
muscle ventromedially. The lateral lamina arises from the anterior border of
the bone and the medial lamina along its inner dorsal surface (Fig. 13), covering
the medial surface of the posterior adductor fibres arising from the medial
surface of the quadrate.
Ventrolaterally to the superficial portion of the external adductor and
anteromedially to the extracolumella, a ligament is present in the position
of the quadrato-maxillary ligament as described by Ingeborg Poglayen-Neuwall
(1953) and Haas (1960). In Typhlosaurus aurantiacus, however, this ligament
arises ventrolaterally from the lateral lamina of the bodenaponeurosis and
extends anteriorly within the upper lip (Figs 5, 7-8), terminating laterally to
the premaxillary. Ventrolaterally to the orbit the integument turns under this
ligament to form the angle of the mouth.
A small horizontal bundle of muscle fibres is associated with the extra-
columella (Fig. 12). It lies in a somewhat similar position to the m. retractor
anguli oris of the amphisbaenids Amphisbaena and Leposternon as described
by Lakjer (1926). However, in Typhlosaurus aurantiacus these fibres extend
between the anteromedial surface of the extracolumella and the lateral lamina
of the bodenaponeurosis and they apparently function to draw the extra-
columella against the lateral surface of the adductor musculature.
The three major divisions of the adductor mandibulae group are readily
identified by virtue or their spatial relationship with the three rami of the
trigeminal nerve (Luther 1914). In Typhlosaurus aurantiacus the external adduc-
tor muscle mass lies laterally to the maxillary and mandibular rami (Figs 10-12),
the internal adductor lies medially to the maxillary but laterally to the ophthal-
mic rami (Fig. 9), and the adductor posterior lies laterally to the mandibular
ramus and ventrally to the external adductor (Fig. 12).
Musculus adductor mandibulae externus
The external adductor musculature arises within the temporal indentation
and its origin is bordered dorsally by a curved ridge on the parietal, extending
from the posterior tip of the postfrontal to the posterior extremity of the parietal
(Figs 9-14).
Three portions of the external adductor musculature—the superficialis,
medius and profundus—are present, either separated by tendinous laminae
or by differences in fibre orientation.
186 ANNALS OF THE SOUTH AFRICAN MUSEUM
The musculus adductor mandibulae externus superficialis (aes, Figs 9-11),
the most lateral portion of the external adductor, lies partially anterior to the
medius and profundus portions. Posteriorly, it is separated from them by, respec-
PAR imm
PF
RMA
oman
oO Nes
PAL TH)
= SS [a X/////| p
O as oo Wil Mle
quem
ee tae
———
me WN GZ \
\ WY
2
~
2
ME
Fig. 10. Cross-section of the skull just behind Figure 9.
CRANIAL AND CERVICAL MUSCLES OF TYPHLOSAURUS AURANTIACUS AURANTIACUS 187
tively, the lateral and the vertical laminae of the bodenaponeurosis. Anteriorly
the superficialis lies lateral to the pseudotemporalis muscle and is separated
from it by the maxillary ramus of the trigeminal nerve. Dorsal to the vertical
lamina of the bodenaponeurosis there is no distinction, other than fibre orien-
tation, between the superficialis and profundus portions. The superficialis,
however, is readily identified by the dorsoventral arrangement of its fibres as
opposed to the anteroventral orientation of the profundus fibres. The two
portions are by no means confluent since they separate easily during dissection.
The most substantial part of the superficialis lies dorsal to the coronoid process,
resulting in a near vertical fibre orientation relative to the long axis of the
lower jaw (Fig. 9).
The superficialis portion arises from the ventral surface of the postfrontal
(Fig. 9), from the dorsolateral ridge on the parietal (Fig. 11) and from the
lateral surface of the profundus portion. The origins are fleshy throughout.
Insertion is effected laterally on to the vertical and lateral laminae of the
bodenaponeurosis (Fig. 9), from the coronoid process posteriorly to a point
dorsal to and almost in line with the anterior tip of the extracolumella.
The superficialis portion is innervated by a posterolateral branch of the
mandibular ramus of V, which runs anterodorsally through the medius portion
to enter the medial surface of the superficialis portion via the medial lamina of
the bodenaponeurosis.
The musculus adductor mandibulae externus medius (aem, Figs 11-14)
lies posterolaterally to the pseudotemporalis muscle and between the boden-
aponeurosis and the quadrate tendon, with the vertical lamina of the latter
piercing it along the ventromedial border. It extends anteroventrally from the
posterolateral border of the parietal to the lower jaw and, being the most ventral
portion of the external adductor muscle, its dorsal border is wedged between the
lower extremities of the superficialis and profundus portions, separated from
them by, respectively, the lateral and medial laminae of the bodenaponeurosis.
Ventrally the muscle is forked, straddling the lower jaw from the coronoid
process to the anterior border of the posterior adductor muscle. At this point
there is, for a short distance, no partition between the fibres of the medius
portion of the external adductor and those of the posterior adductor (Fig. 11).
However, the medius portion is distinctly separated from the posterior adductor
by the lateral and medial laminae of the quadrate tendon over practically its
entire length. Posterior to the bodenaponeurosis and dorsal to the quadrate,
the medius fibres are continuous with those of the more medially situated
profundus portion of the external adductor (Fig. 12). It can, however, be
determined with reasonable accuracy that most of the fibres in this area belong
to the medius portion.
The medius portion arises fleshily from the posterolateral surface of the
parietal (Fig. 12), the lateral surfaces of the supratemporal, the squamosal,
the pro-otic dorsal to the quadrate and the lateral surface of the quadrate
above the extracolumella (Fig. 13).
188 ANNALS OF THE SOUTH AFRICAN MUSEUM
The insertion remains fleshy throughout and the fibres attach on to the
inferior surfaces of the medial and lateral laminae of the bodenaponeurosis, the
superior surfaces of the medial and lateral laminae of the quadrate tendon, the
medial and lateral surfaces of the vertical lamina of the quadrate tendon, and into
5500020)
CH 0,20 05
25066
Fig. 11. Cross-section of the skull at the level of the Gasserian ganglion.
CRANIAL AND CERVICAL MUSCLES OF TYPHLOSAURUS AURANTIACUS AURANTIACUS 189
the lateral groove on the coronoid (Fig. 10), with a few fibres attaching postero-
laterally on to the coronoid process of the dentary. Between the base of the
coronoid process and the foramen for the anterior supra-angular nerve, the
insertion utilizes the medial surface of the surangular, the dorsomedial surface
of the posterior process of the coronoid and a small dorsomedial area on the
prearticular, anterior to the adductor fossa. Dorsally the insertion continues
along the surangular ridge, anterior to the adductor fossa, as well as along
the dorso-lateral surface of the surangular and the dentary forming the
mandibular shelf. The diffuse nature of a small posterior part of the insertion
has been mentioned.
The medius portion is innervated by a posterolateral branch of the man-
dibular ramus of V piercing the muscle medially.
The musculus adductor mandibulae externus profundus (aep, Figs 11-12),
deepest portion of the external adductor, lies against the lateral cranial wall
laterally to the Gasserian ganglion and the maxillary ramus of the trigeminal
nerve, posterolaterally to the pseudotemporalis muscle, and medially to the
superficial and medius portions of the external adductor.
Extending anteroventrally, the profundus portion arises fleshily behind
the postfrontal, on the lateral surfaces of the parietal downgrowth and the
anterior superior process of the pro-otic (Fig. 11), as well as anterolaterally
on the pro-otic proper (Fig. 12). The area of origin is bounded dorsally by
the dorsolateral ridge of the parietal.
This muscle has no direct contact with the lower jaw and inserts fleshily
along the entire medial surface of the vertical lamina, and along the dorsal
half of the upper surface of the medial lamina of the bodenaponeurosis (Fig. 11).
Dorsally to the vertical lamina of the bodenaponeurosis no partition exists
between the superficial and profundus muscles (although the latter remains
discrete owing to the oblique orientation of its fibres, as opposed to the near
vertical orientation of the superficial fibres) (Fig. 11), whereas posterior to the
bodenaponeurosis no distinction is apparent between the profundus and medius
muscles (Fig. 12). However, the extent of each portion may be fairly easily
determined.
The profundus portion is innervated by a branch leaving the mandibular
ramus of V immediately below the Gasserian ganglion and turning dorsally
for a short distance to enter the muscle medially.
Musculus adductor mandibulae posterior
The posterior division of the adductor mandibulae group is present as a
single muscle, the musculus adductor mandibulae posterior (ap, Figs 11-13).
It extends anteroventrally from the quadrate to the mandible and lies laterally
to the mandibular ramus of the trigeminal nerve and medially to the extra-
columella (Fig. 12). It is straddled over its entire length by the medius portion
of the external adductor, although separated from it by the lateral and medial
laminae of the quadrate tendon (Fig. 11).
190 ANNALS OF THE SOUTH AFRICAN MUSEUM
AES
PO
Va
LO
LM
PO
EX
RM
Fig. 12. Cross-section of the skull at the level of the mandibular ramus of V.
The muscle arises fleshily from the slightly concave medial surface of the
quadrate, dorsomedially to and in line with the jaw articulation, as well as
dorsally to the posterior extremity of the protractor pterygoideus muscle and
the attachment of the quadrate ramus of the pterygoid (Fig. 13). The origin is
also ventrolateral to the pro-otic and anterior to the stapes, with a few of the
CRANIAL AND CERVICAL MUSCLES OF TYPHLOSAURUS AURANTIACUS AURANTIACUS 191
most dorsal fibres arising dorsally to the stapes (Fig. 14). Fibres also arise from
the anterior border of the quadrate. The quadrate tendon covers the posterior
adductor completely (Fig. 12) except at the anterior extremity of the muscle in
the region of the anterior supra-angular foramen where, over a short distance,
the fibres of the posterior adductor are continuous with those of the medius
portion of the external adductor. However, apart from obscuring the exact
anterior border of the insertion, this is of little importance since the muscles
are effectively separated, in practice, by the quadrate tendon. In addition, some
of the posterior adductor fibres arise from the inferior surfaces of the lateral
and medial laminae of the quadrate tendon.
The muscle inserts fleshily on to the mandible in an area extending from
the articular facet to the base of the coronoid process. Its posterior extremity
is pierced ventromedially by a small, robust tendon (Fig. 13), receiving the
VIO
EX
Fig. 13. Cross-section of the skull at the level of the quadrate.
192 ANNALS OF THE SOUTH AFRICAN MUSEUM
insertion of the fibres arising in line with the jaw articulation on the medial
surface of the quadrate. This tendon extends anteriorly for a short distance to
attach immediately anterior to the articular facet on the dorsomedial surface
of the articular process. The rest of the fibres straddle the mandible and insert
dorsomedially on the surangular, within and around the adductor fossa (Fig. 11)
on that part of the prearticular forming the ventral rim of the adductor fossa,
and anterior to the fossa on the dorsomedial surface of the prearticular. A few
fibres attach dorsomedially to the posterior process of the coronoid. In addition,
fibres of this part of the muscle also insert on a small, vertical tendon running
dorsally along the surangular ridge and situated within the muscle itself (Fig. 11).
The tendon extends from the anterior border of the muscle to a point in line
with the anterior border of the mandibular ramus of V, and fibres insert along
its lateral and medial surfaces. The lateral fibres of the muscle insert on the
mandibular shelf (Fig. 11), attaching dorsolaterally on to the surangular and
dentary from the articular facet to near the base of the coronoid process.
A posterolateral branch of the mandibular ramus of V ramifies within
the medius portion of the external adductor, with one branch piercing the
medial lamina of the quadrate tendon to innervate the posterior adductor.
Musculus adductor mandibulae internus
According to Lakjer (1926) the third main division of the adductor man-
dibulae group, the internal adductor musculature, consists of two separate
muscles, the m. pseudotemporalis and m. pterygoideus. Both muscles are
present in Typhlosaurus aurantiacus as well-defined groups of fibres situated at
right angles to one another.
The musculus adductor mandibulae internus pseudotemporalis (p, Figs
9-11) is a single, dorsoventral group of fibres located anteriorly within the
temporal indentation, and although the bulk of its fibres are concentrated
dorsomedially to the coronoid process, the muscle extends posteriorly to the
anterior border of the mandibular ramus of V. As required by the classical
definition it is situated medially to the maxillary, laterally to the ophthalmic
and anteriorly to the mandibular rami of V. It lies medially to and against
the superficial, anteromedially to the medius and anteriorly to the profundus
portions of the external adductors; anteromedially to the posterior adductor
muscle; laterally to the levator pterygoideus muscle and the epipterygoid bone,
and anterolaterally to the pterygoideus muscle.
It arises fleshily from the anterolateral surfaces of the parietal downgrowth
and the anterior superior process of the pro-otic, anterior to the origin of the pro-
fundus portion of the external adductor as well as from the lateral surface of
the pro-otic membrane and the dorsal part of the epipterygoid bone.
The muscle inserts on the medial surface of the coronoid bone, utilizing
the medial coronoid groove which extends from the apex of the coronoid
process to the medial surface of the posterior process of the coronoid bone
(Fig. 10). A small tendon attaching to the coronoid bone extends along the
CRANIAL AND CERVICAL MUSCLES OF TYPHLOSAURUS AURANTIACUS AURANTIACUS 193
lower rim of the medial coronoid groove and underlies the pseudotemporalis
muscle, receiving the insertion of its medial fibres (Fig. 9). Behind the coronoid
bone a few fibres insert on to the medial surface of the prearticular bone
(Fig. 11).
The muscle is innervated by a branch of the mandibular ramus of V which
leaves the anterior border of the ramus directly below the Gasserian ganglion
and courses anteriorly for a short distance to enter the muscle posterodorsally.
The musculus adductor mandibulae internus pterygoideus (pts), is the
largest single muscle in the head of Typhlosaurus aurantiacus and extends from
the infra-orbital fenestra to the posterior extremity of the mandible. It has a
small head which expands posteriorly to form a large rounded masticatory
cushion, appropriately termed ‘Kauwulst’ by Lakjer (1926). The ‘Kauwulst’
is the most prominent part of the muscle and, with its opposite number, limits
the aperture of the throat. It lies deep to the intermandibularis posterior muscle
and the mandibulo-hyoid musculature, medially to the insertional tendon of
the cervicomandibularis muscle and laterally to the protractor pterygoidei
muscle and the pterygoid bone.
The muscle arises mainly from the outer surface of the pterygoid bone
(Fig. 9) and from the inner surface of an extensive tendon originating lateral
to the infra-orbital fenestra. Anteriorly the muscle is divided into two short
slips, one arising dorsally from both the posterolateral surface of the palatine
and the anterolateral border of the pterygoid and the second one arising ven-
trally on the posterolateral border of the palatine and the anterolateral surface
of the pterygoid. The two muscle slips become confluent at a point in line with
the anterior border of the levator pterygoideus muscle.
Some fibres arise fleshily along the outer lateral and ventral surface of
the gutter-shaped pterygoid bone (Fig. 11). The major part of the origin,
however, is from the exceptionally strong tendon arising lateral to the infra-
orbital fenestra on the ventral surface of the palatal complex. The tendon arises
behind the ventrolateral process of the maxillary on the ventral surface of the
ectopterygoid, the posteroventral surface of the palatine and on the ventral
surface of the pterygoid in front of the basipterygoid recess. Anteriorly the
tendon covers the origin of the ventral slip of muscle and forms a thick pad
medial to the angle of the mouth (Fig. 9). The pad presumably protects the
fibres from damage by deflecting the coronoid process laterally during adduction
of the jaw. Posteriorly the tendon broadens to cover the anterior two-thirds
of the ventral surface of the muscle. Most of the fibres of the pterygoideus
muscle arise from the inner surface of this tendon.
The muscle inserts on the retroarticular process of the mandible (Fig. 13).
In front of the process the ventral border of the mandible is concave to accom-
modate the lateral fibres of the muscle which wrap around the jaw behind the
posteroventral extremity of the dentary. The muscle envelops the retroarticular
process, with fibres inserting on to its medial, ventral and lateral surfaces.
Medial to the jaw a ribbon-like tendon lies within the muscle and attaches
194 ANNALS OF THE SOUTH AFRICAN MUSEUM
ventrally to the retro-articular process (Fig. 13). Its dorsal and ventral surfaces
are utilized for insertion.
The muscle is innervated by a branch of V which leaves the mandibular
ramus medially, in line with the posterior supra-angular foramen, to pierce
the muscle laterally.
Intermandibular musculature
The intermandibular muscles, innervated by V, form part of the trigeminal
musculature. Situated anteroventrally to the constrictor colli, their fibres
extend transversely between the rami of the lower jaws as thin superficial sheets
of muscle, deep only to the skin and to the insertional tendon of the cervico-
mandibularis muscle. Two portions, an anterior and a posterior, are present.
The m. intermandibularis anterior (ia, Figs 9-10), four to six fibres in
thickness, lies anteriorly between the rami of the lower jaws. It is laterally
interrupted at right angles by bundles of the geniohyoideus muscle which insert
anteroventrally on the jaw. According to Camp (1923) these interdigitations
(of which there are five in Typhlosaurus aurantiacus) are always present in
lizards. The muscle lies superficially to the geniohyoideus and genioglossus
muscles and can be divided into a more posteriorly lying superficial portion
and a more anteriorly lying profundus portion.
The superficial portion arises medially on the jaw (Fig. 10) between the
posterior mylohyoid foramen and the combined foramen for the anterior
mylohyoid and the infra-alveolar nerves, along the medial surface of the pre-
articular, splenial and dentary bones. It inserts anteriorly along the ventral
midline onto the sheet of fascia receiving the insertion of the constrictor colli
muscle. Three bundles of geniohyoideus fibres interdigitate with the superficial
portion of the intermandibularis anterior muscle.
The profundus portion of the m. intermandibularis anterior lies imme-
diately anterior to the superficial portion, posteriorly overlain by the latter.
It arises medially to the jaw from the dorsolateral surface of the sublingual
gland and the outer surface of the buccal lining, dorsal to the gland. The origin
extends from a point in line with the anterior mylohyoid and the infra-alveolar
foramen to just behind the jaw symphysis. The muscle inserts tendinously
in the ventral midline anterior to the superficial portion of the intermandibularis
anterior muscle. Two bundles of the geniohyoideus fibres interdigitate with the
profundus portion.
The intermandibularis anterior is innervated by a branch of the posterior
mylohyoid nerve, which enters the superficial portion posteriorly, and by a
branch of the anterior mylohyoid nerve, which enters the profundus portion
superficially.
The m. intermandibularis posterior (ip, Fig. 12), two fibres in thickness,
lies immediately behind the intermandibularis anterior. The two muscles are
separated by the most posterior interdigitation between the geniohyoideus
and intermandibularis anterior muscles. The intermandibularis posterior does
CRANIAL AND CERVICAL MUSCLES OF TYPHLOSAURUS AURANTIACUS AURANTIACUS 195
not interdigitate with the geniohyoideus muscle and is well separated from the
constrictor colli. It covers the throat ventrally, lateral to the midline, and lies
superficially to the hyoglossus, the genioglossus and the geniohyoideus muscles,
the masticatory cushion (Kauwulst) of the pterygoideus muscle and the hypo-
glossal nerve (Fig. 12).
The m. intermandibularis posterior arises tendinously from a thin sheet
of fascia which covers the pterygoideus muscle laterally (Fig. 12) and attaches
to the lateral surfaces of the surangular and dentary bones between the jaw
articulation and the posterior mylohyoid foramen. The muscle inserts ten-
dinously in the ventral midline on to the same sheet of fascia which receives
the insertion of the constrictor colli muscle and the intermandibularis anterior
muscle (Fig. 12).
It is innervated by a branch of the posterior mylohyoid nerve, entering the
muscle superficially via the posterior mylohyoid foramen.
Tongue musculature
In Typhlosaurus aurantiacus the tongue is of the usual scincid type, bluntly
triangular with a bifurcate apex and posteriorly divided into two roots situated
lateral to the glottis. Behind the apex its dorsal surface is covered with scale-
like papillae, on which glandular surfaces are restricted to the basal portions.
According to Camp (1923) the position of the glandular surfaces is a diagnostic
feature of the Scincomorpha.
The tongue is composed of fibres of both extrinsic and intrinsic muscula-
ture. The intrinsic fibres control the shape of the tongue, while motion is
controlled by the extrinsic fibres. Sondhi (1958) found that in some Indian
reptiles the apparently distinct groups of intrinsic fibres are actually parts of
the hyoglossus muscle and do not deserve independent status. In Typhlosaurus
aurantiacus these fibre groups are equally distinct and in view of the marked
differences between their orientation and that of the hyoglossus muscle, they
will be described separately.
Extrinsic muscles
The m. hyoglossus (hg, Figs 9-12) is a paired longitudinal muscle, extending
parallel to the midline from the hyoid apparatus to the anterior tip of the tongue.
Posteriorly the muscle is dorsoventrally flattened and lies ventrolaterally to the
trachea and the oesophagus, laterally to the first ceratobranchial and dorso-
laterally to the posterior part of the geniohyoideus muscle. Anteriorly it becomes
cylindrical, turning dorsally to the undersurface of the tongue to lie laterally
to the lingual process of the hyoid, medially to the geniohyoideus muscle,
and ventrolaterally to the glottis. It is sheathed by fibres of the vertical intrinsic
musculature. The most anterior part of the muscle lies dorsomedially to the
genioglossus muscle, and tapers towards the jaw symphysis to terminate ven-
trally to the anterior tip of the tongue.
196 ANNALS OF THE SOUTH AFRICAN MUSEUM
The muscle arises superficially in the throat region from the anterior
surface of the first ceratobranchial, medially to the origin of the geniohyoideus
lateralis muscle, ventrolaterally to the oesophagus and dorsolaterally to the
origin of the geniohyoideus medialis muscle. It inserts fleshily along the ventral
surface of the tongue, between the medial and lateral vertical fibres of the
intrinsic musculature and medially to the insertion of the genioglossus muscle.
The muscle is innervated by a branch of the hypoglossal nerve.
The m. genioglossus (ggl, Figs 9-12) is a paired muscle, lying laterally to
the ventral midline and extending from the jaw symphysis to the posterior
border of the tongue. In front it lies dorsolaterally to the geniohyoideus medialis
muscle, ventrolaterally to the hyoglossus muscle, and medially to the sublingual
gland. At the back the muscle extends laterally around the ventral surface of
the sublingual gland and behind it the fibres lie ventrally to the lateral margin
of the oral membrane. A small cylindrical group of fibres separates antero-
ventrally from the genioglossus muscle and extends posteriorly, lying ventro-
medially to the genioglossus muscle and dorsally to the geniohyoideus medialis
muscle. In line with the glottis the fibres of this bundle become confluent with
those of the geniohyoideus medialis muscle. It is innervated by a minute ramus
branching from the hypoglossal nerve.
The genioglossus arises tendinously at the jaw symphysis, taking origin
from the inner ventral surface of the dentary immediately lateral to the sym-
physis, from the connective tissue surrounding the symphysis and, in the mid-
line, from the sheet of fascia covering the throat musculature ventrally. It inserts
along the ventral surface of the tongue, laterally to the hyoglossus muscle
and the lateral group of vertical intrinsic fibres, and interlaces with the transverse
fibres of the intrinsic musculature. The muscle is innervated by a branch of the
hypoglossal nerve.
Intrinsic musculature
The intrinsic muscles are innervated by the hypoglossal nerve and consist
of three groups of fibres, i.e. the vertical lingual, the longitudinal lingual and
the transverse lingual fibres, and although they interweave to some extent each
group remains distinct.
The m. verticalis linguae (vl, Figs 9-11) consists of a superior and an
inferior group of fibres. The superior fibres lie dorsally to the transverse lingual
muscle and form the papillae of the tongue. The inferior fibres lie ventrally
to the transverse lingual muscle, interlace with its fibres, and extend from the
apex of the tongue to the glottis. These fibres consist of a medial and a lateral
group. The lateral group lies between the genioglossus and the hyoglossus
muscles while the medial group lies between the hyoglossus muscle and the mid-
line. The two groups meet tendinously ventral to the hyoglossus muscle.
The m. transversalis linguae (tl, Figs 9-11) lies between the inferior and
superior vertical fibres. Its fibres extend across the width of the tongue from
the apex to the roots and interlace with the fibres of the inferior vertical, the
CRANIAL AND CERVICAL MUSCLES OF TYPHLOSAURUS AURANTIACUS AURANTIACUS 197
hyoglossus and the genioglossus muscles.
The m. longitudinalis linguae (ll, Figs 9-10) lies ventrally along the rim
of the tongue, laterally to the insertion of the genioglossus muscle on either
side. It extends from behind the apex of the tongue to nearly in line with the
glottis as a single cylindrical bundle of fibres, breaking up into smaller bundles
posteriorly and becoming obscured by the insertion of the genioglossus muscle.
The m. geniohyoideus lateralis (ghl, Fig. 12) is a narrow muscle extending
between the ventral surface of the jaw and the hyoid apparatus. It lies ventro-
medially to the jaw, ventrolaterally to the genioglossus muscle, deep to the
intermandibularis posterior muscle, and ventrally to the ‘Kauwulst’ of the
pterygoideus muscle.
It arises fleshily on the ventral surface of the jaw from a short distance
behind the symphysis to the posterior mylohyoid foramen. The origin consists
of five successive bundles of longitudinal fibres interlacing with the fibres of
the intermandibularis anterior muscle. The muscle inserts on the anterolateral
tip of the sole remaining posterior prong of the hyoid apparatus (usually
taken to be the first ceratobranchial). The insertion is fleshy and lies lateral
to that of the geniohyoideus medialis and hyoglossus muscles. The muscle is
innervated by a branch of the glossopharyngeal nerve.
The m. geniohyoideus medialis (ghm, Figs 9-12) is a superficial group
of fibres extending from the anteroventral surface of the genioglossus muscle
to the hyoid apparatus. The muscle lies laterally to the ventral midline, deep
to the intermandibularis musculature, ventromedially to the genioglossus
muscle and below the bundle of fibres extending between it and the genio-
glossus muscle. Posteriorly the muscle is flattened dorsoventrally and lies
ventromedially to the hyoglossus muscle, deep to the intermandibularis
posterior muscle and medially to the hypoglossal nerve. Anteriorly the muscle
becomes triangular in cross section and lies deep to the intermandibularis
anterior muscle.
The muscle arises tendinously behind the origin of the genioglossus
muscle, from the deep surface of the sheet of fascia covering the throat muscula-
ture ventrally and inserts fleshily along the ventral surface of the first cerato-
branchial, ventromedially to the origin of the hyoglossus muscle. The muscle
is innervated by a branch of the glossopharyngeal nerve.
Depressor mandibulae group
The m. depressor mandibulae (Figs 14-16) is a superficial sheet of fibres
behind the adductor musculature. It lies laterally to the stapedial artery, the
lateral head vein and the hyomandibular ramus of VII. On its ventrolateral
surface it is obliquely overlain by the prominent cervicomandibularis muscle
which in its turn is covered by the dorsoventral fibres of the sheet-like constrictor
colli muscle. There is no indication in either Typh/osaurus aurantiacus or Acontias
meleagris that the depressor mandibulae and cervicomandibularis muscles
are continuous.
198 ANNALS OF THE SOUTH AFRICAN MUSEUM
PAR
ay
o
o>
o
5
[wee
oo
18)
Ooyd
of
~
GHL
Fig. 14. Cross-section of the skull at the level of the stapes.
The depressor mandibulae consists of an anterior portion arising cranially
and & posterior portion arising cervically. The anterior portion (dma, Figs 7,
14-15) is spindle-shaped and lies immediately behind the stapes. It arises fleshily
from the anterolateral surface of the fused exoccipital-opisthotic bone, postero-
dorsally to the quadrate head, and extends vertically downwards, laterally to
the posterior half of the stapedial footplate, to insert fleshily on the dorsal
CRANIAL AND CERVICAL MUSCLES OF TYPHLOSAURUS AURANTIACUS AURANTIACUS 199
Fig. 15. Cross-section of the skull at the level of the anterior portion of the depressor
mandibulae.
surface of the retro-articular process behind the tendon connecting the stapes to
the mandible.
The posterior portion (dmp, Fig. 16), a thin triangular sheet of fibres
lateral to the obliquus capitis magnus muscle and the innervations of the longis-
200 ANNALS OF THE SOUTH AFRICAN MUSEUM
simus and episternocleidomastoideus muscles, lies against the posterior border
of the cranial portion of the depressor mandibulae. It arises along the insertional
tendon of the longissimus cervicis muscle, ventrolateral to the spinalis capitis
LV
AT
Fig. 16. Cross-section of the skull at the level of the posterior portion of the depressor
mandibulae.
CRANIAL AND CERVICAL MUSCLES OF TYPHLOSAURUS AURANTIACUS AURANTIACUS 201
muscle and between the origin of the cervicomandibularis muscle and the
posterodorsal border of the adductor mandibulae musculature. The fibres
converge anteroventrally to insert behind those of the cranial portion by means
of a small tendon on to the posterodorsal extremity of the retro-articular process
of the mandible.
The depressor mandibulae is innervated by the hyoid ramus of VII, which
branches off behind the stapedial footplate from the hyomandibular ramus,
pierces the medial surface of the muscle and courses anteriorly for a short
distance within it.
The m. cervicomandibularis (cm, Figs 6-7, 16) is a large elongated sheet
of fibres medial to the constrictor colli muscle. It extends antero-ventrally,
superficial to the cervical musculature, from the level of the seventh dorsal
vertebra to the anteroventral surface of the jaw. It lies lateral to the episterno-
cleidomastoideus muscle, the longissimus cervicis and capitis muscles, and the
anterior paris of the iliocostalis system and the oblique hypaxial muscles.
The muscle arises fleshily from the anterolateral surface of the iliocostalis
system and the oblique hypaxial muscles, from the longissimus cervicis muscle,
and along its border from the dorsal intermuscular septum. Its fibres extend
forward only as far as the posterolateral surface of the ‘Kauwulst’ of the ptery-
goideus muscle where they attach to a thin sheet of fascia which covers the
‘Kauwulst’ laterally and ventrally and lies superficial to the throat muscles.
Anteriorly the tendon becomes aponeurotic and inserts along the ventral
surface of the jaw, from the anterior border of the posterior intermandibular
muscle to just lateral to the symphysis.
The m. cervicomandibularis is innervated by a branch of VII which pierces
the medial surface of the muscle in line with the occipital condyle of the
skull.
The m. constrictor colli (cc, Figs 5, 15-16) is a thin superficial sheet of
fibres covering the cervical region immediately below the skin. Its fibres extend
anteroventrally from the cervical region to cover the pterygoideus muscle
posterolaterally and the cervicomandibularis muscle anteroventrally. The
muscle is triangular in lateral aspect and its anterior border lies immediately
behind the extracolumella. The upper border of the muscle tapers postero-
ventrally and it terminates at a point ventrolateral to and approximately in
line with the third vertebra.
The muscle arises fleshily from a thin superficial sheet of cervical fascia
extending ventrally from the dorsal intermuscular septum, situated between the
transversospinalis and longissimus systems, to insert ventrolaterally in the
cervical region on to a superficial sheet of fascia covering the throat from the
jaw symphysis posteriorly to the rectus abdominis muscles.
The muscle is innervated by a branch of the hyomandibular ramus of VII
which passes through the cervicomandibularis muscle to enter the deep surface
of the constrictor colli in line with the posterior border of the condyle of the
skull.
202 ANNALS OF THE SOUTH AFRICAN MUSEUM
Cervical musculature
In contrast to non-fossorial lizards, the cervical extensor and flexor muscles
in Typhlosaurus aurantiacus play an active role during locomotion and especially
during burrowing movements. However, in spite of this added function the
distribution of the individual muscles retains a pattern common to saurians in
general.
The nuchal ligament is a vertical mid-dorsal sheet in the cervical region
extending from the occiput to the neural spine of the axis. Dorsally the ligament
is continuous with the fascia covering the transversospinalis system. This
fascia is laterally continuous with the dorsal intermuscular septum lying between
the transversosponalis and longissimus systems.
The m. spinalis capitis (sc, Figs 6, 12, 14-16) is a flat, dorsally situated
muscle extending immediately below the skin from the level of the twelfth
vertebra to the occiput. It lies lateral to the nuchal ligament and dorso-medial
to the anterior part of the longissimus dorsi muscle and its cervical derivative,
the longissimus cervicis. The muscle covers the rectus capitis anterior and
obliquus capitis magnus muscles. The lateral fibres arise fleshily from the
lateral margin of the dorsal intermuscular system.
The fibres insert tendinously on to the posterodorsal surface of the parietal
bone. The insertion is confined to a shallow depression on the parietal anterior
to the supra-occipital-parietal suture and lateral to the small mid-dorsal ridge
formed by the ascending process of the tectum synoticum.
The muscle is innervated by a branch of the dorsal ramus of the first
spinal nerve which passes between the rectus capitis posterior and the obliquus
capitis magnus muscles.
The m. rectus capitis posterior and the m. obliquus capitis magnus are
partially fused and appear as a single group of fibres. However, in the interests
of clarity they will be described separately.
The m. rectus capitis posterior (rp, Figs 6, 8) extends from the atlas to
the occiput. It lies ventral to the spinalis capitis muscle, lateral to the nuchal
ligament, dorso-medial to the longissimus cervicis muscle and antero-medial
to the obliquus capitis magnus muscle, which arises posterior to it. In Typhlo-
saurus aurantiacus the rectus capitis posterior is a single muscle and it probably
represents the fused rectus capitis superficialis and profundus muscles still
present, although partially fused, in such forms as Iguana iguana (Olson 1936).
The rectus capitis posterior muscle can be distinguished from the obliquus
capitis magnus muscle by the passage of the dorsal ramus of the first spinal
nerve between them as in Ctenosaura pectinata (Oelrich 1956).
The muscle arises fleshily from the lateral surface of the axial neural arch
and the aponeurotic fascia covering the atlanto-occipital gap. Some fibres also
arise from the dorsal surface of the obliquus capitis magnus muscle.
The fibres insert fleshily, deep to the spinalis capitis muscle and dorso-
medial to the insertion of the obliquus capitis magnus muscle, on to the dorsal
surface of the supraoccipital bone and that part of the fused exoccipital-
CRANIAL AND CERVICAL MUSCLES OF TYPHLOSAURUS AURANTIACUS AURANTIACUS 203
opisthotic bone dorsal to the ridge formed by the lateral semicircular canal
of the internal ear.
The muscle is innervated by a branch of the dorsal ramus of the first
spinal nerve.
The m. obliquus capitis magnus (oc, Figs 6, 8) extends from the fifth
vertebra to the lateral margin of the fused exoccipital-opisthotic bone. It lies
ventral to the spinalis capitis muscle, posterolateral to the rectus capitis posterior
muscle, dorsal to the anterior part of the spinalis dorsi muscle, dorsal to the
rectus capitis anterior muscle and medial to the longissimus cervicus muscle.
It arises fleshily from the dorsal extremities of the neural spines of the
third, fourth and fifth vertebrae, the tendinous tissue connecting the spines
and from the fascia covering the spinalis dorsi muscle.
The fibres extend obliquely forward and insert tendinously on to the
lateral margin of the fused exoccipital-opisthotic bone, attaching to the ridge
formed by the lateral semicircular canal of the internal ear, ventrolateral
to the rectus capitis posterior muscle and dorsomedial to the insertion of the
longissimus cervicis muscle.
The muscle is innervated by the dorsal ramus of the first spinal nerve.
The m. longissimus dorsi lies ventrolateral to the transversospinalis system
and dorsal to the heads of the ribs. At the level of the fourth vertebra it divides
into a dorsal and a ventral group of fibres. The dorsal group, the m. longissimus
cervicis (Ice, Figs 6-8) lies ventrolateral to the spinalis capitis and rectus capitis
posterior muscles and lateral to the obliquus capitis magnus muscle. The
muscle arises fleshily approximately from the level of the seventh to the second
vertebrae from the fibres of the longissimus dorsi muscle. It extends anteriorly
dorsal to the longissimus capitis muscle and lateral to the obliquus capitis
magnus muscle, to insert tendinously on to the lateral margin of the fused
exoccipital-opisthotic bone, lateral to the insertion of the obliquus capitis
magnus muscle and medial to the insertion of the episternocleidmastoideus
muscle.
The muscle is innervated by the dorsal ramus of the first spinal nerve.
The ventral group of fibres, the m. longissimus capitis (Ica, Fig. 8), extends
anteroventrally from the level of the fourth vertebra to the basal tuberosity
of the basi-occipital bone. It lies ventral to the longissimus cervicus muscle,
medial to the episternocleidomastoideus muscle, dorsolateral to the longus
colli muscle and lateral to the rectus capitis anterior muscle. The fibres arise
fleshily from the longissimus dorsi muscle and the synapophysis of the first
four vertebrae to insert tendinously on to the basal tuberosity lateral to insertion
of the rectus capitis anterior muscle and dorsomedial to the insertion of the
longus colli muscle.
The muscle is innervated by a branch of the dorsal ramus of the first
spinal nerve.
The m. rectus capitis anterior (ra, Fig. 8) is a group of short fibres extending
from the axis to the occiput. The muscle lies ventrolateral to the axis, the
204 ANNALS OF THE SOUTH AFRICAN MUSEUM
atlas and the occipital condyle of the skull; medial to the longissimus capitis
muscle and dorsal to the anterior part of the longus colli muscle with which
some of its ventral fibres are confluent.
It arises fleshily on the ventrolateral surface of the axis medial to the
synapophysis and ventrolaterally on the atlas. The fibres insert fleshily on to
the basioccipital and exoccipital-opisthotic bones. The insertion lies ventral
to the insertion of the rectus capitis posterior muscle, dorsal to the insertion
of the longus colli muscle and dorsomedial to the insertion of the longissimus
capitis muscle.
The muscle is innervated by a branch of the first spinal nerve.
Fig. 17. Camera lucida drawing of the left lower jaw; lingual view.
DISCUSSION AND CONCLUSIONS
General
The morphology of fossorial skinks in general is paralleled by that of
snakes in many ways and it is therefore interesting to note that according to the
theory of Walls (1942) snakes originated as fossorial forms. Any evaluation of
the musculature of Typhlosaurus aurantiacus must be made in the light of the
fact that the animal leads a predominantly subterranean existence which does
not involve the construction of burrows. Huey et al. (1974) describes Typhlo-
saurus as a sand swimming lizard which normally moves in a lateral sinuous
path beneath the sand. 7. aurantiacus has a subterminal mouth as in other
fossorial lizards, e.g. Acontias. The position of the mouth prevents soil particles
from entering the buccal cavity during burrowing movements. De Weerdt
(1971) indicates that it is important to a fossorial lizard like Dibamus to use
its mouth in a terminal position. This argument is based on the assumption
that the lizard lives in a burrow and encounters its prey directly in front of it.
Neither Typhlosaurus nor probably Dibamus lives in a burrow and conse-
quently prey may be approached from any angle.
The animal uses its head as a burrowing tool, and the skull and its associ-
ated musculature are consequently strongly modified. Temporal arches and
posttemporal fenestrae are absent. The elongated temporal region is strengthened
by the lateral downgrowth of the parietal and the broad anterior superior
processes of the pro-otic. These structures also serve as areas of origin to the
CRANIAL AND CERVICAL MUSCLES OF TYPHLOSAURUS AURANTIACUS AURANTIACUS 205
adductor musculature. The occipital region is strengthened by the fused exoccipi-
tal and ophisthotic bones and the expanded nature of the occiput allows the
cervical musculature to insert relatively far forward on to the skull.
As in Dibamus (Gasc 1968) Typhlosaurus aurantiacus lacks a m. levator
bulbi. This condition is probably associated with the degeneration of the
eyes. According to Haas (1973) the structural diversity of the m. levator bulbi
in lizards appears not to be controlled by phylogenetic factors but rather by
functional factors such as the presence or absence of a movable lower eyelid
or a general reduction of the visual apparatus.
The m. pseudotemporalis is single in Typhlosaurus aurantiacus as in the
Gekkonidae, the Pygopodidae and the snakes. De Weerdt (1971) describes a
single m. pseudotemporalis in Dibamus but fails to define the position of the
muscle in relation to the maxillary ramus of V. It may well be that the muscle is
in fact part of the m. adductor mandibularis externus. Gasc (1968) describes
am. pseudotemporalis in Dibamus consisting of two parts. Haas (1973) interprets
Gasc’s description as pertaining only to the posterior of the two portions.
However, Gasc, on page 135, clearly states that: ‘Une nappe profonde (fig. 9)
formée par deux chefs. . . . Cette nappe pourrait correspondre, d’aprés ses
insertions, aux deux chefs du m. adductor mandibularis medius (= pseudo-
temporalis); toutefois, la branche maxillaire du trijumeau passe ici au-dessous
de ce plan musculaire.’ From Gasc’s figure 9 it is clear that the maxillary ramus
of V lies medial to the m. pseudotemporalis. It is probable therefore that this
muscle forms part of the m. adductor mandibularis externus and not the m.
adductor mandibularis internus. According to Haas (1973) the gekkonids and
pygopodids lack the m. pseudotemporalis superficialis but retain the profundus
portion of the muscle which consists of an anterior and a posterior part. It
is probable that the single muscle retained in 7. aurantiacus represents the
profundus portion of the m. pseudotemporalis and that the reduction of the
m. pseudotemporalis in this animal is related to the loss of the temporal arches
as Haas (1973) believes it to be in the case of gekkonids, pygopodids and snakes.
The depressor mandibulae is a relatively small muscle and (in theory)
its position close to the fulcrum of the jaw is not functionally optimal. Since
the opening of the jaw is usually assisted by gravity this condition is not a
liability. However, in a lizard which feeds subterraneously the surrounding
pressure of the soil may demand a more sophisticated arrangement of the jaw
opening muscles. In Typhlosaurus aurantiacus the cervicomandibularis is
probably the main jaw opening muscle since its origin on the neck musculature
and ventral insertion on the jaw makes it ideally suited for this purpose. Camp
(1923) notes that the cervicomandibularis muscle is enormously developed in
all burrowers.
From the forwardly extended insertion of the cervical musculature on to
the occiput and the lateral extension of the synapophyses of the axis and the
three following vertebrae it is evident that this group of muscles plays an
important role in locomotion. It serves to flex and extend the skull as well as
206 ANNALS OF THE SOUTH AFRICAN MUSEUM
stiffen the atlanto-occipital joint during burrowing. The degree of fusion
between individual cervical muscles seems logical in the light of their function.
Cranial kinesis
Versluys (1910, 1912) described the movable joints of the reptilian skull.
He divided the skull into an ‘occipital segment’ consisting of the bones of the
braincase and the parasphenoid, and a ‘maxillary segment’ comprising the rest
of the skull. The intracranial movements between the two segments are known
as kinesis and were interpreted by Versluys as a mechanism for increasing the
gape of the mouth by lifting the snout. Various degrees of kineticism exist,
and skulls ranging from akinetic, with little or no movement between the
‘segments’, to amphikinetic, in which more than two movable parts are found.
The constrictor dorsalis group of muscles is responsible for the kinetic move-
ments of the skull.
Versluys based his conclusions on morphological studies, but recent workers
have made use of sophisticated methods to study live material. Frazetta (1962)
who revised Versluys’s terminology, used motion pictures to record the capture
of prey as well as electrical stimulation and biomechanical analysis of the
muscles. Iordansky (1970) used biomechanical analysis to extend the approach
of Frazetta. In contrast to Versluys, Frazetta concluded that kinesis actually
lessens oral gape. However, in spite of the advanced techniques employed by
them, Frazetta and Iordansky are not in full agreement on certain aspects of
kinesis.
It is evident that kinesis is a complex mechanism of which the functional
significance is not yet fully explained. This is also clear from the variable nature
of the constrictor dorsalis group of muscles in a form such as Sphenodon.
Frazetta (1962) describes Sphenodon as akinetic whereas Ostrom (1962) describes
a specimen which has both a levator pterygoidei and a protractor pterygoidei
muscle. According to Ostrom, in previously described specimens of Sphenodon
either one of these muscles were present but never both. The study of cranial
kinesis therefore requires the application of sophisticated techniques to live
specimens as well as the dissections of numerous examples of the same species.
The scope of this paper and the paucity of material precludes an in-depth
study of cranial kinesis in Typhlosaurus aurantiacus but, since the cranial
muscles have been described in detail, a brief summary will be given here.
According to Bellairs (1969) fossorial lizards tend to become monokinetic
or even akinetic. A reduction of intracranial movements seems logical in the
light of strengthening the skull for burrowing. Usually it appears that the
metakinetic bending plane between the supraoccipital and parietal bones is
reduced in favour of the mesokinetic bending plane between the parietal and
frontal bones as in Acontias (De Villiers 1939, Brock 1941, Van der Merwe
1944); Monopeltis capensis (Kritzinger 1946); Anniella (Toerien 1950, Bellairs
1969): Nessia (Bellairs 1969) and Dibamus (De Weerdt 1971). According to
Leonard (1973) the less specialized burrowing skink Scelotes is amphikinetic.
CRANIAL AND CERVICAL MUSCLES OF TYPHLOSAURUS AURANTIACUS AURANTIACUS 207
In Typhlosaurus the presence of a small levator pterygoidei muscle, a
substantial protractor pterygoidei muscle, a synovial articulation between
the quadrate and the skull and a movable basipterygoid articulation clearly
indicate a certain amount of intracranial movement. Because of the posterior
expansion of the skull, the occipital bones have become fused and the insertional
areas of the cervical musculature have increased to such an extent that the
spinalis capitis muscle inserts dorsally on the posterior part of the parietal,
anterior to the position of the metakinetic bending plane.
The position of this muscle suggests a sharp reduction, if not total absence,
of a functional metakinetic bending plane, despite the presence of the carti-
laginous ascending process of the tectum synoticum. The post-orbital bar is
incomplete and according to Leonard (1973) this is a prerequisite for meso-
kinesis. Smit (1964) agrees that metakinesis is reduced or absent and states
that movement is clearly possible between the parietal and the frontal bones,
indicating that the Typhlosaurus skull is definitely mesokinetic.
Jaw mechanics
According to Ostrom (1964) the vertebrate lower jaw operates as a lever
of the third class during adduction. This arrangement ensures maximum
depression of the jaws with a minimum length of adductor muscle fibres.
In a system of this kind the mechanical advantage is directly proportional
to the length of the moment arm if the applied force (adductor musculature)
remains constant. The moment arm is defined as the perpendicular distance
between the line of applied force and the fulcrum (Fig. 18A). In the jaw the
moment arm represents the distance between the jaw articulation and the tip
of the coronoid process.
If the line of applied force functions in a posterior direction, at an angle
of less than ninety degrees to the long axis of the lever, the moment arm FB
(Fig. 18A) is no longer perpendicular to the applied force and is effectively
displaced to position FB’ (Fig. 18B) with the result that the length of the moment
arm is decreased and the system functions at a disadvantage. However, this
condition can be overcome by raising the point of attachment (development
of a coronoid process) of the applied force (Fig. 18C).
From Figure 18C it follows that: m? = x? + y?.
The moment arm (m) is therefore a function of x (the distance between
the coronoid process and the jaw articulation) and y (the height of the coronoid
process). If y is constant x will determine the line of muscle action (@) and vice
versa, because tan 6 = >.
Consequently, with y constant any decrease in x will result in a more
posteriorly directed line of muscle action, or, alternatively, an increase in x
will result in a more perpendicular orientation of the adductor fibres. If x is
kept constant and y decreased the fibre orientation would become more vertical
208 ANNALS OF THE SOUTH AFRICAN MUSEUM
A
B
A. F
R
B.
c 4
Fig. 18. Diagram to illustrate the action of the lower jaw.
and if y is increased the fibre orientation would become more posteriorly
directed. It is clear therefore that the coronoid process in terms of height (y)
and distance from the articulation (x) is functionally important in determining
the action of the lower jaws and its associated musculature, and not as De
Weerdt (1971) suggests, mainly a strengthening device.
CRANIAL AND CERVICAL MUSCLES OF TYPHLOSAURUS AURANTIACUS AURANTIACUS 209
The vertebrate lower jaw, seen in terms of the mechanics of a third class
lever, would theoretically function at optimal efficiency with the adductor
fibres orientated perpendicular to the long axis of the lower jaw and inserted
on to the jaw as close as possible to the symphysis.
Ostrom (1964) notes two disadvantages in this arrangement. As a result
of increasing x the gape of the mouth will correspondingly decrease. However,
De Mar & Barghusen (1973: 626) state: ‘If the increase in relative lengths
of the moment arms achieved by increasing y and increasing x are the same,
the increase in distance that the muscle must stretch to achieve a given gape
is the same for the two methods. Thus . . . reduction of gape is the same and
not a consideration per se in making the comparison.’
Secondly, Ostrom (1964) indicates that because of their vertical orientation
the origin of the adductor fibres would encroach on to the facial region, restrict-
ing their size and power. This statement is true except in cases where the orbit
is anteriorly placed or decreased in size, making it possible to extend the tem-
poral origin of the jaw adductors anteriorly. In Typhlosaurus the eye is degenerate
and lacks eye muscles. The orbit is consequently reduced and the temporal
region elongated. The lateral downgrowth of the parietal and the forward
extension of the anterior superior process of the pro-otic makes additional
areas of origin available for the adductor musculature, compensating for the
loss of such structures as the supratemporal arch. The relatively forward
position, therefore, of the adductor muscles suggests a difference in the line
of muscle action as compared to a non-fossorial skink like Mabuia.
From the work of De Mar & Barghusen (1973) it is clear that the height
and position of the coronoid process is influenced by the line of muscle action.
Any difference, therefore, in the line of muscle action would be reflected in
the proportions of the lower jaw.
In comparing the lower jaws of Typhlosaurus and Mabuia the outstanding
feature is their proportional similarity in terms of x and y. The tooth row,
however, is shorter in Typh/osaurus than in Mabuia because of the subterminal
mouth of the former. Consequently, the force of the bite at the jaw symphysis
will probably be proportionately greater in Typhlosaurus than in Mabuia. The
similarity of the two lower jaws suggests that the mean line of muscle action
is identical in both forms, and that there is probably no difference of any
consequence in the action of the jaw.
It therefore appears that the primary adaptation for a fossorial habit is
streamlining of the body and its various parts. Loss of limbs and attenuation
of the body is associated with the new mode of locomotion. Strengthening
of the skull for burrowing results in the lateral downgrowth of the parietal
bone and the extension of the anterior superior process of the pro-otic bone,
the lengthening of the temporal region, and the loss of metakinesis and the
supratemporal arch.
It may be concluded therefore that the distribution of the jaw muscles
in Typhlosaurus represents the optimal functional arrangement to maintain a
210 ANNALS OF THE SOUTH AFRICAN MUSEUM
mode of jaw action essentially similar to that of a non-fossorial lizard such as
Mabuia capensis, within a skull that is proportionally different because of
marked changes resulting from a fossorial mode of life.
ACKNOWLEDGEMENTS
I wish to thank the following persons: Professor M. E. Malan of the
Zoological Institute of the University of Stellenbosch, who suggested the
project, for her guidance and assistance during the research; Dr M.A. Cluver
of the South African Museum for critically reading the manuscript; Messrs
M. N. Bester, A. J. Lindvelt, D. J. van Eeden, D. P. Mostert and Mrs
R. Semmellink, all of the Zoological Institute at Stellenbosch for assistance
rendered at various stages of the work; Mrs I. Chesselet of the South African
Museum, her family and Mr P. J. Louw of Klein Botrivier for their fieldwork;
Mr N. J. Eden of the South African Museum for taking the photographs;
Miss A. E. Louw and Mrs P. D. Eedes for helping with the typing, and my wife
Lorna for her valued assistance.
REFERENCES
Apams, L. A. 1919. A memoir on the phylogeny of the jaw muscles in Recent and fossil
vertebrates.— Ann. N.Y. Acad. Sci. 28: 51-166.
AUFFENBERG, W. 1962. A review of the trunk musculature in the limbless land vertebrates. —
Am. Zool. 2: 183-190.
BARGHUSEN, H. R. 1973. The adductor jaw musculature of Dimetrodon (Reptilia, Pelycosauria).
—J. Paleont. 47: 823-834.
Bettairs, A. d’A. 1969. The Life of Reptiles. London: Weidenfeld and Nicholson.
Bonin, J. J. 1965. The eye of Agamadon anguliceps Peters (Reptilia, Amphisbaenia). — Copeia
1965: 324-331.
*BOULENGER, G. A. 1887. Catalogue of the Lizards in the British Museum (Natural History) 3.
2nd ed. London: British Museum.
*Boyb, E. 1969. The cranial morphology of the African limbless skink Melanoseps. Unpub-
lished MSc. thesis, University of Stellenbosch.
BROADLEY, D. G. 1968. A revision of the African genus Typhlosaurus Wiegmann (Sauria:
Scincidae).— Arnoldia (Rhodesia) 3 (36): 1-20.
Brock, G. T. 1941. The skull of Acontias meleagris, with a study of the affinities between
lizards and snakes.—J. Linn. Soc. (Zool.) 41: 71-88.
Camp, C. L. 1923. Classification of Lizards.— Bull. Am. Mus. nat. Hist. 48: 289-481.
*CLuver, M. A. 1965. The cranial morphology of the limbless skink Typhlcontias. Unpub-
lished MSc. thesis, University of Stellenbosch.
De Mar, R. & BARGHUSEN, H. R. 1973. Mechanics and the evolution of the synapsid jaw. —
Evolution Lancaster, Pa. 26: 622-637.
De Vituiers, C. G. S. 1939. Uber den Schiadel des Siidafrikanischen Schlangenartigen Scin-
ciden Acontias meleagris.— Anat. Anz. 88: 320-347.
De Weerpt, A. 1971. The skull, jaw muscles and cranial kinesis of Dibamus novae guineae
with special reference to the systematic position of the Dibamidae.— Unpublished MSc.
thesis, University of Stellenbosch.
*Du Tort, F. L. 1971. The cranial morphology of the African burrowing lizard Feylinia
polylepis (Bocage).—Unpublished MSc. thesis, University of Stellenbosch. ;
EpGewortH, F. H. 1935. The cranial muscles of vertebrates. Cambridge: Cambridge University
Press.
CRANIAL AND CERVICAL MUSCLES OF TYPHLOSAURUS AURANTIACUS AURANTIACUS 211
Evans, F. G. 1939. The morphology and functional evolution of the atlas-axis complex from
fish to mammals.—Ann. N.Y. Acad. Sci. 39: 29-104.
FitzSimons, V. F. M. 1943. The lizards of South Africa.— Transv. Mus. Mem. 1: i-xv, 1-528.
FRAZETTA, T. H. 1962. A functional consideration of cranial kinesis in lizards.—J. Morph.
111: 287-320.
Gapow, H. 1901. The Cambridge Natural History. 8. Amphibia and Reptiles. London:
Macmillan.
Gasc, J. P. 1965. Les adaptions anatomique du lézard apode Feylinia currori Gray au fouissage
par reptation ondulante.—C. r. hebd. Séanc. Acad. Sci., Paris 260: 1248-1251.
Gasc, J. P. 1967a. Introduction a |’étude de la musculature axiale des squamates serpenti-
formes.—Mém. Mus. natn. Hist nat. Paris (A) 48 (2): 69-124.
Gasc, J. P. 19675. Retentissement de I’adaptation a la locomotion apode sur le squelette des
squamates. — Colloques int. Cent. natn. Rech. scient. Evolution des Vertébrés. 163: 373-394.
Gasc, J. P. 1967c. Un cas particulier de l’adaptation a la vie souterraine: le lézard serpenti-
forme Dibamus, Dumeril et Bibron.—C. r. hebd. Sci. Acad. Séanc., Paris (D) 265: 41-43.
Gasc, J. P. 1968. Contribution a l’ostéologie et a la myologie de Dibamus novaeguineae Gray
(Sauria, Reptilia).—Ann/s Sci. Nat. (Zool.) (12) 10: 127-150.
Greer, A. E. 1970. A subfamilial classification of scincid lizards.— Bull. Mus. comp. Zool.
Hary. 139: 151-184.
*HAACKE, W. D. 1964. Description of two new species of lizards and notes on Fitzsimonsia
brevipes (FitzSimons) from the central Namib Desert.— Scient. Pap. Namib Des. Res. Stn.
25: 1-15.
*Haas, G. 1930. Uber die Kaumuskulatur und Schadelmechanik einiger Wihlschlangen. —
Zool. Jb. 52: 95-218.
Haas, G. 1934. Beitrag zur Frage der Homologisierung der Kiefermuskulatur der Ophidia
und Sauria.— Biologia gen. 10: 211-226.
Haas, G. 1960. On the trigeminus muscles of the lizards Xenosaurus grandis and Shinisaurus
crocodilurus.— Amer. Mus. Novit. (2017): 1-54.
Haas, G. 1973. Muscles of the jaws and associated structures in the Rhynchocephalia and
Squamata. Jn: GANs, C., BELLAIRS, A. d’A. & PARsons, T. S. eds. Biology of the Reptilia
4: 285-539. London & New York: Academic Press.
Hewitt, J. 1929. On some Scincidae from South Africa, Madagascar and Ceylon.—Ann.
Transv. Mus. 13: 1-8.
HorsTeTTerR R. & Gasc, J. P. 1969. Vertebrae and ribs of modern Reptiles. Jn: GANs, C.,
BeLvairs, A. d’A. & Parsons, T. S. eds. Biology of the Reptilia 1: 201-310. London &
New York: Academic Press.
Hotper, L. A. 1960. The comparative morphology of the axial skeleton in the Australian
Gekkonidae.—J. Linn. Soc. (Zool.) 64: 300-335.
Huey, R. B., PIANKA, E. R., EGAN, M. E. & Coons, L. W. 1974. Ecological shifts in sympatry:
Kalahari fossorial lizards (Typhlosaurus).— Ecology 55: 304-316.
IORDANSKY, N. N. 1970. Structure and biomechanical analysis of functions of the jaw muscles
in the lizards.— Anat. Anz. 127: 383-413.
KRITZINGER, C. C. 1946. The cranial anatomy and kinesis of the South African amphisbaenid
Monopeltis capensis Smith.—S. Afr. J. Sci. 42: 175-204.
Lakser, T. 1926. Studien iiber die Trigeminus-versorgte Kaumuskulatur der Sauropsiden.
Kopenhagen: C. H. Reitzel.
LANGENBARTEL, D. A. 1968. The hyoid and its associated muscles in snakes.—J/linois biol.
Monogr. 38: 1-156.
LEONARD, C. J. 1973. The cranial morphology of the didactyle sand skink Scelotes bipes
sexlineatus (Harlan). Unpublished MSc. thesis, University of the Western Cape.
List, J. C. 1966. Comparative osteology of the snake families Typhlopidae and Leptotyphlo-
pidae. — I/linois Biol. Monogr. 36: 1-112.
LusoscH, W. 1933. Untersuchung iiber die Visceralmuskulatur der Sauropsiden.— Morph.
Jb. 72: 584-663.
Luter, A. 1914. Uber die vom N. trigeminus versorgte Muskulatur der Amphibien mit
einem vergleichenden Ausblick itiber den Adductor mandibulae der Gnathostomen und
einem Beitrag zum Verstandnis der Organisation der Anurenlarven.—Acta Soc. Scient.
fenn. 44 (7): 1-151.
212 ANNALS OF THE SOUTH AFRICAN MUSEUM
Mertens, R. 1955. Die Amphibien und Reptilien Siidwest-Afrikas.— Abh. senckenb. natur-
forsch. Ges. 490: 1-172.
NisH1, S. 1916. Zur vergleichenden Anatomie der eigentlichen (genuinen) Riickenmuskeln. —
Morph. Jb. 50: 168-318.
OELRICH, T. M. 1956. The anatomy of the head of Crenosaura pectinata (Iguanidae).— Misc.
Publs Mus. Zool. Univ. Mich. 94: 1-122.
OLson, E. C. 1936. The dorsal axial musculature of certain primitive Permian tetrapods. —
J. Morph. 59: 265-311.
Ostrom, J. H. 1961. Cranial morphology of the hadrosaurian dinosaurs of North America.
— Bull. Am. Mus. nat. Hist. 122: 35-186.
Ostrom, J. H. 1962. On the constrictor dorsalis muscles of Sphenodon.— Copeia 1962: 732-735.
Ostrom, J. H. 1964. A functional analysis of jaw mechanics in the dinosaur Triceratops.—
Postilla 88: 1-35.
Peters, W. C. H. 1882. Naturwissenschafliche Reise nach Mossambique, auf Befehl seine,
Majestdat des Konigs Friedrich Wilhelm IV. In den Jahren 1842 bis 1848 ausgefuhrt. Zoologie.
Ill. Amphibien. Berlin: G. Reimer.
POGLAYEN-NEUWALL, Ingeborg. 1953. Untersuchen iiber die trigeminusmuskulatur von
Hatteria.—Z. wiss. Zool. 157: 57-67.
POGLAYEN-NEUWALL, Ingeborg. 1954. Die Kiefermuskulatur der Eidechsen und ihre Inner-
vation.—Z. wiss. Zool. 157: 97-132.
POGLAYEN-NEUWALL, Ivo. 1953a. Untersuchungen der Kiefermuskulatur und deren Inner-
vation an Krokodilen.— Anat. Anz. 99: 257-276.
POGLAYEN-NEUWALL, Ivo. 19535. Untersuchungen der Kiefermuskulatur und deren Innerva-
tion bei Schildkréten.— Acta zool. Stockh. 34: 241-292.
Romer, A. S. 1956. Osteology of the Reptiles. Chicago: University of Chicago Press.
RuGe, G. 1897. Ueber das peripherische Gebiet des Nervus facialis bei Wirbelthieren. Jn:
Festschrift zum 70. Geburtstag von Cari Gegenbaur. 3: 193-348. Leipzig: Engelmann.
SAVE-SGDERBERGH, G. 1945. Notes on the trigeminal musculature in non-mammalian Tetra-
pods.— Nova Acta R. Soc. Scient. upsal. (1V) 13 (7): 1-59.
Smit, E. P. 1964. Die skedelmorfologie van die genus Typhlosaurus (Familie: Scincidae).
Unpublished MSc. thesis, University of Stellenbosch.
SonpDHI, K. C. 1958. The hyoid and associated structures in some Indian reptiles. — Ann.
Zool., Agra. 2: 155-239.
Soop, M. S. 1948. The anatomy of the vertebral column in serpents.—Proc. Indian Acad.
Sci. (B) 28 (1): 1-26.
*STANNIUS, H. 1849. Nouveau Manuel d’ Anatomie Comparee 2. Animaux vertébrés. Paris: Roret.
TOERIEN, M. J. 1950. The cranial morphology of the Californian lizard Anniella pulchra
(Gray).—S. Afr. J. Sci. 46: 321-342.
*VaLLois. H. V. 1922. Les transformations de la musculature de I’episome chez les vertébrés. —
Archs Morph. gén. exp. 13: 1-538.
VAN DER Merwe, N. J. 1944. Die Skedelmorfologie van Acontias meleagris (Linn.).—Tydskr.
Wet. Kuns. 5: 59-88.
VersLuys, J. 1910. Streptostylie bei Dinosauriern.— Zool. Jb. (abt. f. Anat.) 30: 175-260.
VerSLuys, J. 1912. Das Streptostylie— Problem und die Bewegung im Schadel bei Sauropsiden.
—Zool. Jb. (Suppl.) (15) 2: 545-716.
*VeTTER, B. 1874. Untersuchungen zur vergleichenden Anatomie der Kiemen- und Kiefer-
muskulatur der Fische.—Jena Z. Naturw. 8: 405-458.
*VeTTER, B. 1878. Untersuchungen zur vergleichenden Anatomie der Kiemen- und Kiefer-
muskulatur der Fische.—Jena Z. Naturw. 12: 431-450.
WALLS, G. L. 1942. The vertebrate eye and its adaptive radiation. — Bull. Cranbrook Inst. Sc. 19.
*WIEGMANN, A. F. A. 1834. Herpetologica mexicana, seu Descriptio Amphibiorum Novae
Hispaniae . . . Pars prima Saurorum species: 1—54. Berlin.
Witte be, G. F. & LAURENT, R. 1943. Contribution a la systématique des formes dégradées
de la famille des Scincidae apparentees au genre Scelotes Fitzinger.—Mem. Mus. r. Hist.
nat. Belg. (2) 26: 1-44.
ZANGERL, R. 1945. Contributions to the osteology of the post-cranial skeleton of the Amphis-
baenidae.— Am. Midl. Nat. 33: 764-780.
*Not seen in the original.
CRANIAL AND CERVICAL MUSCLES OF TYPHLOSAURUS AURANTIACUS AURANTIACUS
mq
ms
nl
oc
ABBREVIATIONS
anterior process of the coronoid
m. adductor mandibulae externus medius
m. adductor mandibulae externus profundus
m. adductor mandibulae externus superficialis
anterior inferior process of the pro-otic
m. adductor mandibulae posterior
articular
anterior superior process of the pro-otic
stapedial artery
bodenaponeurosis
buccal cavity
buccal lining
basioccipital
basipterygoid process
brain
basal tuberosity
coronoid
m. constrictor colli
internal carotid artery
m. cervicomandibularis
coronoid process
dentary
anterior portion of the m. depressor mandibulae
posterior portion of the m. depressor mandibulae
eye
m. episternocleidomastoideus
extracolumella
Gasserian ganglion
m. genioglossus
m. geniohyoideus lateralis
m. geniohyoideus medialis
groove for the insertion of the pseudotemporalis muscle
fascia supporting the extracolumella anteriorly
pad of fibrocartilage
foramen for the chorda tympani
m. hyoglossus
m. intermandibularis anterior
infraorbital fenestra
m. intermandibularis posterior
insertional tendon of the m. cervicomandibularis
insertional tendon of the m. pterygoideus
lateral lamina of the bodenaponeurosis
m. longus colli
m. longissimus capitis
m. longissimus cervicis
longitudinal lingual fibres
m. levator pterygoidei
lateral lamina of the quadrate tendon
lateral semicircular canal
lateral head vein
musculus
medial lamina of the bodenaponeurosis
Meckelian cartilage
medial lamina of the quadrate tendon
lateral mandibular shelf
nuchal ligament
m. obliquus capitis magnus
213
214 ANNALS OF THE SOUTH AFRICAN MUSEUM
oe fused ophistotic-exoccipital
p m. pseudotemporalis
pa pre-articular
pal palatine
par parietal
pat ascending process of the tectum synoticum
pb __ parasphenoid — basisphenoid
pe __ posterior process of the coronoid
pd __ tendinous pad on m. pterygoideus
pe _ lingual process
pf postfrontal
pm _ posterior mylohyoid foramen
pn palatine nerve
po pro-otic
prp m. protractor pterygoidei
pt pterygoid
pts m. pterygoideus
q quadrate
qa quadrate tendon
r rostral
ram. rectus capitis anterior
rm mandibular ramus of V
rma maxillary ramus of V
rop opthalmic ramus of V
rpm. rectus capitis posterior
S stapes
sa surangular
sc spinalis capitis
sd =m. spinalis dorsi
sp ___ splenial
sq. squamosal
st supratemporal
t tendinous sheath of the extra columella
tc tendinous connection between supratemporal and quadrate
tl tranverse lingual fibres
to tongue
tr __ trachea
ts tendon between the squamosal and quadrate
vb _ vertical lamina of the bodenaponeurosis
vl _ vertical lingual fibres
vq vertical lamina of the quadrate tendon
6. SYSTEMATIC papers must conform with the International code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. n., sp. n., comb. n.,
syn. n., etc.
An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.
Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (figs 8a—b).
Nucula largillierti Philippi, 1861: 87
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9
Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma separates consecutive numbers
Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.
In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:
Holotype
SAM-A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach,
Port Elizabeth (33.51S, 25.39E), collected by A. Smith, 15 January 1973.
Note standard form of writing South African Museum registration numbers and of date.
7. SPECIAL HOUSE RULES
Capital initial letters
(a) The par Maps and Tables of the paper when referred to in the text
e.g. . the Figure depicting C. namacolus .
. in C. namacolus (Fig. 10) .
(b) The eda of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A. L. du Toit
Von Huene _—i but _-F. von Huene
(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian
Punctuation should be loose, omitting all not strictly necessary
Reference to the author should be expressed in the third person
Roman numerals should be converted to arabic, except when forming part of the title of a
book or article, such as
‘Revision of the Crustacea. Part VIII. The Amphipoda.’
Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.
JURI A. VAN DEN HEEVER.
THE CRANIAL AND CERVICAL MUSCLES OF TH
SOUTH AFRICAN LIMBLESS LIZAR
TYPHLOSAURUS AURANTIACUS AURANTIAC
PETERS (REPTILIA, SAURIA
JOLUME 69 PART 9 APRIL 1976 ; ISSN 0303-2515
S Lif - C/ayehrim ti/H7
OF THE SOUTH AFRICAN
MUSEUM >
‘APE TOWN
INSTRUCTIONS TO AUTHORS
1. MATERIAL should be original and not published elsewhere, in whole or in part. When
accepted, copyright becomes the property of the Trustees of the South African Museum.
2. LAYOUT should be as follows:
(a) Masthead to consist of
Title: informative but concise, without abbreviations and not including the names of new genera or species
Author’s(s’) name(s)
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(b) Abstract of not more than 200 words, intelligible to the reader without reference to the text
(c) Table of contents giving hierarchy of headings and subheadings
(d) Introduction
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(th) References
(i) Abbreviations, where these are numerous
3. MANUSCRIPT, to be submitted in triplicate, should be typewritten, double spaced with
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The number of the figure should be marked on the back of each illustration.
5. REFERENCES cited in text and synonymies should all be included in the list at the end of
the paper, using the Harvard System (ibid., idem., loc. cit., op. cit. are not acceptable):
(a) Author’s name and year of publication given in text, e.g.:
“Smith (1969) describes...’
‘Smith (1969: 36, fig. 16) describes...”
‘As described (Smith 1969a, 1969b; Jones 1971)...’
“As described (Haughton & Broom 1927)...’
‘As described (Haughton ef al. 1927)...’
Note: no comma separating name and year
Pagination indicated by colon, not p.
names of joint authors connected by ampersand
et al. in text for more than two joint authors, but names of all authors given in list of references
(b) Full references at the end of the paper, arranged alphabetically by names, chronologically
within each name, with suffixes a, b, etc. to the year for more than one paper by the same
author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a).
For books give title in italics, edition, volume number, place of publication, publisher.
For journal article give title of article, title of journal in italics (abbreviated according to the World list of
scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part
number (only if independently paged) in parentheses, pagination (first and last pages of article).
Examples (note capitalization and punctuation)
BuLLouGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
Fiscuer, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques.—J. Conch., Paris 88: 100-140.
FiscHER, P.-H., DuvaL, M. & RAFFy, A. 1933. Etudes sur les échanges respiratoires des littorines.— Archs
Zool. exp. gén. 74: 627-634.
Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. —
Ann. Mag. nat. Hist. (13) 2: 309-320. :
Konn, A. J. 19605. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. —
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. — Denkschr. med.-naturw. Ges. Jena 16: 269-270. ae
continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 69 Band
April 1976 April
Part 9 Deel
THE PLIOCENE FOSSIL OCCURRENCES IN
‘E’ QUARRY,
LANGEBAANWEG, SOUTH AFRICA
By
O.. B.. BENDEY
Cape Town Kaapstad
The ANNALS OF THE SOUTH AFRICAN MUSEUM
are issued in parts at irregular intervals as material
becomes available
Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000
Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
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Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad 8000
OUT OF PRINT/UIT DRUK
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Price of this part/Prys van hierdie deel
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Trustees of the South African Museum © Trustees van die Suid-Afrikaanse Museum
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ISBN 0 949940 88 7
Printed in South Africa by In Suid-Afrika gedruk deur
The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk.,
Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap
THE PLIOCENE FOSSIL OCCURRENCES IN ‘E’ QUARRY,
LANGEBAANWEG, SOUTH AFRICA
By
Q. B. HENDEY
South African Museum, Cape Town
(With 6 figures and 4 tables)
[MS. accepted 27 November 1975}
ABSTRACT
The Pliocene Varswater Formation in the vicinity of Langebaanweg, Cape Province, is
comprised of three main units, now named the Gravel, Quartzose Sand and Pelletal Phosphorite
Members. The lowest unit in the succession, the Gravel Member, has yielded a largely marine
fauna, including fifteen shark species, incorporated in rocky and sandy beach deposits. The
principal source of fossils, the Quartzose Sand Member, was laid down in a variety of
depositional environments in and near an estuary. The economically important unit, the
Pelletal Phosphorite Member, is fossiliferous in a relatively limited area which was situated in
the immediate vicinity of the river mouth. The fossils from the deposits overlying the Gravel
Member represent a wide variety of marine, freshwater and terrestrial invertebrates and
vertebrates. About seventy-five mammalian species are recorded, including a few belonging to
groups not previously recorded from Africa.
CONTENTS
PAGE
Introduction : : . : Pe eA ls:
Geology . . : : : = = 218
Depositional environments . a 222
Fauna . : ; : : : = 23
Plotay. ‘ f F : : = 243
Dating . : ’ : : : ee 2AS
Conclusions : : ‘ : . 245
Acknowledgements . ; ‘ . 245
References . : : : . ~ 246
INTRODUCTION
Large-scale production of phosphate from deposits on the farm Langeberg
near Langebaanweg, Cape Province, was commenced in 1953 and several years
later the presence of fossils in these deposits was reported (Singer 1961). Initially
few fossils of good quality were collected, but prospecting revealed the presence
of highly fossiliferous deposits south of a small open-cast mine, ‘E’ Quarry.
During 1964 these deposits were exposed in a trench about 240 metres long,
120 metres wide and ranging in depth from 2 to 30 metres (Fig. 1). Soon after
the 1964 excavations were commenced it became apparent that the deposits
would yield well-preserved fossils in large quantities. Mining of ‘E’ Quarry (the
New Varswater Mine) has been continuous since then and although the number
215
Ann. S. Afr. Mus. 69 (9), 1976: 215-247, 6 figs, 4 tables.
216 ANNALS OF THE SOUTH AFRICAN MUSEUM
of fossils recovered has varied from year to year, the original expectations have
been fulfilled. Certainly no other recorded African fossil occurrence of Pliocene
age has produced so large an assemblage of specimens representing so wide a
variety of species.
During 1968 the South African Museum commenced an investigation of the
fossiliferous deposits exposed in ‘E’ Quarry, this project following on from a
similar undertaking directed for a period of ten years by R. Singer of the
University of Chicago. The first phase of the Langebaanweg project had also
taken into account the mined-out occurrences at ‘C’ Quarry on the farm
Langeberg and Baard’s Quarry on the farm Muishondsfontein. The fossils from
these sites are limited in both quality and quantity and there are some still
unresolved problems relating to their geological associations.
Several publications have resulted from the second phase of the Langebaan-
weg project, including reviews (Hendey 1970a; 1973; 1974a), accounts of the
geology (Tankard 1974a; 19746; 1975), discussions on dating (Hendey 19706;
1972a; 19746), as well as descriptions of some of the fossils recovered (e.g.
Simpson 1971; Kensley 1972; Gentry 1974).
The geological study of the ‘E’ Quarry deposits, which has now been
largely completed, formed part of a broadly-based investigation of late Cenozoic
deposits in the south-western and southern Cape Province, and was undertaken
independently of the palaeontological study, although the two have been
mutually complementary.
Recently research on the fossils from Langebaanweg has decreased, although
there has been an increase in the amount of material collected. The latter
development is a direct result of changes in the mining programme. The expected
back-filling of ‘E’ Quarry (Hendey 1973) was commenced during 1974 and
collecting was accelerated in those areas to be covered by new mine dumps. In
addition, the mining company (Chemfos Limited) removed a large quantity of
fossiliferous deposit from one of the threatened areas (East Stream, see Fig. 1)
and this is being screened by the first permanent field assistants on site. The
nature of the present undertaking is being further influenced by the fact that
during 1975 mining of the last of the phosphatic deposits known to be highly
fossiliferous was commenced. Until recently these deposits were not scheduled
to be mined until about 1990.
Since its inception, the prime object of the present phase of the Langebaan-
weg project has been the recovery and identification of fossils, with the collecting
being as comprehensive as possible. The recent changes in the mining programme
have added urgency to this aspect of the undertaking since leisurely collecting
and excavation over certain areas of the mine are no longer possible and the
areas thus affected will increase with the passage of time. Many of the specimens
still in the deposits will be lost if they are not salvaged promptly. In addition,
the volume of fossiliferous deposit elsewhere is being steadily reduced by the
mining and although some is likely to remain indefinitely, technical difficulties
may make it inaccessible to further exploitation. The termination of active field
‘i
]
2
PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA
‘SaTJOU OST AjoyeuTxosdde our] ayeog *AreNg <O, NO
“Poul OY “CT “UONPARSXO HOG] OY} JO JU9}XO oJeUNIXOIdde ayvoIPuUl g punose soul] pojdding *poq wag ‘D “gee pag Jo soinsodxgq “g “WIROINIS SPI VY
JO MOIA [BOY “T “SIF
‘(PL6L A[ne) SomueegosueT
“AlIeNg)
As |
,
218 ANNALS OF THE SOUTH AFRICAN MUSEUM
work at the site in the not too distant future has now become a distinct
possibility.
The collecting of fossils could be expedited by being more selective, but as
far as possible this approach is avoided, mainly because it would adversely affect
that part of the undertaking concerned with the recording of the nature of the
fossil occurrences. This aspect of the investigation already suffers because of its
lower priority rating. It is hoped that eventually an analysis of specimens from
any given area in terms of species and body part representation, together with
their condition and associations, will provide information on environments of
deposition, the taphonomy of the fossils and aspects of the ecology of the area
at the time of deposition. Some information of this nature is already available.
Although more time has recently been spent on the collecting of fossils and
directly related technical matters, some progress has been made in the
palaeontological research, and the present report updates some of the
information and opinions previously recorded.
GEOLOGY
The phosphatic and fossiliferous deposits exposed in ‘E’ Quarry make up
what is now termed the Varswater Formation (Hendey 1974a; Tankard 1975).
This formation is Pliocene in age and it is underlain by the Miocene Saldanha
Formation (Tankard in press). The overlying deposits, informally termed the
‘surface bed’, are largely, or entirely Pleistocene and Holocene in age. The
Varswater Formation is comprised of three main units which were referred to
by a variety of names in earlier publications. The member names used in the
present report (Table 1) are those which are now considered most appropriate
(A. J. Tankard pers. comm.), while those of the beds are purely informal and
likely to be modified at a later date. The Gravel and Pelletal Phosphorite
Members are both unequivocally described by their names, but the name of the
Quartzose Sand Member refers to the dominant lithological element. This
member also includes horizons of carbonaceous sand and clay (the ‘peat bed’,
Fig. 1), clayey sands and silt. The non-geographic names of the members are
justified in terms of Section 3.10(d) of the South African Code of Stratigraphic
Terminology and Nomenclature (1971: 118).
Since Tankard (1975) has described the Varswater Formation in detail, the
only other comments on the geology of ‘E’ Quarry which are included here are
those which have a bearing on the palaeontology of the deposits.
The Quartzose Sand Member is the most highly fossiliferous of the three
units, while the Pelletal Phosphorite Member is poorly fossiliferous, except for
an area still exposed of the west wall of the mine (‘bed 3a’, Fig. 1). There are
only limited exposures of the Gravel Member in a few areas of the mine and
relatively little attention has been paid to this unit during the current phase of
the Langebaanweg project.
PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 219
TABLE |
Stratigraphy of the ‘E’ Quarry exposures of the Varswater Formation
Present report
Hendey (1974) | Tankard (1975)
MEMBERS BEDS
| beds 3aS & 3aN
Pelletal Pelletal (fossiliferous)
z Bed 3 Phosphorite Phosphorite and other un-
g Member Member | named beds
< | ee ee eS eee
z Fluvial Quartzose tidal mud flat
2 | Bed 2 | Sand Sand bed, peat bed
x Member Member and other un-
E named beds
2 (fossiliferous)
4 ———_— qt sq_i—m—~:
S$ Beach Gravel | —
Bed 1 | Gravel Member
| Member
Over most of ‘E’ Quarry the three members of the Varswater Formation
are readily identifiable. The Gravel Member is always unmistakable, although
the sandy element of this member may on occasion have been regarded as part
of the overlying Quartzose Sand Member. Recognition of the Pelletal Phospho-
rite Member has, for the most part, not been difficult, although its lower limit
has not always been clearly defined. The problems encountered in recognizing
the lower limit of this member have been due to a variety of factors. Theoretically
the mining is cut off at the base of the Pelletal Phosphorite Member, although in
practice this is impossible and irregularities on the floor of the mine do not
necessarily reflect the nature of the boundary between the Pelletal Phosphorite
and Quartzose Sand Members. In most areas the mining has actually extended
into the latter unit where the exposed surface may be contaminated and obscured
by spillage from the excavator and slumping of deposit from the vertical mine
faces. Furthermore, the Quartzose Sand Member was truncated prior to the
deposition of the Pelletal Phosphorite and where it is very thin it has been
difficult or impossible to recognize. In certain critical areas the deposits of the
two members are superficially similar and although detailed sediment analyses
would no doubt resolve the issue in problematical instances, no such analyses
have been undertaken.
From the preceding comments it should be clear that difficulties have centred
largely on the identification of the upper and lower limits of the Quartzose Sand
Member. Added to this is the fact that this member is the most complicated unit
in the succession in terms of lithology and the variable character of the deposits
has itself led to some confusion in the past. The sometimes striking differences
220 ANNALS OF THE SOUTH AFRICAN MUSEUM
TABLE 2
The stratigraphy of the East Stream and peat areas of ‘E’ Quarry
— __ rereoeoro—Xs¥KXn—w—X—n—“n=“—xowoeneoeOeOwooOOO a si I
EAST STREAM PEAT AREA
PELLETAL Phosphatic sand Phosphatic sand
z PHOSPHORITE (? non-fossiliferous) (? non-fossiliferous)
) MEMBER
Py;
S Silt
2 QUARTZOSE (invertebrates common)
= SAND Quartzose sand |-—--—-—-—-—-—--------
> MEMBER (vertebrates common) Peat
2 (vertebrates common)
wz eee eee ee ee
S GRAVEL Sand & gravel Sand & gravel
MEMBER (marine fossils common) | (marine fossils common)
in the nature of the Quartzose Sand Member are illustrated by examples of the
‘E’ Quarry succession given in Table 2.
The difficulty experienced in identifying the units of the Varswater Forma-
tion has resulted in some of the fossils collected being of doubtful provenance.
In this connection it should also be noted that before the basic three-unit
succession was recognized in 1969, little or no data on the source of specimens
was recorded and the provenance of many specimens collected before that date
may never be known. The same often applies in the case of specimens picked up
by mine workers. Although the problem of unprovenanced material is not as
serious as it might have been, it is an unfortunate complication in the
palaeontological investigation.
In the faunal lists given later, records of unknown or uncertain origin are
excluded. The source of most of the more significant specimens from ‘E’ Quarry
is well documented and all the species identified to date are represented by at
least one specimen of known provenance.
The characteristics of the units comprising the Varswater Formation are
accounted for by the sequence of events which occurred during deposition of the
formation. These events have a direct bearing on the interpretation and identifi-
cation of depositional environments (vide infra). They were summarized by
Tankard (1974a: 219) who stated that, ‘In the Pliocene a transgressing sea
pushed deltaic marsh sediments ahead of it until it reached a temporary still-
stand .. . [with the] temporarily stable conditions [allowing] a barrier bar to
build up, behind which estuarine conditions prevailed. The estuary was fed by a
river from the north-east. The final transgression reworked the older sediments.’
It was at the time of the stillstand that the Quartzose Sand Member was
accumulated, while the Pelletal Phosphorite Member was laid down during the
final transgression (Fig. 2).
Sea
Breaker- bar
PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA
—-*
- THE GRAVEL MEMBER
THE QUARTZOSE SAND MEMBER
AS SQeF Se SS OWE eer —- --- -----~--------------
ai
THE PELLETAL PHOSPHORITE MEMBER
Miocene deposits (Saldanha Formation) reworked by a _ Pliocene marine transgression
SG Spo SSP BS TAs EA ~~ ~~~ --------
Sse
22 oe
transgression -
PesFa fae =.
SMP oBze same ara Shade
SO SOS Sess gOS o Hea
Se
aqeaeas: Se FSP5 Sac
SS F405
Mud flats
(a5 o*
225222
Oo
mQ=—s
&
2790 Porn 23 ae.
—_—_- ~~
23°. x
CN a egw ne og a
=
ood
SEB
Soo?
POF S0¢
'
i=
o
—_
”
”
oe
rs
i— 1)
”
c
os
-
~-
oe
J
~
Cc
usc
c
o
-
"”
_—
=
-
a”
>
<
oo
=
°
a
=
a
~
oc
i=
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—
-
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-
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=
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i—|
Deposits accumulated during the final phase of the
SPELES
221
Fig. 2. The depositional history of the Varswater Formation.
222 ANNALS OF THE SOUTH AFRICAN MUSEUM
Although the geology of the Varswater Formation is now well known,
there is still scope for further detailed work. Of particular interest to the
palaeontological investigation is geological evidence indicating depositional
environments. The relationships between fossils and the deposits in which they
occur has hitherto received little attention.
DEPOSITIONAL ENVIRONMENTS
It is now generally accepted that the Varswater Formation accumulated
during a marine transgression at a time when a river discharged into the sea in
the immediate vicinity of ‘E’ Quarry. It follows that marine, fluvial and terrestrial
environments were then present in the area, together with some of the attendant
micro-environments peculiar to each. Theoretically it is therefore possible that
deposition of this formation took place in more than one of these environments
and also that as the sea transgressed macro- and/or micro-environmental
changes might have occurred in any given area now exposed in the mine.
Tankard (1975) has given an account of the depositional environments of
the three units comprising the Varswater Formation in ‘E’ Quarry. They were
as follows:
Gravel Member—Rocky and sandy marine beach environments.
Quartzose Sand Member—Essentially estuarine and fluvial environments,
although a peat deposit, probably representing a marsh environment,
is also mentioned.
Pelletal Phosphorite Member—A_ shallow-water environment situated
between a beach bar and beach.
The means by which the vertebrate fossils in deposits overlying the Gravel
Member came to be incorporated in these deposits has been the subject of some
dispute in the past (Tankard 1975: 281). There is in fact no single answer to this
question and it is clear that just as the depositional environments of the sediments
varied, so too did those of the fossils. The characteristics of the deposits change
both vertically and horizontally, the most noticeable and frequent changes being
in the lower levels of the succession, that is, those levels in which fossils occur
most commonly. Although no detailed study of the relationships between
sediments and fossils has yet been undertaken, it is obvious that differences in
the deposits go together with palaeontological differences.
Some observations on the environments in which the ‘E’ Quarry fossils were
laid down have already been recorded (Hendey 1974a), but it is probably worth
while at this stage to elaborate on earlier statements.
The Gravel Member
The fauna of the Gravel Member is comprised overwhelmingly of marine
species, both invertebrates and vertebrates, and the fossils of this member were
undoubtedly accumulated along a marine shoreline. The invertebrates indicate
PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 223
the presence of both rocky and sandy habitats. Remains of terrestrial vertebrates
do occur, but they are invariably fragmented and rolled. This also applies to
many of the marine fossils. The condition of the fossils is evidently the result of
wave action and it differs from that of fossils from fluvial deposits in the
Quartzose Sand and Pelletal Phosphor te Members.
The Quartzose Sand Member
The situation in respect of Quartzose Sand Member depositional environ-
ments is complicated. Tankard (1975) has recognized both estuarine and fluvial
facies within this member and many of the fossils recovered from the ‘E’ Quarry
exposures must have been deposited in the river and estuary. For example, some
of the fossiliferous exposures in the eastern parts of the mine are medium to
coarse sands which are a westerly extension of the fluvial deposits referred to by
Tankard (1975: 274). On the other hand, the nature of some fossil occurrences
suggests that specimens were deposited in areas adjacent to the river and estuary
in both subaerial and subaqueous situations (e.g. floodplain and pond).
Evidence suggesting that certain of the Quartzose Sand Member fossils
were accumulated on land surfaces has been mentioned elsewhere (Hendey
1974a: 349-353). Since this discussion related to carnivore activity rather than
environments of deposition, some points concerning the latter were omitted or
insufficiently emphasized.
In certain of the Quartzose Sand Member exposures (e.g. the floodplain
deposits of East Stream) the condition of the fossils and the nature of their
occurrence contrasts with the situation where there is incontrovertible evidence
for subaqueous deposition of material (e.g. in the fluvial deposits of the Pelletal
Phosphorite Member). The fossils in the latter deposits tend to be abraded and
fragmented, while elements of single skeletons are dispersed. Exceptions to these
rules were probably specimens which had been protected by soft tissues or which
had not been subjected to prolonged transport. The fossils of subaerially
accumulated assemblages are generally perfectly preserved and damage to, or
dispersal of, specimens can usually be ascribed to carnivore activity or fires.
The fact that there are certain Quartzose Sand Member fossil occurrences
where only terrestrial species are recorded, or where they are much more
commonly represented than aquatic species, also tends to suggest that there was
subaerial accumulation of specimens. The presence of some aquatic species
could be explained by periodic inundations of land surfaces. The persistent
presence of an aquatic environment would lead to higher proportions of
aquatic species being represented and there are occurrences in the Quartzose
Sand Member where this is the case. In one such occurrence in the East Stream
area a relatively high proportion of fish bones went together with appreciable
quantities of abraded bone. These fossils were probably deposited in a channel,
a feature for which there was no other obvious evidence.
An example of burnt bone having suggested that a skeleton cannot have
been moved after it was partially burnt was given elsewhere (Hendey 1974a:
224 ANNALS OF THE SOUTH AFRICAN MUSEUM
351). The nature of this particular occurrence was the important factor, since
burnt bone itself is not necessarily proof that there were fires over areas
presently exposed in the mine. Burnt bone might easily have been washed in
from elsewhere. There is another recently discovered example of burnt bone
which must have been in situ. This was a concentration of several hundred bones
of small vertebrates, mainly rodents and insectivores, most of which are heavily
charred. This occurrence is likely to represent the residue of a burnt owl pellet
accumulation, which cannot have been moved after burning without the bones
becoming dispersed.
A hitherto unrecorded factor which supports the theory that some subaerial
accumulation of fossils occurred concerns the presence of coprolites in the
deposits. Coprolites of at least three types have been recovered from the
Quartzose Sand Member. The first and largest type were evidently produced by
large carnivores, probably hyaenas. The second type are smaller and contain
fragmented bones belonging to small vertebrates. They were probably produced
by one or more of the smaller carnivores recorded from this member. An
account of such coprolites was recently given by Mellett (1974). The last type are
small, with no visible bone and tending to be cylindrical when not deformed.
Their source is not known but they are extremely abundant in certain areas
(e.g. East Stream).
Coprolites have been recovered only in certain parts of the mine and, except
for the smallest kind, they are nowhere common. Their condition varies, some
being remarkably fresh in appearance, while others are fragmented and distorted.
One of the ?hyaena coprolites, which was found together with four others, is
flattened, its appearance suggesting that it was trampled when fresh. Some of the
smallest coprolites, which are very fine-textured, show clear impressions made
by leaves, although they may be otherwise undistorted.
In order for faeces to be preserved intact they must almost certainly have
been dropped on a land surface and have been fairly rapidly buried thereafter.
Fresh faeces dropped in and then transported by water is unlikely to have
survived intact for long. It is also unlikely that groups of specimens would have
remained together if they had been transported and there are four instances
recorded where groups of 3, 3, 5 and 7 ?hyaena coprolites were found in close
association. These groups, together with several isolated specimens, came from
a relatively limited area in the eastern part of the mine. Unexcreted faeces could
have been carried to its final resting place while still inside a carcass, but none
of the coprolites has been found in direct association with other remains of their
possible producer. In addition, the shape of the better preserved specimens, and
the presence of leaf impressions on some, indicates that they had actually been
excreted.
Taken in conjunction the various factors referred to above are here regarded
as convincing evidence for subaerial accumulation of at least part of the
Quartzose Sand Member fossil assemblage.
The earlier reference to periodic inundations of land surfaces was intended
PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 225
to indicate seasonal flooding of the river and estuary, but there is also evidence
for tidal flooding. A recently exposed deposit in the south-western part of the
mine includes an invertebrate fauna which indicates that the depositional
environment was a tidal mud flat. The nature of the deposits is in keeping with
the fauna which it contains. The limited exposures of this deposit examined prior
to their becoming temporarily inaccessible suggested that it overlies the peat bed
mentioned earlier (see Table 2).
The faunas of the peat and tidal mud flat beds are strikingly different from
one another and both differ from the fauna of more typical exposures of the
Quartzose Sand Member elsewhere. The mud flat bed is an exceptional
occurrence in this member, being the only one from which large numbers
of well-preserved invertebrate fossils have been recovered. Vertebrate remains
are rare. By contrast vertebrate fossils occur commonly in the peat bed
and in certain other exposures of the Quartzose Sand Member. As fossil-
collecting in this member has progressed, it has become apparent that while
certain vertebrate species are fairly ubiquitous, others occur only in certain
areas, while the overall representation of species and the condition of specimens
varies from place to place. While it may not yet be possible to interpret all such
evidence meaningfully, a superficial comparison of the peat bed and East Stream
faunas should serve to illustrate that different sediments go together with faunal
differences.
In this instance the sediment differences are visually striking, the black
sands and clays of the peat bed contrasting sharply with the white quartzose
sands of the East Stream area. The deposits in the latter area are here regarded
as a largely floodplain accumulation, while the peat area is thought to have been
a marsh.
The fossils of the peat bed tend to be less complete than those from East
Stream, although this may in part, or even largely, be due to the recent disturb-
ance of the peat by mining activities. This disturbance may also account for the
fact that whereas at East Stream a number of instances are recorded where
partial skeletons of individuals were preserved, nothing on a comparable scale
has so far been observed in the peat bed. A notable exception was the discovery
of the distal extremities of a sivathere fore- and hindlimb, elements of which
were found in articulation standing more or less vertically in the deposit,
seemingly all that survived of an animal trapped in the marshy deposits. At East
Stream elements of single skeletons were, with few exceptions, found slightly
dissociated from one another and they tended to lie more or less horizontally in
the deposits. This suggests that they were accumulated on a firm surface.
Although a single species of land tortoise (Chersina sp.) is the most
commonly represented vertebrate in both areas, there are otherwise some
marked differences in the representation of species. There are many species
recorded from East Stream which are not known from the peat bed, although
the reverse either does not apply or is at least much less obvious. Another
example concerns the pig, Nyanzachoerus, which is known from the peat bed on
226 ANNALS OF THE SOUTH AFRICAN MUSEUM
the basis of only a few isolated teeth and bones, whereas at East Stream the
remains of at least fifteen individuals, some represented by incomplete skeletons,
have been collected. Birds, which are common in both areas, provide another
example. In the East Stream assemblage the most common species is a francolin,
which is a terrestrial bird, but in the peat bed it is rare, while waterbirds are
relatively more common (G. Avery pers. comm.).
Another rather curious difference between the two faunas concerns the
representation of the seal, Prionodelphis capensis. This species is not common in
either of the faunas, but at East Stream it is represented almost exclusively by the
remains of very young individuals, whereas only a few isolated teeth and bones
of adult seals are known from the peat bed. There are also differences in the
coprolite occurrences in the two areas. East Stream is one of the areas in the
mine where the smallest type of coprolite occurs in great numbers, while the
larger type with visible bone and those of ?hyaenas are rare. The only coprolites
known from the peat bed area few specimens of the type which contains visible bone.
A complete analysis of the two assemblages will no doubt provide further
and more precise examples of their similarities and differences, which are
presumably more than just fortuitous.
The potential importance of faunal analysis in determining depositional
environments is illustrated by the fact that there are obvious differences in
faunal assemblages even where deposits are superficially little different or
indistinguishable. Such differences have been observed at the same level in
apparently homogeneous deposits over distances of only a few metres. An
example mentioned earlier was the occurrence in the East Stream area of a
‘channel’ containing a relatively high proportion of fish remains and abraded
bones. A second example concerns a quartzose sand exposure in the vicinity of
the peat bed which was found on screening to contain large numbers of frog
bones, but very few terrestrial vertebrate fossils. In similar deposits elsewhere the
representation of fossils was reversed, with frogs being rare and terrestrial
vertebrates common. The presence of a pond, which left no other obvious
traces, could account for the amphibian-rich occurrence.
Although the question of depositional environments within the Quartzose
Sand Member has received only passing attention during the current phase of
the palaeontological investigation, there is evidence for fossils having accumu-
lated in estuarine, fluvial, marsh, mud flat, pond and floodplain environments.
The Pelletal Phosphorite Member
The Pelletal Phosphorite Member covers a wide area and is much the
thickest of the units in the Varswater Formation, but it is known to include
vertebrate fossils in large numbers only in a relatively limited area, exposures of
which still exist along the more northerly part of the west wall of the mine. These
fossiliferous deposits are informally termed ‘bed 3a’ and reasonably large fossil
samples have been recovered from two exposures, designated ‘bed 3aS’ and
‘bed 3aN’ (Fig. 3).
227
PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA
*(S901}
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fT ay &
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228 ANNALS OF THE SOUTH AFRICAN MUSEUM
The bed 3aS deposits are exposed along the lower mining face (“bottom cut’)
of the west wall, while the bed 3aN deposits are approximately 100 metres
north-north-west on the upper mining face (‘top cut’) at elevations of between
2 and 4 metres higher. The bed 3aN deposits are close to the northerly limit of
economically recoverable phosphate and are only between 2 and 3 metres thick,
of which only the lowest 0,5 to 1 metre is fossiliferous. Some fossils have been
recovered from the uppermost levels of the Pelletal Phosphorite in this area, but
they are not regarded as part of the bed 3aN sample. The bed 3aS deposits are
also between 2 and 3 metres thick in an area sampled by controlled excavations
during 1969 and 1970. In this area fossils were concentrated in three distinct
levels spread over the lowest 1,5 metres of deposit. Bed 3aS deepens in a southerly
direction, where there may be more than three levels of concentration. The
thickness of both beds 3aS and 3aN is considerably less than the 25 metre
maximum development of the Pelletal Phosphorite Member.
In the bed 3aN area the base of this member is marked by a 0,75 metre thick
phosphatic sandstone. The surface of the rock at its most northerly exposure
appears to be fairly smooth and more or less horizontal, while the overlying
deposits are apparently not fossiliferous. A little further south the rock surface
dips markedly to the south-west and it becomes progressively more irregular.
Crevices and potholes filled with coarse sand and gravel are common and the
rock eventually becomes discontinuous. Fossils occur in abundance where the
rock surface is irregular, being concentrated in the irregularities and becoming
progressively less common upwards in the overlying finer-grained sediments.
The latter have a clay component in the more northerly exposures, but this is
absent in the southerly exposures where the deposits which immediately overly
the rock are unconsolidated sands. The ill-defined boundary between these two
types of deposit runs from north-east to south-west. Indications are that there
was a channel of fast-flowing water directed in a south-westerly direction in that
area where the rock surface is irregular and where the overlying deposits are
unconsolidated sands. The clayey-sands apparently formed the northern bank
of the channel.
The nature of the bed 3aS fossiliferous occurrences is essentially similar,
except that in this instance there is no rock horizon at the base of the deposits,
there are no clayey deposits indicating a channel bank and fossils are concen-
trated at more than one level in the deposits. The exact stratigraphic relationship
between bed 3aS and bed 3aN is not known, but the lower elevation and more
southerly situation of the former suggests that it was laid down earlier during the
marine transgression than bed 3aN. There may, however, be a direct link
between the highest of the bed 3aS levels of fossil concentration and bed 3aN.
Indications are that bed 3a was laid down in, or in the direct path of the
river which discharged into the sea most of the sediment making up the Pelletal
Phosphorite Member and that the course of the river shifted northwards as the
sea transgressed. A structure contour map of the base of the Pelletal Phosphorite
Member (Tankard 1975: fig. 3), a modified version of which is reproduced here
PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 229
Vv U T Ss R ,a iP 0 N 4
G2
38
Ez
3
1
FE
34 = ee
30
e—7
QUARRY
18
8
2 eee ae ree Set Se contour interval 2m
as metres
ES 10
Fig. 4. Structure contour map of the base of the Pelletal Phosphorite Member. Arrowed lines
indicate the probable courses of the river at the time that beds 3aS and 3aN were laid down.
(Adapted from Tankard 1975: Fig. 3.)
(Fig. 4), supports the suggestion that the lower course of the river was directed
towards the area where bed 3a is now exposed.
The nature of the bed 3a fossil occurrences, and particularly those of bed
3aN, provide clear evidence for deposition by fast-flowing water. Specimens
trapped by irregularities in the rock surface had protruding parts either com-
pletely abraded away or broken up and the fragments scattered in a south-
westerly direction. In many instances specimens on the rock surface had their
lower parts abraded, apparently by the coarser sediment fraction carried along
the rock surface by the flowing water. Individual elements of single skeletons
were dispersed and in some instances were traced by following connected series
of irregularities in the rock surface.
A feature of the faunas of both beds 3aS and 3aN was the large number of
specimens of the seal, Prionodelphis capensis, which are represented. Several well-
preserved skulls of this species were recovered from bed 3aN, while the several
skulls of the similarly-sized hyaena, Hyaena abronia, found in the same deposits
were not as well preserved or as complete. The more numerously represented
and better preserved remains of the aquatic carnivore suggests that deposition
230 ANNALS OF THE SOUTH AFRICAN MUSEUM
was in a subaqueous environment and that the remains of the terrestrial species
had suffered transport over longer distances. As a general rule the terrestrial
fossils from bed 3a are more fragmented and less well preserved than similar
specimens from some of the Quartzose Sand Member deposits.
While most of the bed 3a fossils probably were deposited subaqueously, the
possibility cannot be ruled out that some subaerial accumulation of material
also took place (e.g. on the river banks and/or sand bars). A ?hyaena coprolite
was recovered from the clayey deposits of bed 3aN, that is, those deposits
regarded as having formed the northern bank of the channel at the time that the
fossils of bed 3aN were being accumulated. The smallest type of coprolite is not
uncommon in bed 3aS, but the majority of these specimens are fragmented, their
condition being in marked contrast to the generally very well preserved specimens
from the Quartzose Sand Member.
The previously stated opinion that some of the fossils from what is now
termed bed 3aS were derived from the Quartzose Sand Member (Hendey 19705:
122), may be relevant to the question of the bed 3aS coprolites. These specimens
could well be part of the derived element of the bed 3aS assemblage, having
hardened sufficiently while incorporated in the Quartzose Sand Member to
survive transport to, and redeposition in the Pelletal Phosphorite Member. On
the other hand, they were in most cases not hard enough to survive the trans-
portation intact.
While there is a strong likelihood that the bed 3aS deposits include fossils
derived from the Quartzose Sand Member, there is no evidence to suggest that
this was the case with bed 3aN. Truncation of the Quartzose Sand Member after
the 30 metre stillstand must have been confined to the early stages of the final
transgression, that is, the time when the lower levels of bed 3aS were being laid
down. The great majority of the bed 3aS coprolites come from the lower levels.
The relatively high elevation of bed 3aN virtually precludes the possibility of it
containing fossils derived from the Quartzose Sand Member.
Occasional fossils have been recovered from exposures of the Pelletal
Phosphorite Member other than beds 3aS and 3aN. Some specimens have been
collected in the west wall area from deposits overlying bed 3a, including a few
from the uppermost level. These specimens were probably also transported into
the area by the river. Their rarity may be due to the river mouth having been
some distance away when these deposits were laid down, the fossils representing
the remnants of occasional carcasses which had floated out to sea from the river
mouth. Specimens have also been collected from several different levels in the
most north-easterly exposures of the Pelletal Phosphorite Member. These
fossils are probably the remains of animals accumulated near the southern side
of the river mouth, while those of bed 3a accumulated in, ahead of and on the
northern bank of the river mouth (Fig. 4).
PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 231
FAUNA
During the past six years large numbers of fossils have been collected from
‘E’ Quarry, most material having come from surface collecting, excavations at
random and screening of deposit, but some also having been recovered in
controlled excavations. The deposits sampled extend over an area of about
28 hectares (70 acres), with specimens having come from all the known fossili-
ferous horizons within the Varswater Formation. There is considerable variation
in the size of assemblages from individual occurrences within the deposits, while
the total assemblages from each of the three members are also of unequal size.
The fauna of the Quartzose Sand Member is the largest and best known, since it
has been exposures of this member which have been the principal focus of
attention during the current phase of the Langebaanweg project (Fig. 5). Mining
of highly fossiliferous Pelletal Phosphorite Member deposits (bed 3a) was
recommenced during 1975 so that there has recently been a substantial increase
in the amount of material collected from this member. Relatively little time has
been devoted to the collecting of fossils from the Gravel Member, exposures of
which are limited.
In terms of the requirements for taxonomic studies, many species, mainly
amongst the smaller vertebrates, are more than adequately represented in
existing collections and in such instances the addition of further material may
be of little or no significance. There are, however, many more species which are
poorly represented and in these instances there is still a real need to build up
sample sizes. Since 1969 the annual additions to the collections have always
included several new records for the site, while there has also been further
identification of material already in the collections. The ‘E’ Quarry fauna is, in
general, comparatively well known, although most of the specimens have still
to be studied in detail.
The following summary accounts of groups represented in the fauna include
references to recent new records, recent and current studies, significant additions
to previously existing species assemblages, as well as other comments on avail-
able material.
Invertebrates
Many of the invertebrate fossils recovered from ‘E’ Quarry have already
been described (Kensley 1972), this material having come from the Gravel
Member, while Tankard (1975) has mentioned other invertebrates from the
Varswater Formation. Additional material is now available, including the first
substantial invertebrate assemblage from the Quartzose Sand Member. This
material is from the tidal mud flat deposit mentioned earlier and includes
marine, freshwater and terrestrial molluscs. Many of the specimens are remark-
ably well preserved, some even retaining traces of their original colour. This
material is being studied by B. Kensley (South African Museum) and P. Nuttall
(British Museum (Natural History)).
ANNALS OF THE SOUTH AFRICAN MUSEUM
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PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 233
Lower vertebrates
No progress has been made in the identification of bony fish, amphibians
and reptiles, but P. A. Hulley (South African Museum) has completed a pre-
liminary investigation of the cartilaginous fish from the Gravel Member
(Table 3). Of particular interest was the fact that the Selachii proved more
diverse than anticipated, with the assemblage being complicated by the presence
of derived specimens (see p. 243).
TABLE 3
A provisional list of the cartilaginous fish from the Gravel Member of the Varswater
Formation, including derived material. (Identified by P. A. Hulley of the South African
Museum.)
SELACHII
Hexanchidae _
Notidanus serratissimus
Carcharhinidae
Carcharhinus melanopterus
Carcharhinus limbatus
Galaeorhinus sp.
Prionace glauca
Negaprion sp. or Hypoprion sp.
Odontaspidae
Odontaspis accutissima
Odontaspis sp. B
Odontaspis sp. C
Otodontidae
Megaselachus megalodon
Carcharodontidae
Carcharodon sp.
Isuridae
Isurus sp.
Squalidae
Squalus sp.
Squatinidae
Squatina africana
Squatina sp.
BATOIDEI
Rajidae
Raja sp.
Trygonidae
Gen. & sp. indet.
Myliobatidae
Myliobatis sp.
Birds
Although the ‘E’ Quarry birds probably constitute the largest late Tertiary
avian assemblage from anywhere in Africa, they have received only superficial
attention, with only one species, a penguin, having been positively identified
(Simpson 1971). The original study of the penguin remains suggested the
presence of a second species and this has now been confirmed by more recently
discovered material, although the second species remains unidentified (Simpson
234 ANNALS OF THE SOUTH AFRICAN MUSEUM
1975). G. Avery (South African Museum) has identified to the family or genus
level about a dozen other birds, but there are many more which are completely
unclassified. The Quartzose Sand Member has been the source of the largest
number and the best preserved of the specimens. New material includes
incomplete skeletons of two large raptors and another belonging to a stork-like
species. The tidal mud flat deposits of the Quartzose Sand Member contain
fragments of bird egg-shell, some of which retain their colour.
Mammals
The mammalian fossils from the ‘E’ Quarry exposures of the Quartzose
Sand and Pelletal Phosphorite Members have been the principal focus of the
current phase of the Langebaanweg project. The number of species recorded
from these deposits has grown steadily over the years and approximately
seventy-five have now been positively or tentatively identified (Table 4). Only
about one-third have been described and even these include a number which are
incompletely classified, largely because of inadequacies in available material. In
some instances newly discovered material has made positive identifications
distinctly possible.
Mammalian microfauna
The updated list of “E’ Quarry mammals differs most markedly from those
previously published by including provisional identifications of many of the
small mammals from the Quartzose Sand Member. This part of the list was
provided by T. N. Pocock (Vanderbijlpark, Transvaal). Of interest is the first
record of a bat from the site. The small mammals of the Pelletal Phosphorite
Member, which are generally represented by more fragmentary material, remain
unstudied. The rodents are to be studied by Craig C. Black (Carnegie Museum,
Pittsburgh).
Primates
Although a cercopithecoid is tentatively recorded, there has been increasing
doubt about its presence in view of the continued lack of positively identifiable
material. Primates feature prominently in the late Cenozoic fossil record of
Africa and their great rarity in, or complete absence from the Varswater
Formation is a perplexing aspect of its fauna. A feature of this fauna is its great
diversity and the situation which exists in respect of so successful a group as the
primates is indeed curious. The situation of Langebaanweg in a coastal environ-
ment at the southern continental extremity is likely to be related to the rarity or
absence of this group. The Quaternary fossil record of the south-western Cape
Province is characterized by a similar dearth of primates.
Carnivores
In contrast with the primates, carnivores are an exceptionally well repre-
sented and diverse group. Fully one third of the Varswater Formation mammals
identified to date are carnivores.
PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 235
TABLE 4
A provisional list of the mammals from the Quartzose Sand and Pelletal Phosphorite Members
of the Varswater Formation.
Quartzose Pelletal
Sand Phosphorite
Member Member
INSECTIVORA
Chrysochloridae
SRVSOCHIOMISSD. 3) 9% 2 42. Oe x eR x x
Soricidae
Myosorex sp. : ' Z : , fs 4 ; >
Suncus sp. ‘ : : . ‘ ; x
Soricidae gen. & sp(p). indet. : : : ; : x
Macroscelididae
Elephantulus sp. . 2 ; : : ; : . x Bc
CHIROPTERA
Vespertilionidae
Eptesicus sp. . - . : : ; : : : Xx
? PRIMATES
? Cercopithecidae
Gen. & sp.indet.. . . : : ‘ : : x
PHOLIDOTA
Manis sp. — 2, os : we ESR SS ve
TUBULIDENTATA
Orycteropus sp... : : : F : : : x x
CARNIVORA
Canidae
Vulpes sp. ee ote Ne er ae Pe ee x
Ursidae
Agriotherium africanum : : : : ; : oe
? Procyonidae
SICH cerape InGcin a Pee ee See x
Mustelidae
Mellivora aff. punjabiensis . : Xx
Mellivorinae gen. & sp. indet. (af. Plesiogulo) ; %
Enhydriodon africanus . : : x
Viverridae
Viverra leakeyi
Viverrinae gen. & sp. indet.
Genetta sp.
Herpestes sp. A
Herpestessp.B.
Herpestinae sp. C .
Herpestinae sp. D .
Herpestinae sp. E .
Hyaenidae
Percrocuta australis : : . ‘ ‘
Hyaenictis preforfex . ‘ 2 é ‘ : : Be
Euryboas sp. nov. .
Hyaena abronia
Hyaenidae sp. B : ' : : 5 : : x
Hyaenidae sp.E . : : , , : : : x
xX XX XK XK XK XK XK
x
x
xX X
x
236 ANNALS OF THE SOUTH AFRICAN MUSEUM
Quartzose Pelletal
Sand Phosphorite
Member Member
CARNIVORA (cont.)
Felidae
Machairodus sp.
Homotherium sp.
Felis sp. (small)
Felis aff. issiodorensis
Felis obscura .
Dinofelis diastemata ; ; s
Carnivora (possibly Lutrinae) gen. & sp. indet.
PINNIPEDIA
Phocidae
Prionodelphis capensis . ; 3 ; ‘ ; F x x
PROBOSCIDEA
Gomphotheriidae
Gen waciSP GES hen ee) Gace ea eee ee x x
Elephantidae
Mammuthus subplanifrons . F ; : ; : x Hi
HYRACOIDEA
Procavia cf. antiqua : ‘ irune : : : x ?
PERISSODACTYLA
Equidae
Hipparion sp. A. : ; 2 : M ‘ : x
Hipparionsp.B. : : : : ; j ; x
Hipparion namaquense . : d F : : ‘ x
Rhinocerotidae
Ceratotherium praecox . : ; F : ; : x ?
ARTIODACTYLA
Tayassuidae
Gen. & sp. indet. . : ; . ; : ; : ><
Suidae
Nyanzachoerus sp(p). . , : : : ; ; x< x
Hippopotamidae
Hippopotamus sp. . A : ‘ : : ; ; x
Giraffidae
Sivatherium sp. ; : : : - ; : : x
Giraffa sp. : : : : : ; ; : ; x
Bobidae
Tragelaphus aff. angasi . ; ; : ; : ; x
Mesembriportax acrae . : — ‘ + eer *
Bovini gen. & sp. indet. : : : : ‘ : x
Reduncini sp. A ‘
Reduncinisp.B .
Alcelaphini sp. A .
Alcelaphini sp. B : 3 ; : ;
Raphicerus sp. : ; : : : ; ; 2 x
Gazella aff. vanhoepeni . : = : : :
2Ovibovini gen. & sp. indet. : : . , 2 x
LAGOMORPHA
Gen. & sp. indet. . : ; , ; ; : ; se x
xX X ~~ xX
x X X
x &X
x X KX X K XK K K XK
PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 237
Quartzose Pelletal
Sand Phosphorite
Member Member
RODENTIA
Bathyergidae
Bathyergus sp. a é i ! . 2 F : x x
Cryptomys sp. - : : : : : : : Xx
Hystricidae
Gen. & sp. indet. . : ; : : ? : : x
Muscardinidae
Graphiurus sp.
Cricetidae/Muridae
Aethomys sp. A
Aethomys sp. B
Mus sp. A
Mus sp. B
Rhabdomys sp. :
Otomyinae gen. & sp. nov. .
Mystromys sp. A
Mystromys cf. darti
Mystromys cf. hausleitneri
Desmodillus sp.
Dendromus sp.
Steatomys sp. : : : : : ‘ :
Rodentia gen. & spp. indet. : : : : : : x
x
x XK XK XK XK K KK XK XK XK
CETACEA
Gen. & spp. indet. PMS.) Son a ae Ae x x
By far the best represented species is the seal, Prionodelphis capensis
(Hendey & Repenning 1972). At the time that it was described, the species
assemblage was already unusually large for a fossil phocid and since then many
new specimens have been collected. P. capensis is evidently the best represented
fossil phocid in the world. New material includes several nearly complete skulls
(Fig. 6), parts of many more and hundreds of postcranial bones. Most of the
material is from the Pelletal Phosphorite Member, the newest and best specimens
having come from bed 3aN.
In view of the environments of deposition in the Varswater Formation, other
aquatic carnivores are surprisingly rare. Only one specimen belonging to the
otter, Enhydriodon africanus, is known, although the assemblage does include one
unidentified carnivore which might also have been an otter (Hendey 1974a).
Until recently Canidae were known only on the basis of a few isolated teeth
belonging to a small species and, as with the primate, there had been a growing
doubt about identification. This doubt was dispelled by the discovery of a skull
and associated postcranial bones belonging to the species. The skull has
characters which indicate that the animal concerned was a fox (Vulpes sp.). It is
known only from the Pelletal Phosphorite Member.
Another of the carnivores from this member, the bear, Agriotherium
africanum (Hendey 19726), remains one of the rarer elements in the fauna,
although some additional postcranial bones were found recently.
238 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 6. Dorsal, lateral and ventral views of a skull of the seal, Prionodelphis capensis
(SAM-PQ-L 31976) from bed 3aN, Pelletal Phosphorite Member, ‘E’ Quarry. Scale in
centimetres.
PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 239
Agriotherium was an unexpected addition to the ‘E’ Quarry fauna, being the
first African record of the subfamily to which it belongs. Another similarly
‘exotic’ species was recently tentatively identified, this time a procyonid. The
Procyonidae were widely distributed in Eurasia and North America during the
late Tertiary, but have not previously been recorded from Africa. The Lange-
baanweg procyonid, if that is indeed what it is, may prove to be an enduring
problem since it is poorly represented, with the only specimens having come
from an area now covered by a mine dump. The specimens in question are
fragments of the skull of an immature individual and parts of the hindfoot of an
adult.
The Mustelidae include another of the new records for the site and this
species is as exotic as the Agriotherium and ?procyonid in terms of the known
distribution of its closest relatives. The new species is a giant form whose size
approaches that of Megalictis, from the North American Miocene, which is the
largest of all known mustelids. It belongs to a group of wolverine-like carnivores
which includes Megalictis and which has previously been recorded only in
Eurasia and North America. The presence in Africa of a member of this group is
in a sense even more unexpected than the presence of the bear and ?procyonid
since, unlike them, the giant wolverines are not recorded from southern Asia,
which is the Eurasian region with the greatest faunal resemblances to Africa.
The giant wolverine and ?procyonid, which are both from the Quartzose
Sand Member, join the list of Langebaanian species that belong to groups with
an essentially Eurasiatic (and North American) record (Hendey 1974a: 61).
Apart from the otter, or otters, and the giant wolverine, the only other
mustelid known from ‘E’ Quarry is a small, poorly represented species of
Mellivora (Hendey 1974a), one additional specimen of which was recently
collected from the Pelletal Phosphorite Member.
The Viverridae is the most diverse of the carnivore families known from the
Varswater Formation. Previously only one civet (Viverra leakeyi) had been
recorded, but new material indicates that there is at least one other species as
well. The available civet material is problematical largely because it is so frag-
mentary. A large number of new specimens belonging to smaller viverrids have
been discovered. Most of the material belongs to herpestines, but a small genet
is also represented. When the herpestines were first studied (Hendey 1974a) only
two species were recognized (Herpestes spp. A and B) and they are now the most
commonly represented viverrids in the ‘E’ Quarry assemblage. The new material
includes a few specimens which apparently belong to three additional species
(Hendey 19746: 157).
The most common of the larger terrestrial carnivores from the site are
hyaenas, which now include one additional record, an apparently new species of
Euryboas. The more advanced species of this genus were the long-legged and
sharp-toothed ‘hunting hyaenas’ which occurred in Africa and southern Europe
early in the Pleistocene (Hendey 1975). Four hyaena species had previously been
recorded, namely, Percrocuta australis, Hyaena abronia, hyaenid species B and
240 ANNALS OF THE SOUTH AFRICAN MUSEUM
Hyaenictis preforfex, while a fifth, hyaenid species E, was tentatively identified
(Hendey 1974a). The status of the latter remains unresolved. Additional speci-
mens, readily identifiable with P. australis, H. abronia and Species B, have been
discovered, but H. preforfex is still only known from the remains of a single, aged
individual. A detailed study of the hyaenid material now available is warranted.
H. abronia, the best represented of the species, may prove particularly useful in
resolving the problem of whether or not the Quartzose Sand and Pelletal
Phosphorite Members are substantially different in age (vide infra).
The Felidae also include a recent new record, a small wildcat-sized species
from the Quartzose Sand Member. Although specimens in addition to those
already described have been discovered, only the false sabretooth, Dinofelis
diastemata, is reasonably well represented.
Proboscideans
The Varswater Formation has yielded relatively few proboscidean specimens
and those that have been found are generally fragmentary. The only exception
is the incomplete skeleton of an elephant from the East Stream exposures of
the Quartzose Sand Member (Maglio & Hendey 1970; Hendey 1974a: 349).
Remains of an unidentified gomphothere occur more commonly than those of
the elephant, Mammuthus subplanifrons.
Perissodactyls
Only one rhinoceros, Ceratotherium praecox, is known from the Varswater
Formation (Hooijer 1972). It is perhaps the most commonly represented of the
larger mammals from the Quartzose Sand Member, but only a few fragmentary
specimens from the Pelletal Phosphorite Member may belong to this species.
The situation in respect of the Equidae is more complex. The material from
the Quartzose Sand Member is here regarded as belonging to two species of
Hipparion, an opinion rejected by Hooijer (in preparation). A few specimens
thought to be from the Pelletal Phosphorite Member were excluded from
Table 4 because of uncertainties about their provenance. Otherwise the only
significant specimen from this member is an incomplete lower dentition from the
uppermost level of the deposits in the bed 3aN area. Hooijer (in preparation)
has referred it to Hipparion namaquense Haughton, 1932. It is notable in being
the only record of this species in the Langebaanweg area. Also no other speci-
mens from the same horizon have yet been positively identified.
The Equidae, perhaps more than any other group represented in the various
fossil occurrences near Langebaanweg, are potentially important in resolving the
problems of relative age and stratigraphic relationships of the deposits in which
they occur. For example, although the Gravel Member in ‘E’ Quarry has
produced few useful terrestrial vertebrate fossils, one interesting exception is an
incomplete equid tooth, which, together with two teeth from the same horizon
in ‘C’ Quarry, is tentatively identified with the Miocene species, Hipparion
primigenium. Hooijer (in preparation) also rejects this identification, but if the
PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 241
material does indeed represent a Miocene species, it must have been reworked
from the underlying Saldanha Formation. This formation would therefore be
younger than 12,5 m.y. B.P., since before this date Hipparion was not present in
Africa, or elsewhere in the Old World (Hooijer 1975).
The Baard’s Quarry assemblage includes both Hipparion and Equus, the
latter indicating that at least part of the fauna is younger than that from the
Varswater Formation. The Hipparion material from Baard’s has recently proved
to be as controversial as that from ‘E’ Quarry, but the assemblage is here
regarded as being comprised of two species, the more commonly represented
being distinct from the ‘E’ Quarry hipparions. Earlier it had been indicated that
only one Hipparion was represented in the Langebaanweg occurrences (Hendey
1972a; 1974a), but this is now discounted. The various issues relating to the
Langebaanweg equids should eventually be resolved satisfactorily.
Artiodactyls
Perhaps the most unexpected of the recent new records from ‘E’ Quarry is
that of a peccary. Previously the species concerned had been regarded as a
miniature pig (Hendey 1974a: 47), but additional material from bed 3aN led to
the revised identification being suggested by both A. W. Gentry (British Museum
(Natural History)) and H. B. S. Cooke (Dalhousie University) (pers. comm.).
This is the first African record of the family Tayassuidae and it is also the most
recent Old World occurrence. The nearest peccary record in both a geographic
and temporal sense is the Miocene Pecarichoerus orientalis from the Siwalik
Hills of India (Colbert 1933). The Langebaanweg peccary, which is one of the
smallest ever recorded, was the subject of a recent preliminary study (Hendey
in press).
Nyanzachoerus is the only pig which occurs in the Varswater Formation and
it is known from most of the fossiliferous exposures of the Quartzose Sand
Member, having been particularly common in the East Stream area. The species
from this member is remarkably well represented, the available assemblage being
larger than those of previously described members of this genus. Recently
Nyanzachoerus was recorded from the Pelletal Phosphorite Member for the first
time. The specimen concerned, a fragmented and incomplete skull, differs in
some respects from the Quartzose Sand Member specimens and probably
represents a second species. Although not yet studied in detail, the Quartzose
Sand Member Nyanzachoerus has already proved useful in the relative dating of
the deposits (Hendey 1973), and the Pelletal Phosphorite Member species
promises to be equally useful in this respect (see p. 244).
The apparent absence of hippopotamus from the ‘E’ Quarry fauna has
previously given rise to comment, since the depositional environments of the
Varswater Formation were such that this animal might have been expected to
occur quite commonly (Hendey 1974a: 48). A few fragmentary hippo remains
were found for the first time during 1975 in the bed 3aN exposures of the
Pelletal Phosphorite Member. The fact that hippos are rare in this member and
242 ANNALS OF THE SOUTH AFRICAN MUSEUM
that they are still not recorded from the extensively sampled Quartzose Sand
Member has yet to be satisfactorily explained.
The ‘E’ Quarry Giraffidae are fairly well represented by elements of the
postcranial skeleton, but cranial material is rare and invariably fragmented and
incomplete. Some of the best giraffid specimens, belonging to both Sivatherium
and Giraffa, have come from bed 3aN. The material is being studied by
J. M. Harris (Kenya National Museum).
One additional bovid species, a reduncine, was recently recorded from
‘E’ Quarry. Curiously, Bovidae are not as well represented at this site in terms of
numbers of individuals as they are at some of the Pleistocene fossil occurrences
in the region. In addition, there are relatively few species recorded from the
Quartzose Sand Member, which is otherwise remarkable for the diversity of
species represented. On the other hand, certain deposits tend to include
individuals of particular species in numbers which are disproportionately high
in terms of their bovid assemblages as a whole. For example, the boselaphine,
Mesembriportax acrae (Gentry 1974), is by far the most commonly occurring
bovid in the Quartzose Sand Member, while in bed 3aS it is alcelaphines which
are abundant. At least some of the Quartzose Sand Member boselaphine
remains are believed to have accumulated sub-aerially, indicating that this
species was an inhabitant of the area now exposed in the mine. Gentry (1974)
suggested that the boselaphine was an open woodland species and this sort of
habitat may well have existed in the immediate vicinity of the old estuary. The
situation in respect of the Pelletal Phosphorite alcelaphines was probably quite
different. Many, and perhaps all of the specimens were apparently washed to
their final resting-places by the river and their carcasses may have originated
upstream where the river crossed open plains, the probable preferred habitat of
the alcelaphines.
Other mammals
Other terrestrial mammals recorded from the Varswater Formation include
a pangolin, an aardvark, a dassie (hyrax) and a porcupine. All are rare. Cetacea
are more common, but are represented mainly by undiagnostic postcranial
bones. Both whales and dolphins occur, remains of the latter having been found
for the first time during 1975.
The porcupine is known only from a fragmented skull of a large and
unidentified species which is currently being studied by Judy M. Maguire
(Bernard Price Institute for Palaeontological Research). This specimen was
found in the Quartzose Sand Member shortly after publication of a comment on
the supposed absence of porcupines from the Varswater Formation (Hendey
1974a: 42).
The dassie, Procavia cf. antiqua, was originally identified on the basis of
very fragmentary specimens, but more and better material is now available.
The most complete specimens are from easterly exposures of the Quartzose
Sand Member.
PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 243
The aardvark is amongst the most poorly represented of the ‘E’ Quarry
mammals and until recently the same applied to the pangolin. The situation in
respect of the latter has been slightly improved by the discovery of an incomplete
skeleton, including parts of the skull, much of the tail and parts of all four limbs.
FLORA
Prior to the exposure of the peat bed the only botanical remains recovered
from the ‘E’ Quarry exposures of the Varswater Formation were some fossil
root fragments from the Quartzose Sand Member. A series of samples from the
peat bed submitted to the Institute for Environmental Sciences at the University
of the Orange Free State included one which was pollen-rich. Approximately
92 per cent of the sporomorphae belonged to one unidentified taxon (E. M.
van Zinderen Bakker pers. comm. to A. J. Tankard). This may represent a
locally abundant marsh plant. Interestingly, both tree and grass pollens were
identified and this lends support to the earlier suggestion that these vegetation
types must have been present in the area at the time that the deposits were laid
down (Hendey 1973).
During 1975 two boreholes were sunk from the floor of the mine into
deposits underlying the Varswater Formation. They intersected two peat
horizons which are evidently part of the Miocene Saldanha Formation. These
peats contain both visible and microscopic plant remains.
DATING
A Pliocene age for the Varswater Formation, with an inferred chronometric
date of 4-5 m.y. B.P., is still accepted, but there have been new developments
concerning the dating of fossils from the Gravel Member and the age difference
between the Quartzose Sand and Pelletal Phosphorite Members.
The Gravel Member is composed largely of an abraded and fragmented
phosphatic rock which is known to contain bone fragments and which is
undoubtedly pre-Pliocene in age. The rock, and other deposits with which it
may have been associated, was eroded during the early stages of the Pliocene
marine transgression (Fig. 2). The possibility that some fossils might have been
reworked from the deposits truncated by the transgression has been recognized,
but was not substantiated until a study of the Gravel Member Selachii by
P. A. Hulley revealed that some specimens are evidently of pre-Pliocene age.
The derived fossils may also include the few isolated Hipparion teeth referred to
earlier (see p. 240). The derived pre-Pliocene element in the Gravel Member
fossil assemblage is likely to make up but a small part of the assemblage as a
whole.
Still unresolved is the question of the time taken for the Varswater Forma-
tion to accumulate. There were clearly intervals of time between the deposition
244 ANNALS OF THE SOUTH AFRICAN MUSEUM
of fossils at different levels in the succession, but so far as is known the only one
which may have been of sufficient duration to be palaeontologically significant
was that between the deposition of the Quartzose Sand and Pelletal Phosphorite
Members. The original opinion that the faunas of the two members were
broadly contemporaneous (Hendey 19705) was based on the fact that they co
have species in common and on the belief that differences between them were
due simply to a general dissimilarity in the modes and environments of
deposition of the fossils.
On the other hand, the Pelletal Phosphorite Member does postdate the
Quartzose Sand Member and it has been recognized that the time factor may
have been significant enough to be reflected in the characteristics of individual
species represented in the succession and to have influenced the overall compo-
sition of the two faunas. Previous studies on species common to the two faunas
have provided no conclusive evidence of significant evolutionary changes.
For example, the suggestion that the Pelletal Phosphorite Viverra leakeyi might
be a more advanced variety of the same species from the Quartzose Sand Member
(Hendey 1974a: 81) has still not been substantiated. In this and other instances
comparisons have been complicated mainly by inadequacies in the available
material.
The most important indication to date that the two faunas may be separated
by a substantial period in time came with the discovery of the first Nyanzachoerus
specimen from the Pelletal Phosphorite Member. The new specimen is in
certain respects more advanced than the Quartzose Sand Member Nyanzachoerus
and if there was an ancestor/descendant relationship between the two forms,
their difference in age may be of the order of several hundred thousand years.
Although comparisons with Nyanzachoerus species recorded elsewhere are
‘complicated by the uniqueness of the ‘E’ Quarry forms’, indications are that
the one from the Pelletal Phosphorite is unlikely to be less than 4 m.y. old, while
that from the Quartzose Sand dates back probably no more than 5 m.y.
Now that more material from the Pelletal Phosphorite Member is becoming
available, the potential for meaningful comparisons between the two main
faunas from ‘E’ Quarry is greatly increased.
Relative dating of the ‘E’ Quarry fossils and deposits has been based on
comparisons of some mammals with their counterparts at various localities in
East Africa. The fact that this dating is neither very secure nor very precise is
due to the small number of species which have been appropriate for such
comparisons. It is increasingly apparent that the Langebaanweg fauna is a good
deal more unique than had hitherto been supposed and although individual
species do have much in common with their contemporaries further north, they
are not always identical. If the fauna does indeed include local endemics, and
also perhaps late survivors of lineages which were already extinct elsewhere, the
faunal dating of the occurrences will become complicated and less satisfactory.
PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 245
CONCLUSIONS
The ‘E’ Quarry fossil occurrences, which are the most important in the
Langebaanweg area, are significant for a variety of reasons. There are very few
recorded sites in southern Africa which have yielded vertebrate fossils of
comparable age and the assemblages from such sites are minuscule by com-
parison. Consequently the only substantial information on the nature of the
subcontinental vertebrate fauna of 4-5 million years ago is derived from the
‘E’ Quarry record. The nearest important fossil occurrences which are broadly
contemporaneous are in East Africa, about 4 500 kilometres away (e.g. Vogel
River Series, Lower Kaiso Formation, Mursi Formation of the Omo Group,
Kanapoi/Lothagam and Kubi Algi at East Rudolf). However, not even these
occurrences have produced assemblages so large and so diverse as that from
‘E’ Quarry. Elsewhere in Africa the Pliocene fossil record is either poor or non-
existent, so Langebaanweg is an especially important source of information on
the animal life of this epoch not only in a local sense, but for Africa as a whole.
It has the additional merit of being the only important Pliocene occurrence on
the continent where both marine and terrestrial faunas are represented.
Certain of the ‘E’ Quarry species assemblages are already impressively large
and, since collecting is continuing, they are still growing. The preservation of
specimens is generally very good, all skeletal elements are represented (and
collected) and unequivocal associations of cranial and postcranial material are
not uncommon. It should therefore ultimately prove possible to provide
comprehensive definitions of many species and this may well assist in resolving
identification problems in the smaller assemblages from other African sites.
The species diversity is remarkable by any standards and as a result the
fauna as a whole will be better known than those of many other sites where
important elements may be lacking (e.g. vertebrate microfauna, birds, aquatic
species).
The ‘E’ Quarry fauna, like many others in southern Africa, has so far been
dated only in a relative sense and in this respect it is more problematical than
many from East Africa. However, the Langebaanweg occurrences are amongst
the very few of any importance in southern Africa where generally acceptable
geological dating is also possible.
Perhaps the biggest drawback of the site lies with its situation at the
southern continental extremity, far from the main focus of African late Cenozoic
palaeontological investigation (i.e. East Africa). On the other hand, this may
ultimately invest it with a particular interest since, as the contemporary East
African fauna becomes better known, similarities and differences of
zoogeographic significance may emerge.
ACKNOWLEDGEMENTS
During the past two years a number of persons have contributed directly
and indirectly to the furthering of the Langebaanweg research project and I am
246 ANNALS OF THE SOUTH AFRICAN MUSEUM
greatly indebted to them for their interest and contributions. Particular thanks
are due to Mr G. Avery, Dr P. A. Hulley and Dr B. Kensley (South African
Museum), Dr A. W. Gentry (British Museum (Natural History)), Dr J. M. Harris
(Kenya National Museum), Dr D. A. Hooijer (Rijksmuseum van Natuurlijke
Historie), Dr R. G. Klein (University of Chicago), Mr T. N. Pocock (Vanderbijl-
park) and Dr G. G. Simpson (The Simroe Foundation and University of
Arizona). Dr A. J. Tankard (South African Museum), who has undertaken the
geological study at Langebaanweg, patiently discussed the geology of the
deposits with the result that several points concerning the depositional environ-
ments of the fossils were clarified. Mr G. Benfield, the Mine Superintendent at
Langebaanweg, has been unfailingly helpful and, together with other mine
employees, has kept the project viable.
The fieldwork at Langebaanweg is financed by the South African Council
for Scientific and Industrial Research. The undertaking is aided on site in many
ways by Chemfos Ltd., a subsidiary of Samancor. The South African Air Force
has provided aerial photographs. The Wenner-Gren Foundation for Anthropo-
logical Research (New York) provided the vehicle used for the fieldwork
(Grant no. 2752-1834). The assistance of these organizations is gratefully
acknowledged.
REFERENCES
COLBERT, E. H. 1933. An Upper Tertiary peccary from India.—Am. Mus. Novit. 635: 1-9.
GENTRY, A. W. 1974. A new genus and species of Pliocene boselaphine (Bovidae, Mammalia)
from South Africa.— Ann. S. Afr. Mus. 65: 145-188.
HENDEY, Q. B. 1970a. A review of the geology and palaeontology of the Plio/Pleistocene
deposits at Langebaanweg, Cape Province.— Ann. S. Afr. Mus. 56: 75-117.
HENDEY, Q. B. 1970b. The age of the fossiliferous deposits at Langebaanweg, Cape Province. —
Ann. S. Afr. Mus. 56: 119-131.
HENDEY, Q. B. 1972a. Further observations on the age of the mammalian fauna from Lange-
baanweg, Cape Province. —Palaeoecol. Afr. 6: 172-175.
HENDEY, Q. B. 19726. A Pliocene ursid from South Africa.— Ann. S. Afr. Mus. 59: 115-132.
HENDEY, Q. B. 1973. Fossil occurrences at Langebaanweg, Cape Province.— Nature, Lond.
244: 13-14.
HENDEY, Q. B. 1974a. The late Cenozoic Carnivora of the south-western Cape Province. — Ann.
S. Afr. Mus. 63: 1-369.
HENDEY, Q. B. 19746. Faunal dating of the late Cenozoic of southern Africa, with special
reference to the Carnivora.— Quat. Res. 4: 149-161.
HENDEY, Q. B. 1975. Relationships of North American hyaenas.—S. Afr. J. Sci. 71: 187.
HENDEY, Q. B. In press. Fossil peccary from the Pliocene of South Africa. Science.
HENDEY, Q. B. & REPENNING, C. A. 1972. A Pliocene phocid from South Africa.— Ann. S. Afr.
Mus. 59: 71-98.
Hoower, D. A. 1972. A late Pliocene rhinoceros from Langebaanweg, Cape Province. —
Ann. S. Afr. Mus. 59: 151-191.
Hoover, D. A. 1975. The hipparions of the Baringo Basin sequence.— Nature, Lond. 254:
211-212.
KENSLEY, B. 1972. Pliocene invertebrates from Langebaanweg, Cape Province.— Ann. S. Afr.
Mus. 60: 173-190.
MAGLIo, V. J. & HENDEY, Q. B. 1970. New evidence relating to the supposed stegolophodont
ancestry of the Elephantidae.—S. Afr. archaeol. Bull. 25: 85-87.
MELLETT, J. S. 1974. Scatological origin of microvertebrate fossil accumulations.— Science
185: 349-350,
PLIOCENE FOSSIL OCCURRENCES IN LANGEBAANWEG, SOUTH AFRICA 247
Simpson, G. G. 1971. Fossil penguin from the late Cenozoic of South Africa.— Science 171:
1144-1145.
Simpson, G. G. 1975. Notes on variation in penguins and on fossil penguins from the Pliocene
of Langebaanweg, Cape Province, South Africa.— Ann. S. Afr. Mus. 69: 59-72.
SmNGER, R. 1961. The new fossil sites at Langebaanweg, South Africa.—Curr. Anthrop. 2:
385-387.
SoUTH AFRICAN CODE OF STRATIGRAPHIC TERMINOLOGY AND NOMENCLATURE. 1971. Trans.
geol. Soc. S. Afr. 74: 111-131.
TANKARD, A. J. 1974a. Petrology and origin of the phosphorite and aluminium phosphate rock
of the Langebaanweg—Saldanha area, south-western Cape Province.— Ann. S. Afr. Mus.
65: 217-249.
TANKARD, A. J. 1974b. Chemical composition of the phosphorites from the Langebaanweg—
Saldanha area, Cape Province.—Trans. geol. Soc. S. Afr. 77: 185-190.
TANKARD, A. J. 1975. Varswater Formation of the Langebaanweg-Saldanha area, Cape
Province. —Trans. geol. Soc. S. Afr. 77: 265-283.
TANKARD, A. J. In press. The Saldanha Formation: a Neogene transgressive complex. — Trans.
geol. Soc. S. Afr.
6. SYSTEMATIC papers must conform with the International code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. n., sp. n., comb. n.,
syn. n., etc.
An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.
Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (figs 8a—b).
Nucula largillierti Philippi, 1861: 87
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
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dash, not comma, separates consecutive numbers
Synonymy arrangement according to chronology of bibliographic references, whereby
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not acceptable.
In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:
Holoty,
SAM-A13535 i in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach,
Port Elizabeth (33.51S, 25.39E), collected by A. Smith, 15 January 1973.
Note standard form of writing South African Museum registration numbers and of date.
7. SPECIAL HOUSE RULES
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(a) The tease Maps and Tables of the paper when referred to in the text
e.g. . the Figure depicting C. namacolus .
. in C. namacolus (Fig. 10) .
(b) The ae of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A. L. du Toit
Von Huene _—ibut:~ F. von Huene
(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian
Punctuation should be loose, omitting all not strictly necessary
Reference to the author should be expressed in the third person
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book or article, such as
“Revision of the Crustacea. Part VIII. The Amphipoda.’
Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.
Q. B. HENDEYs
THE PLIOCENE FOSSIL OCCURRENCES IN
| ‘E? QUARRY
LANGEBAANWEG, SOUTH AFRICA
/OLUME 69 PART 10 JUNE 1976 ISSN 0303-2515
> ~ A 4 - Copetouor MUS. COMP. ZOOL.
; ) LIBRARY
AUG 14 5 4976
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(a) Author’s name and year of publication given in text, e.g.:
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Examples (note capitalization and punctuation)
BuULLouGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan,
FIscHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques.—J. Conch., Paris 88: 100-140.
FIscHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines.— Archs
Zool. exp. gén. 74: 627-634. c
Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. —
Ann. Mag. nat. Hist. (13) 2: 309-320. .
Konn, A. J. 19605. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. —
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
TuHrELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer.— Denkschr. med.-naturw. Ges. Jena 16: 269-270. Ag
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 69 Band
June 1976 Junie
Part 10 #£Deel
NOTES ON THE ADDUCTOR JAW MUSCULATURE
OF VENJUKOVIA, A PRIMITIVE ANOMODONT
THERAPSID FROM THE PERMIAN OF THE USS.S.R.
By
HERBERT R. BARGHUSEN
Cape Town Kaapstad
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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap
NOTES ON THE ADDUCTOR JAW MUSCULATURE OF VENJUKOVIA,
A PRIMITIVE ANOMODONT THERAPSID FROM THE PERMIAN
OF THE U:S.S.R.
By
HERBERT R. BARGHUSEN
Departments of Anatomy and Oral Anatomy, University of Illinois Medical
Center, Chicago, Illinois 60680
(With 4 figures)
LMS accepted 9 March 1976]
ABSTRACT
The primitive anomodont Venjukovia, from Zone II of the Russian Permian, has been
thought to bridge the morphological gap between dinocephalians and dicynodonts. Much
of the pattern of adductor jaw musculature in Venjukovia conforms closely to that found in
dicynodonts and is thereby consistent with the hypothesis that these are closely related forms.
The most important similarity is the probable presence in Venjukovia of a distinctive lateral
division of the external adductor. The presence of this division is a derived character previously
known only in dicynodonts. However, as far as can be determined, similarities between
Venjukovia and dinocephalians only involve joint possession of a primitive therapsid arrange-
ment of various other parts of the jaw musculature. Therefore, the muscle pattern does not
provide evidence of a closer relationship with dinocephalians than with other primitive
therapsid groups. Moreover, in contrast to primitive dinocephalians (brithopodids), Venju-
kovia lacks an extensive area of origin of the external adductor from the dorsal surface of the
temporal roof. The absence of this specialized area of origin suggests (tentatively) that Venju-
kovia retained the primitive therapsid condition of the temporal roof and that the lines leading
to Venjukovia and dicynodonts on the one hand and to dinocephalians on the other diverged
before the primitive dinocephalian condition was achieved.
CONTENTS
PAGE
Introduction . . « «. . + « 249
Maténalieore sso fs 2 = re 250
Adductorjaw musculature . . . 252
Gonclusions.= sege.00 & a se 2 ee
Acknowledgements . . . . . 259
References: 5 . = . « « « 209
Abbreviations aN Sa. Wat Fee) 8 200
INTRODUCTION
The Russian anomodont Venjukovia has long attracted attention because
it shows structural resemblances to tapinocephalid dinocephalians on the
one hand (Efremov 1940) and to dicynodonts on the other (Watson 1942,
1948). Efremov noted general resemblances between Venjukovia and tapino-
cephalids in the structure of the teeth, face and palate. Watson agreed but
also noted that the lower jaw of Venjukovia shows many features which are,
in essence, identical with those in dicynodonts. Romer (1956) regarded Venju-
Kovia as seeming to bridge the morphological gap between dinocephalians
249
Ann. S. Afr. Mus. 69 (10), 1976: 249-260, 4 figs.
250 ANNALS OF THE SOUTH AFRICAN MUSEUM
and dicynodonts. It is not the purpose of this paper to extensively analyse
the phylogenetic position of Venjukovia; the issues involved are complex
and demand a thorough phylogenetic analysis of therapsids in general which
is the subject of a paper in preparation. However, the present study, which
involves a partial reconstruction of adductor jaw musculature in Venjukovia,
has obvious potential for defining character states useful in such an analysis.
In addition, some general but very tentative conclusions concerning the possible
relationships between Venjukovia, dinocephalians and dicynodonts can be
advanced based on a comparison of their patterns of adductor jaw musculature.
Accordingly, such comparisons will be made in so far as it is possible to do
so in this paper.
A rationale for jaw muscle reconstruction in synapsid reptiles and a
reconstruction of this musculature (based on a reptilian model) in the pelycosaur
Dimetrodon has already been provided (Barghusen 1973). The evaluation of
direct evidence of muscle attachment as well as the arguments concerning the
distribution of individual jaw muscles presented for Dimetrodon also apply
to the reconstructions made here. For this reason, the reader is referred to
Barghusen (1973) for clarification of the issues involved. In addition, by virtue
of the phylogenetic position of Dimetrodon within the pelycosaur family from
which therapsids were derived, the pattern of jaw musculature which it shows
constitutes the pattern primitive to the evolution of this musculature in therapsid
reptiles. Therefore, comparisons of the reconstructed musculature in Venju-
kovia will also be made with that of Dimetrodon exemplifying the pre-therapsid
arrangement from which the musculature of Venjukovia was derived.
MATERIAL
The reconstruction of adductor jaw musculature presented here for Venju-
kovia is based on information gained from a skull of V. prima (PIN 2793/1)
and three lower jaws of V. invisa (PIN 157/1111, 157/1112, 157/5) housed in
the Palaeontological Institute, U.S.S.R. Academy of Sciences, Moscow. The
skull is exceptionally well-preserved except posteriorly where much of the bone
forming the posterior margin of the lateral temporal fenestra, the posterior
root of the zygomatic arch, and the occiput has flaked from the matrix. Never-
theless, the bone which is present, and impressions of bone in the matrix,
clearly indicate that the outlines of the lateral temporal fenestra, temporal
fossa, zygomatic arch, and the position of the quadrate correspond to that
shown in Figures 1A and 1C. Matrix has not been cleared from the temporal
fossae or the orbits. For this reason it is impossible to estimate the area of
origin of m. adductor mandibulae internus pseudotemporalis, m. adductor
posterior, and, if present, m. adductor mandibulae internus pterygoideus
anterior. Consequently, these muscles will not be considered in this recon-
struction. However, the medial surface of the lower jaw (Fig. 1D) has not
departed sufficiently from that of Dimetrodon to suggest that the general areas
NOTES-ON THE ADDUCTOR JAW MUSCULATURE OF VENJUKOVIA 251
Fig. 1. Reconstruction of the skull and lower jaw of Venjukovia.
A. Dorsal view of skull. B. Ventral view of skull with left lower jaw in place. C. Lateral view
of skull and lower jaw. D. Medial view of lower jaw. The areas of origin and insertion of the
external adductor and posterior pterygoideus jaw musculature are indicated by parallel
lines. The outline of the posterior margin of the skull and the position of the quadrate were
determined from bone impressions in the matrix. (Skull reconstructed from PIN 2793/1;
lower jaw reconstructed from PIN 157/1111, 157/1112, 157/5.)
of insertion of these muscles were significantly different from those in
Dimetrodon.
The reconstruction of brithopodid jaw musculature is also based on
specimens housed in the Palaeontological Institute, Moscow. These include an
exceptionally well-preserved skull and lower jaw of Titanophoneus potens
(PIN 157/1). Details of the temporal fossa were also gained from Notosyodon
gusevi (PIN 2505/1).
Much of the information upon which the reconstruction of musculature
in Lystrosaurus is based was collected in 1965 at Yale University from an
acid-prepared skull (SAM-4325). Unfortunately this skull was subsequently
252 ANNALS OF THE SOUTH AFRICAN MUSEUM
destroyed in transit. More recently, information has also been gained from
acid-prepared material examined while the author visited the South African
Museum, Cape Town. This material included a skull and lower jaw of
L. declivis (Nat. Mus. C 403).
ADDUCTOR JAW MUSCULATURE
M. ADDUCTOR MANDIBULAE EXTERNUS
The temporal region of Venjukovia exhibits the posterodorsal enlargement
of both the lateral temporal fenestra and temporal fossa which is characteristic
of most therapsids as opposed to sphenacodontid pelycosaurs (cf. Figs 2A, C).
In dorsal view (Fig. 1A) the temporal fossa is broadly exposed due to the
extensive but as yet incomplete reduction in the width of the temporal roof.
The degree of reduction is comparable to that seen in some dicynodonts (ef.
Emydops, Crompton & Hotton 1967, Fig. 1B) but not as extensive as that
found, for example, in Lystrosaurus (Fig. 3A). The presence of a temporal
crest (Fig. 1A, temp cr) in Venjukovia provides direct evidence that the external
adductor took origin from the lateral face of that part of the postorbital forming
= B
inf mar
MAME (lat) MAME (lat)
Fig. 2. Lateral views of the skull and lower jaw of Dimetrodon, A, hypothetical condition,
B, Venjukovia, C, and Lystrosaurus, D, forming a morphological series which illustrates
stages in the development of the lateral division of the external adductor jaw musculature
characteristic of dicynodonts. In A the zygomatic arch is positioned close to the adducted
lower jaw. In B the zygomatic arch is dorsally displaced creating an access route for that
part of the external adductor originating from the anterior face of the quadratojugal and
quadrate to invade the lateral surface of the squamosal. In C and D the arch is further dis-
placed and the invasion, creating the lateral division of the external adductor, has taken
place; in addition, an area of insertion for the lateral division is established on the dorsolateral
surface of the lower jaw. (A after Romer and Price.)
NOTES ON THE ADDUCTOR JAW MUSCULATURE OF VENJUKOVIA 253
MAME
(med)
att apon
Fig. 3. The skull and lower jaw of Lystrosaurus.
A. Dorsal view of skull. B. Ventral view of skull with left lower jaw in place. C. Lateral view
of skull and lower jaw. The areas of origin and insertion of the external adductor and posterior
pterygoideus muscles are indicated by parallel lines.
the lateral margin of the temporal roof. The establishment of muscle origin
here probably represents an invasion of muscle attachment from the anterior
face of that part of the squamosal forming the posterior wall of the temporal
fossa. This condition, which is a distinct departure from that found in sphena-
codontids, was probably common among primitive therapsids as it is also
found in gorgonopsids, Biarmosuchus, and Eotitanosuchus (personal observa-
tions). The extent of this muscle attachment on the postorbital also approaches
that seen in dicynodonts (e.g. Lystrosaurus, Fig. 3A; also see Crompton &
Hotton 1967; Cluver 1975). Venjukovia, however, does not achieve the special-
ized condition seen in brithopodids, the most primitive known dinocephalians.
In brithopodids, the external adductor took extensive origin from the dorsal
surface of the temporal roof (Fig. 4A; also see Watson 1948; Olson 1962;
Boonstra 1963; Barghusen 1973). The condition in brithopodids contrasts
254 ANNALS OF THE SOUTH AFRICAN MUSEUM
sharply with that found in other primitive therapsids (i.e. gorgonopsids, Eotita-
nosuchus, Biarmosuchus, and ictidorhinids) as well as in Venjukovia. All these
animals retain the sphenacodontid arrangement, in which no adductor jaw
musculature attaches to the dorsal surface of the temporal roof. In contrast
to more primitive dinocephalians, many tapinocephalid dinocephalians (e.g.
Moschops) reverted back to the primitive therapsid and sphenacodontid arrange-
ment in the sense that the attachment of the external adductor to the dorsal
surface of the temporal roof was eliminated. Evidence strongly suggests that
this reversal in tapinocephalids was related to the development of head-butting
behaviour in these animals (Barghusen 1975). Conceivably a reversal may also
have taken place in the ancestry of Venjukovia. However, Venjukovia does not
exhibit any of the specializations for use of the dorsal surface of the head in
butting which are displayed by tapinocephalids and which, if present in Venju-
kovia, would suggest tapinocephalid relationships. At the moment it seems
more likely, therefore, that the condition manifested by Venjukovia is a retention
of the primitive therapsid and pelycosaur condition as there are no obvious
functional reasons to suggest that the absence of muscle attachment to the
dorsal surface of the temporal roof was secondarily derived. If so, Venjukovia
reflects a morphological stage, with regard to the attachment of part of the
adductor musculature, resembling that which must have been antecedent to
the stage reached by the most primitive known dinocephalians. Such a con-
clusion appears to be implicit in Boonstra’s (1963) remarks to the effect that
when the dinocephalian and dicynodont lines diverged, the intertemporal skull
table was broad and the origin of the adductor muscles showed a fairly primi-
tive condition. This conclusion runs counter to suggestions (Efremov 1940)
that Venjukovia was derived from tapinocephalids.
Preparation of the temporal fossa is not complete in the skull of Venjukovia.
However, there are reasons to believe that the undersurface of much of the
retained portion of the temporal roof served for the attachment of the external
adductor. This is expected from the distribution of the attachment of this
muscle in living reptiles. In addition, a depression on the undersurface of the
roof indicates that this was the case in Dimetrodon (see Barghusen 1973) and
a similar scar or depression has been found in all therapsids examined in which
the temporal region was sufficiently prepared and which showed a degree of
development of the temporal roof similar to that of Venjukovia. These therapsids
include Titanophoneus (see Orlov 1958, fig. 21) and Notosyodon among the
dinocephalians and Leontocephalus (see Kemp 1969, fig. 5) among the gor-
gonopsids. This evidence clearly suggests that such an arrangement of muscle
attachment was present prior to and during the evolution of those therapsids
that did not completely eliminate the temporal roof. It is also expected (Barg-
husen 1973) that the dorsal and ventrolateral parts of the posterior wall of
the temporal fossa, including the squamosal and quadrate, served as an area
of origin of the external adductor muscle in Venjukovia. The squamosal portion
of this area of attachment to the posterior wall of the fossa would have served
NOTES ON THE ADDUCTOR JAW MUSCULATURE OF VENJUKOVIA 255
att apon
Fig. 4. The skull and lower jaw of Titanophoneus.
A. Dorsal view of skull. B. Ventral view of skull. C. Lateral view of skull and lower jaw.
The areas of origin of the external adductor and posterior pterygoideus muscles are indicated
by parallel lines. (Outlines after Orlov.)
as the original site from which the invasion of muscle attachment onto the
lateral surface of the postorbital in therapsids, including Venjukovia, took
place.
The posterodorsal tip of the dentary, forming the apex of the coronoid
eminence in Venjukovia, marks the probable site of attachment of a bodenapo-
neurosis (Fig. 1D, att apon) serving for the insertion of much of that part of
the external adductor described above. This function for the coronoid eminence
is indicated by comparisons with living reptiles (Barghusen 1973) and is con-
sistent with direct evidence of tendon attachment in Dimetrodon (Barghusen
1968) and theriodont therapsids. A similar insertion would also have been
present in all dinocephalians (Fig. 4C, att apon) and, as Cluver (1975) has
256 ANNALS OF THE SOUTH AFRICAN MUSEUM
recently pointed out, in dicynodonts, in which the posterodorsal tip of the
dentary marks a site of tendinous attachment (Fig. 3C, att apon) identical in
position to that in Venjukovia.
By far the most important question concerning the external adductor in
Venjukovia is whether or not a lateral division was present comparable to that
found in dicynodonts (Figs 2D, 3B, C). The external adductor jaw musculature
independently established an extensive area of attachment on the zygomatic
arch and exposed lateral surface of the lower jaw at least twice and probably
three times in therapsid history. This definitely occurred in cynodonts (Barg-
husen 1968) and dicynodonts (Watson 1948; Crompton & Hotton 1967;
Cluver 1975) and probably occurred in gorgonopsids (Barghusen 1968; Kemp
1969). In each group the manner in which this muscular modification occurred
is highly distinctive. In cynodonts a laterally bowed zygomatic arch created a
channel through which developing masseter musculature descended from the
temporal fossa to insert on the lateral surface of the dentary; subsequent
events led to the establishment of muscular origin on the entire zygomatic
arch. An invasion also occurred in gorgonopsids but in this case the muscle
attachments were confined to the lateral surface of the angular and the posterior
root of the zygomatic arch. In dicynodants, a channel was created for the
invasion of musculature from the confines of the temporal fossa through the
dorsal displacement of the zygomatic arch relative to the dorsal surface of the
lower jaw (Crompton & Hotton 1967). This invasion resulted in a newly added
lateral division of the external adductor originating from the arch and expanded
lateral surface of the squamosal and inserting on a distinctive lateral shelf of
the dentary (Crompton & Hotton 1967; also see Figs 3B—C). This muscular
development is a dicynodont hallmark, as is the developing masseter muscle
for cynodonts.
Despite the limitations of incomplete preservation posteriorly, the mor-
phology of Venjukovia indicates that this animal closely conforms to, and may
reflect, an initial stage in the development of the condition seen in dicynodonts.
The zygomatic arch is dorsally displaced relative to the jaw articulation. This
created the necessary condition whereby external adductor musculature origi-
nating on the ventrolateral part of the posterior wall of the temporal fossa
(the quadratojugal and quadrate as reconstructed in Dimetrodon (Barghusen
1973)) had free access to invade the lateral surface of the squamosal by a route
passing inferior to the level of the zygomatic arch (Fig. 2). It is also apparent
(Fig. 2) that the zygomatic arch of Venjukovia and dicynodonts was dorsally
displaced to the level of the inferior margin (Fig. 2A, inf mar) of the attachment
of the external adductor to the inner surface of the cheek as reconstructed in
Dimetrodon on the basis of stretch capabilities of muscle (see Barghusen 1968).
This suggests that much of the length of the inferior and medial surfaces of the
arch could have served for the origin of musculature with sufficient fibre length
to allow for a substantial gape (also see Watson’s 1948 discussion of gape in
dicynodonts). Finally, this displacement removes the zygomatic arch from
NOTES ON THE ADDUCTOR JAW MUSCULATURE OF VENJUKOVIA 25)
immediate proximity to the lateral surface of the lower jaw which, thereby,
becomes a potential site of muscle insertion.
The extent of invasion of muscle attachment onto the lateral surface of
the squamosal is impossible to determine in Venjukovia due to insufficient
preservation of the posterior root of the zygomatic arch. However, the presence
of a fossa (Fig. 1C, fossa) excavated into the lateral surface of the surangular
and most posterior part of the dentary, as a departure from the primitive
therapsid condition, strongly suggests that such invasion did take place. The
fossa indicates the incipient development of muscle attachment to the dorso-
lateral surface of the jaw corresponding to the insertion of the lateral division
of the external adductor in dicynodonts (Fig. 3C); the insertion has merely
expanded anteriorly in dicynodonts to include more of the dentary. The fossa
thus indicates the presence of a laterally placed muscle whose area of origin
may well have included the lateral surface of the squamosal. If so, the impression
of the squamosal left on the matrix suggests that the posterior root of the
zygomatic arch was shaped as in Figure 1C and capable of supporting an area
of origin from its lateral surface as is illustrated.
M. ADDUCTOR MANDIBULAE INTERNUS PTERYGOIDEUS (POSTERIOR)
The area of origin of the posterior pterygoid muscle in Venjukovia is
clearly very similar to that reconstructed in Dimetrodon (Barghusen 1973).
A boss on the distal end of the transverse process of the pterygoid as well as
indications of a medial pterygoid crest (Fig. 1B, m pt cr) provide direct evidence
of tendinous attachment in the manner of sphenacodontids, theriodonts, and
brithopodids (Fig. 4B), as well as many living reptiles. The ventral surface of
the quadrate ramus of the pterygoid is also expected to have served as an area
of origin for this muscle mass (Fig. 1B, MAMIPt (post)). The major difference
in area of origin of the posterior pterygoid muscle between Venjukovia, on the
one hand, and Dimetrodon and brithopodids, on the other, is that the transverse
process is less massive and projects anterolaterally in Venjukovia (cf. Figs 1B,
4B). The anterolateral projection approaches (but by no means achieves) the
condition in dicynodonts, in which the transverse process has lost its role in
bracing and controlling the movements of the lower jaw and projects almost
straight forward (Fig. 3B, tp). In this regard, Venjukovia resembles a morpho-
logical stage which was probably antecedent to the condition in dicynodonts.
It is generally assumed that the transverse process of the pterygoid is either
reduced (Romer 1956) or entirely eliminated (Watson 1948; Crompton &
Hotton 1967) in dicynodonts. It appears, however, that this structure is merely
redirected forward and in many cases is well developed. The structure labelled
as the transverse process in Figure 3B has the same general topographic rela-
tionships to the subtemporal fossa and the ectopterygoid (when the latter is
present) as does the structure which is clearly the transverse process in Venju-
kovia. Moreover, direct evidence in one acid-prepared specimen (SAM-4325)
of Lystrosaurus indicates that the relationship to the posterior pterygoid muscle
258 ANNALS OF THE SOUTH AFRICAN MUSEUM
is also the same. The ventral surface of the distal portion of the process is
slightly rugose and striated, suggesting tendon attachment. In addition, a line
of parallel striations (Fig. 3B, str) extends posteromedially from the ventral
edge of the anteriorly directed process to terminate near the level of the basi-
pterygoid joint. The position of this line corresponds to that of the medial
pterygoid crest and also provides direct evidence of probable tendon attachment.
The area of insertion of the posterior pterygoid muscle in Venjukovia
(Fig. 1B) is expected to have included the ventromedial surface of the articular
and probably the anteromedial face of the retroarticular process. This recon-
struction contrasts with many made in the past in which it was assumed that
pterygoideus musculature ran beneath the jaw to insert on the lateral surface
of the main body of the angular medial and posterior to the reflected lamina.
This has been thought to have been the case in sphenacodontid pelycosaurs
and in therapsids in general (see Barghusen 1968, 1973; Crompton & Hotton
1967; Watson 1948). The muscle involved has been given various names and
reconstructed with a number of different areas of origin; however, it is the
posterior pterygoid muscle as reconstructed by Barghusen (1973). While an
insertion on the lateral surface of the angular appears to be valid for sphena-
codontids, its validity is questionable in most therapsids. As pointed out by
Allin (1975), the very close relationship between the reflected lamina and the
lateral surface of the main body of the angular in the vast majority of therapsids,
including brithopodids, Venjukovia, and dicynodonts, argues against the inter-
vention of musculature between these structures. The confinement of muscu-
lature within such a narrow space would cause functional inefficiency since
the amount of muscle is necessarily very small and, as Allin remarks, the
muscle ‘would be unable to shorten without bulging against its skeletal confines
as well as constricting off its own blood supply, although an associated venous
plexus might provide volumetric compensation’. It may be added that if a
venous plexus were present in the space between the lamina and main body
of the angular, the amount of musculature present would have been negligible.
It is more reasonable, as persuasively argued by Allin, to regard this space as
housing an air-filled chamber existing as a tympanic or pharyngeal diverti-
culum and functioning in sound reception.
CONCLUSIONS
The pattern of adductor jaw musculature reconstructed for Venjukovia
suggests that this animal is closely related to dicynodonts—perhaps more
closely related to dicynodonts than to any other known therapsid group.
A close relationship is indicated by: 1. the probable presence of a similar
lateral division of the external adductor in both groups in contrast to the
condition in other therapsids; and 2. the fact that the structure of the area of
origin of the posterior pterygoid in Venjukovia is consistent with the structure
postulated for a theoretical morphological stage transitional to the dicynodont
condition. The muscular morphology of Venjukovia contrasts with that of
NOTES ON THE ADDUCTOR JAW MUSCULATURE OF VENJUKOVIA 259
primitive dinocephalians in that there is no specialized origin of the external
adductor from the dorsal surface of the temporal roof in Venjukovia. Unless
a reversal (i.e. secondary elimination of this area of origin) is postulated,
animals which would be classified as dinocephalians were not involved in the
ancestry of Venjukovia. At the moment, there is no evidence which suggests
that such a reversal took place. The therapsid ancestry of Venjukovia is, the
author believes, an open question.
ACKNOWLEDGEMENTS
I am indebted to the following individuals and institutions for making
fossil material in their care available for study: Dr Peter K. Chudinov of the
Palaeontological Institute, U.S.S.R. Academy of Science, Moscow; Dr Michael
A. Cluver of the South African Museum, Cape Town; and Dr A. W. Crompton,
formerly of Yale University, New Haven, Connecticut. Thanks are also due
to Dr James Hopson for critical commentary on the manuscript.
This research was supported by NSF grant GB-40061.
REFERENCES
ALLIN, E. F. 1975. Evolution of the mammalian ear.—J. Morph. 147: 403-438.
BARGHUSEN, H. R. 1968. The lower jaw of cynodonts (Reptilia, Therapsida) and the evolu-
tionary origin of mammal-like adductor jaw musculature.—Postilla 116: 1-49.
BARGHUSEN, H. R. 1973. The adductor jaw musculature of Dimetrodon (Reptilia, Pelyco-
sauria).—J. Paleont. 47: 823-834.
BARGHUSEN, H. R. 1975. A review of fighting adaptations in dinocephalians (Reptilia, Therap-
sida). — Paleobiology 1: 295-311.
Boonstra, L. D. 1963. Early dichotomies in the therapsids.—S. Afr. J. Sci. 59: 176-195.
Cuuver, M. A. 1975. A new dicynodont reptile from the Tapinocephalus Zone (Karoo System,
Beaufort Series) of South Africa, with evidence of the jaw adductor musculature. — Ann.
S. Afr. Mus. 67: 7-23.
Crompton, A. W. & Hotton, N. 1967. Functional morphology of the masticatory apparatus
‘of two dicynodonts (Reptilia, Therapsida).—Postilla 109: 1-51.
Erremovy, I. A. 1940. Preliminary description of new Permian and Triassic terrestrial verte-
brates from the U.S.S.R.—Trudy paleon. Inst. 10: 1-140.
Kemp, T. S. 1969. On the functional morphology of the gorgonopsid skull.— Phil. Trans. R.
Soc. (B) 256: 1-83.
Otson, E. C. 1962. Late Permian terrestrial vertebrates, U.S.A. and U.S.S.R.—Am. phil.
Soc. Trans. 52: 1-224.
Or.ov, J. A. 1958. Predatory dinocephalians from the fauna of Isheyevo (Titanosuchia).—
Trudy Paleon. Inst. Akad. Nauk. 72: 1-114.
Romer, A. S. 1956. Osteology of the reptiles. Chicago: University of Chicago Press.
Watson, D. M. S. 1942. On Permian and Triassic tetrapods.— Geol. Mag. 79: 81-116.
Watson, D. M. S. 1948. Dicynodon and its allies.— Proc. Zool. Soc. Lond. 118: 823-877.
ABBREVIATIONS
an angular
art articular
att apon attachment of the bodenaponeurosis
bo basioccipital
260
bs
d
ect
f
inf mar
}
1
m
MAME
MAME (lat)
MAME (med)
MAMIPt (post)
m pt cr
sm
sp
sq
str
temp cr
tp
Vv
Nat. Mus.
PIN
ANNALS OF THE SOUTH AFRICAN MUSEUM
basisphenoid
dentary
ectopterygoid
frontal
inferior margin of the origin of the external adductor from the inner surface
of the cheek
jugal
lacrimal
maxilla
external adductor muscle
lateral division of the external adductor muscle
medial division of the external adductor muscle
posterior pterygoid muscle
medial pterygoid crest
nasal
parietal
palatine
paroccipital process
prearticular
postfrontal
premaxilla
postorbital
prefrontal
pterygoid
quadrate
surangular
septomaxilla
splenial
squamosal
striations
temporal crest
transverse process of the pterygoid
vomer
National Museum, Bloemfontein, South Africa.
Palaeontological Institute, Moscow, U.S.S.R.
South African Museum, Cape Town, South Africa.
6. SYSTEMATIC papers must conform with the International code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., Syn. nov., etc.
An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.
Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 185 : 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (figs 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
comma separates author’s name and year
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figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers
Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.
In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:
Holotype
SAM-A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach,
Port Elizabeth (33.51S, 25.39E), collected by A. Smith, 15 January 1973.
Note standard form of writing South African Museum registration numbers, date and geographical positions.
7. SPECIAL HOUSE RULES
Capital initial letters
(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. *... the Figure depicting C. namacolus...’; ‘*. ..in C. namacolus (Fig. 10)...’
(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene
(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian
Punctuation should be loose, omitting all not strictly necessary
Reference to the author should be expressed in the third person
Roman numerals should be converted to arabic, except when forming part of the title of a
book or article, such as
‘Revision of the Crustacea. Part VIII. The Amphipoda.’
Specific name must not stand alone, but be preceded by the generic name or its abbreviation
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Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of Bio-
logical Abstracts.
HERBERT R. BARGHUSEN
NOTES ON THE ADDUCTOR JAW
MUSCULATURE OF VENJUKOVIA,
A PRIMITIVE ANOMODONT THERAPSID
FROM THE PERMIAN OF THE USSS.R.
OLUME 69 PART 11 JUNE 1976 Ss
ISSN 0303-2515
; > -Af- Cepetow WwW MUS. COMP. ZOOL.
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BuL.LouGu, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FiscHer, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques.—J. Conch., Paris 88: 100-140.
FiscHER, P.-H., DuvaL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines.—Archs
Zool. exp. gén. 74: 627-634.
Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. —
Ann. Mag. nat. Hist. (13) 2: 309-320. p
Konn, A. J. 19605. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. —
Bull. Bingham oceanogr. Coll. 17 (4): 1-51. .
THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. — Denkschr. med.-naturw. Ges. Jena 16: 269-270. d
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 69 Band
June 1976 Junie
Part 11 Deel
ISOPODAN AND TANAIDACEAN CRUSTACEA
FROM THE ST PAUL AND AMSTERDAM ISLANDS,
SOUTHERN INDIAN OCEAN
By
BRIAN KENSLEY
Cape Town Kaapstad
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ISOPODAN AND TANAIDACEAN CRUSTACEA FROM THE ST PAUL
AND AMSTERDAM ISLANDS, SOUTHERN INDIAN OCEAN
By
BRIAN KENSLEY
South African Museum, Cape Town
(With 26 figures)
LMS accepted 26 February 1976]
ABSTRACT
A collection of land and marine Isopoda and Tanaidacea from the St Paul and Amsterdam
Islands, southern Indian Ocean, is dealt with. Three species of tanaids and thirty-four species
of isopods are discussed. Of these, seven new species are described, viz. Eisothistos crateris,
Panathura amstelodami, Cymodocella sapmeri, Munnogonium subtilis, Coulmannia unicornis,
Echinomunna uroventralis, and Janisera trepidus. The latter species belongs to the new genus
Tanisera, while a species of Janira, viz. J. angusta Barnard, is transferred to the new genus
Taniroides. The isopod fauna is analysed into zoogeographical components. It is shown that
there is a strong endemic fauna (27%), as well as a South American/Antarctic/Subantarctic
and a widespread component, but by far the largest component (35 %) is that group common
to the islands and South Africa (mostly the west coast of South Africa). It is concluded that
the isopod fauna of the St Paul and Amsterdam Islands falls into the cold-temperate faunal
category with strong affinities to the fauna of South Africa.
CONTENTS
PAGE
introduction. v2.02) 2... as! i 261
Review of published work. . 262
Specieslist. Sos es, O23
Stationilish, = <3). 0G: <2 268
Systematic discussion . . . 27]
Zoogeographical discussion . 318
Acknowledgements. . . . 321
References ©. =. sw ee. «6
INTRODUCTION
During 1971-2, a reséarch programme centred on the St Paul and Amster-
dam Islands was sponsored by Terres Australes et Antarctiques, with the
logistic support of the Société Anonyme de Péche Maritime et de Ravitaille-
ment (S.A.P.M.E.R.). As part of this programme, J. Beurois of the Station
Marine d’Endoume et Centre Oceanographie, Marseille, made extensive
collections of invertebrates from these islands, both intertidally and subtidally.
The marine and the few terrestrial isopods collected were submitted to the
author for identification. The following is an account of the species found,
with a discussion of their zoogeographical implications. Most of the type
specimens are deposited in the Paris Museum of Natural History. A few para-
types are deposited in the South African Museum, and are designated ‘SAM’.
261
Ann. S. Afr. Mus. 69 (11), 1976: 261-323, 26 figs.
262 ANNALS OF THE SOUTH AFRICAN MUSEUM
REVIEW OF PUBLISHED WORK ON ISOPODA FROM THE ST PAUL
AND AMSTERDAM ISLANDS
The St Paul and Amsterdam Islands, situated at 38.43S, 77.32E and
37.55S, 77.40E respectively (see fig. 1), almost midway in the southern
Indian Ocean, and just north of the Subtropical Convergence, have been
visited by biologists at infrequent intervals. The Austrian frigate Novara called
at the islands during its circumnavigation of the earth in 1857-9. A preliminary
report on the isopods by C. Heller was published in 1861, while the full report
appeared in 1865. This dealt with five species of isopods and one tanaid: Jdotea
nitida, Cleantis granulosa, Porcellio paulensis, Sphaeroma perforata, Cirolana
rugicauda and Tanais gracilis.
Brocchi (1877) reported on a collection of isopods made by Velain and
d’Lisle during the French mission sent to observe the passage of Venus. Nine
species were dealt with, viz. Idotea nitida, Porcellio paulensis, Sphaeroma
perforata, Sphaeroma tuberculata, Cymodoce picta, Cirolana rugicauda, Rocinela
major and Cymothoa gadorum. This material, unfortunately, cannot be located
and must be presumed lost.
The German South-Polar Expedition of 1901-3 visited the islands on the
Gauss. Vanhoffen (1914) reported on the marine isopods, listing seven species
and one tanaid: Cirolana rugicauda, Cycloidura perforata, Dynamenella brunnea,
Jaeropsis paulensis, Antias hispidus, Antias marmoratus, Janira sp. and Tanais
gracilis. Budde-Lund (1906) dealt with the land isopods of this expedition and
mentions Deto armata from St Paul. Finally, André (1932) lists three species,
viz. Paridotea ungulata, Cycloidura perforata and Porcellio paulensis.
To date, the total number of isopods from the St Paul and Amsterdam
Islands numbers 16 species, 4 of these being of uncertain identity. The present
collection includes 35 species, bringing the total number of isopods to 44,
this more-than-doubling of the number being a reflection of the very thorough
collecting carried out by J. Beurois.
Amsterdam Is
St Paul Is
»
Kerguelen Is
Fig. 1. Map showing position of the St Paul and Amsterdam Islands.
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 263
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268
ANNALS OF THE SOUTH AFRICAN MUSEUM
STATION LIST
AMSTERDAM ISLAND (AMS)
Station
No.
al
a2
a3
a4
a5
a6
a7
a8
a9
al0
Date
10.2.1971
10.2.1971
10.2.1971
11.2.1971
11.2.1971
12-1971
11.2.1971
27.2.1971
27.2.1971
27 -2ASTA
12.2.1971
12241971
12.2.1971
12.2.1971
12.2.1971
14.2.1971
14.2.1971
14.2.1971
21971
19.2.1971
19.2.1971
19.2.1971
19.2.1971
22.2.1971
23.2.1971
23.2.1971
24.2.1971
aT 2ASTSA
9.3.1971
9.3.1971
9.3.1971
5.12.1971
5.121971
11.12.1971
13.12.1971
13.12.1971
14.12.1971
Depth (metres)
upper infralittoral
upper infralittoral
upper infralittoral
upper infralittoral
upper infralittoral
upper infralittoral
upper infralittoral
upper infralittoral
midlittoral
between midlittoral
and sublittoral
littoral
midlittoral
lower midlittoral
midlittoral
upper infralittoral
midlittoral
midlittoral
upper infralittoral
upper infralittoral
low tide
0,5
40-50
midlittoral
upper midlittoral
midlittoral
upper sublittoral
120
80-100
80-100
80-100
80-100
Locality and ecological data
north coast; obtained from scraping 400 cm?
amongst abundant algae
north coast; obtained from scraping 400 cm?
amongst Ulva and Splachnidium
north coast; obtained from scraping 400 cm?
amongst Splachnidium
north coast; among rock crevices and kelp
holdfasts
north coast; in Laminaria holdfasts
north coast, amongst algae
north coast, under stones, and in clean
sediment
north coast, under stones
north coast among boulders
north coast, in wet sediment
north—north-east coast; amongst algae and
sponges
north—north-east coast; amongst small
molluscs and algae
north—north-east coast
north—north-east coast amongst Porphyra
and Splachnidium
north—north-east coast under stones
east coast; scrapings from algal growth on
wall
east coast; scrapings from wall
east coast; scrapings from cavity
north coast; algae from lobster pot
north coast, from Macrocystis holdfast
washed ashore
north coast, from Macrocystis holdfast
washed ashore
north coast, from Laminaria holdfast
north coast, from Laminaria holdfast
north coast, under stones
north coast, natural tide-pool
north coast, under stones
south-east coast from lobster pot
north coast, under stones
north coast, in tide-pools
north coast
north coast, amongst red algae
on Macrocystis fronds
north-east coast, from coralligenous bottom,
with bryozoa, sponges, and corals
north-east coast, from coralligenous bottom
north-east coast from antipatharian
epifauna
north-east coast, from encrusting fauna of
stones, sponges, bryozoans, etc.
north-east coast, from antipatharian
epifauna
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 269
Station
No. Date Depth (metres) Locality and ecological data
48 16.12.1971 80-100 north-east coast from bryozoans encrusting
antipatharian trunk
60 21,1972. 80-100 south-east coast from gorgonacian
Acanthogorgia
64a 34.1972 80 south-east coast epifauna from cable sub-
merged for one year
64b 3.1.1972 south-east coast, from buoy and 3-metre
floating cable
73 6.1.1972 80-100 north-east coast, from coralligenous bottom
74 8.1.1972 north-east coast, from encrusting fauna of
stone
78 9.1.1972 25 from buoy cable
83 9.1.1972 south-east coast, from encrusting fauna of
stones
94 13.1.1972 60-100 north-east coast, from bryozoans encrusting
antipatharian trunk
96 13.1.1972 80-100 north-east and south coasts, washed from
variety of coralligenous organisms
100 14.1.1972 0-5 north-east coast, on Macrocystis fronds
101 14.1.1972 80-100 east coast, from epifauna of antipatharian
103 15.1.1972 80-100 north-east coast, from epifauna of anti-
patharian
111 16.1.1972 80-100 north-east coast, from encrusting fauna of
stones
119 20.1.1972 80 north-east coast, from epifauna of bryozoan
132 22 ARIS7Z 80-100 east coast, from epifauna of gorgonacian
Acanthogorgia
133 23.1.1972 80-100 north coast, from sponges and corals
142a 25.1.1972 50 south-west coast, from epifauna of bryo-
zoans and sponges
142b 25.1.1972 50 south-west coast, from epifauna of algae and
sponges
143 25.1.1972 25-30 south-west coast, from amongst red algae
and sponges
147 25.1.1972 60 west coast, from amongst algae, corals, and
sponges
148 25.1.1972 80 west coast, from sponge on antipatharian
base
173 12.2.1972 50-60 north-east coast, from epifauna of anti-
patharian trunk
D1 6.12.1971 80-90 south-east of island
D7 2.1.1972 60-80 east coast
D12 23.2.1972 40-50 north coast
D9 23.1.1972 50-60 east coast
B9 17.12.1971 50 north-east coast
In the following stations, the date of collection is also the station number:
Date Depth (metres) Locality and ecological data
2.5.1969 east coast, from amongst alga Pterocladia
27.3.1970a sublittoral north coast, from crevices in rocks
27.3.1970b sublittoral north coast, from crevices in rocks
28.3.1970a sublittoral north coast, from crevices in rocks and
under stones
28.3.1970b sublittoral north coast, from rocky wall and pools
270
Date
12.12.1970
16.1.1971
NG Ke All
Depth (metres)
ANNALS OF THE SOUTH AFRICAN MUSEUM
Locality and ecological
on dead antipatharian trunk
north-east coast, from Macrocystis holdfast
east coast, from encrusting fauna of rock
Isopods from the following stations were all taken from the stomach and digestive tracts of
the fish Acantholatris monodactylus, all taken with a line, from 2 to 3 metres depth, in February—
March 1971:
Pl; P2, P6, P7, Pl3a, P13b; Pid, P16, Pl7, P23, P26; P27, P28; P29) P30; P31, P33; Ps4ee3e
P37, P38, P39, P42, P44, P46, P47, P48, P49, PSO.
Isopods from the following station were taken from the stomach contents of the rock lobster
Jasus paulensis:
Je
ST PAUL ISLAND
Station
No.
Date
19.12.1971
19.12.1971
19.12.1971
20.12.1971
20.12.1971
20.12.1971
20.12.1971
20.12.1971
21.12.1971
21.12.1971
21.12.1971
21.12.1971
21.12.1971
21.12.1971
22.12.1971
22.12.1971
22.12.1971
22.12.1971
22.12.1971
22 AZ 1ST
22.12.1971
23.12.1971
23.12.1971
23.12.1971
23.12.1971
23.12.1971
23.12.1971
26.12.1971
Depth (metres)
2-3
sublittoral
sublittoral
sublittoral
sublittoral
mid- to sublittoral
upper sublittoral
littoral
sublittoral
sublittoral
sublittoral
sublittoral
sublittoral
sublittoral
sublittoral
upper sublittoral
upper sublittoral
littoral
littoral
littoral
Locality and ecological data
north edge of crater, from amongst sponges,
bryozoans, and ascidians growing on sub-
merged gill net
inside crater, amongst algae
inside crater, amongst Ulva
inside crater, amongst algae
inside crater, from under stones
inside crater, amongst stones and algae
inside crater, from black sediment between
stones and algae
inside crater, amongst organic debris and
green algae
exterior of north jetty, amongst red algae
exterior of north jetty, from amongst
Splachnidium rugosum
exterior of north jetty, on rocks
inside crater, amongst encrusting algae,
sponges, and ascidians
from rock scrapings
from rock scrapings
inside crater, amongst encrusting corals,
bryozoa, and worm tubes on boulders
inside crater, amongst encrusting algae on
boulders
inside crater, amongst encrusting ascidians,
sponges and bryozoans
amongst encrusting algae, sponges and
bryozoans
amongst encrusting algae, sponges, and
ascidians
from Laminaria holdfast
from Laminaria holdfast
exterior of crater, east coast
exterior of crater, east coast
amongst algae exterior of crater, east coast
inside crater, from Laminaria holdfast
inside crater, from Laminaria holdfast
inside crater, from Macrocystis fronds
inside crater, amongst sponges, ascidians
and algae
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 27]
Station
No. Date Depth (metres) Locality and ecological data
59 28.1.1972 from stomach contents of fish Thyrsites
atun
67 30.1.1972 80-100 south-east coast, amongst encrusting algae,
bryozoans and ascidians
T7a 22 AGT2 from Macrocystis holdfast
T7b 2.2.1972 from Macrocystis holdfast
82 4.2.1972 on operculum of fish, Latris lineata
85 4.2.1972 60-70 north-east coast, from encrusting sponges,
worm tubes, and corals
90 15.2.1972 sublittoral inside crater, from encrusting sponges, bryo-
zoans, ascidians, etc.
91 15.2.1972 sublittoral inside crater, from encrusting worm tubes,
and green algae
B7 24.12.1971 45 inside crater, from encrusting bryozoans,
worm tubes and algae on stones
B19 27.1.1972 30 exterior of crater, amongst red algae and
bryozoans
D3 30.1.1972 50-80 north-east coast, from coarse sandy bottom
DSa 30.1.1972 3-4 east coast, from Laminaria holdfast
D5b 30.1.1972 344 east coast, from Laminaria holdfast
DSc 30.1.1972 34 east coast, from red algae
D6 16.2.1972 40-50 north-east coast, from red algae, and
bryozoans
D8 16.2.1972 40 north-east coast, from coarse sediment
In the following stations the date of collections is also the station number:
Date
1970
29.12.1970
1.1.1971
SL.1971
29.1.1971
Depth (metres)
25-30
80-100
Locality and ecological data
exterior of crater, from Macrocystis
exterior of crater, amongst sponges,
ascidians and hydroids
on floating buoy
interior of crater, found in fishing boat
exterior of crater, from submerged cable
SYSTEMATIC DISCUSSION
Order TANAIDACEA
Family Tanaidae
Anatanais gracilis (Heller)
Tanais gracilis Heller, 1865: 133, pl. 12 (fig. 3). Vanhdffen, 1914: 468, fig. 6a-g. Barnard,
192S5a: 381; 1940: 489.
Previous records
Liideritzbucht to Durban, St Paul and Amsterdam Islands, Ceylon.
Die ANNALS OF THE SOUTH AFRICAN MUSEUM
Material
AMS a4 1 St Paul 6c 1
b3 1 Ta 2)
11 1 Tb 1
74 1 8a 1 ovig. 2 + 1
142b 1 DS5/c 1 ovig. 9+ 1
18 2
20 3
22b = 1 ovig. 9
Til Bice wed
90 3
93 2
Leptochelia barnardi Brown
Leptochelia barnardi Brown, 1957: 406, figs 4a—c, Sa.
Previous records
False Bay, Table Bay.
Material
AMS a4 6 St Paul 90 1
Leptochelia savignyi (Kroyer)
Leptochelia savignyi: Brown, 1957: 404, fig. 5b (synonymy).
Previous records
Liideritzbucht to Mocgambique, north and south Atlantic, Mediterranean,
Indo-Pacific.
Material
AMS a4 7} St Paul B7 1
Tb 1
D5/c 1
19 1
90 2
Order ISOPODA
Suborder VALVIFERA
Family Idoteidae
Idotea metallica Bosc
Idotea metallica: Barnard, 1914: 203; 1940: 507. Schultz, 1969: 78, fig. 97.
Previous records
Wide-ranging, almost cosmopolitan: Cape, Mocambique, Natal, Tristan
da Cunha, Greenland, Nova Scotia, Straits of Magellan.
Material
AMS 781 ovig. 2
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 273
Genus Paridotea Stebbing
Heller (1861, 1865) and Brocchi (1877) in their lists of isopods from St Paul
and Amsterdam Islands both include /dotea nitida. Hale (1924), André (1932),
and Sheppard (1957) place J. nitida in the synonymy of Paridotea ungulata
(Pallas). Examination of the idoteids from St Paul and Amsterdam Islands in
the present collection, however, revealed two abundant species of Paridotea,
neither being P. ungulata. Examination of Heller’s type material, and com-
parison with P. ungulata from the Cape, make it clear that P. nitida is a valid
species.
Paridotea nitida (Heller)
Fig. 2B, D, F
Idotaea nitida Heller, 1861: 497.
Idotea nitida Heller, 1865: 131, pl. 12 (fig. 1). Brocchi, 1877: 97.
Previous records
St Paul Island.
Material
AMS 2.5.69 164 5 29 St Paul D5/b 1°
a4 28a, 229 Dales, 3 a0 499
a5 2 29 6c 233 1 ovig. 2
a6 1633 2499 17juv. Ta 233
a7 233 14 26 53 21 99
a8 13 19 19 23d 2 ovig. 22
Cl 12 gd 18 99 20 192 1 ovig. 2
C2 13 LS 28 6gg 292
C3 3gg 392 59 1o
16 3 gd
3 LOSS) L3sSS sSyuv-
143 2 juv.
P16 1Q
P13 13
P29 13 19
P30 233 299
P34 433 399
|e ff 1¢
P38 83d 1199 1 ovig. 2
P42 1g
P46 1¢
P49 13 5 292
P50 2 $2
Remarks
P. nitida can be separated from P. ungulata, with which it was for many
years confused, by the structure of the pleotelsonic apex (distal corners rounded,
separated by shallow concave distal margin in P. nitida, distal corners acute
and spinose, with more concave distal margin in P. ungulata), the uropodal
ramus (distinctly broader than long in P. nitida, as long as wide in P. ungulata),
and the shape of the coxal plates of the last four pereional segments. (See
me. 2A, C, E.)
274 ANNALS OF THE SOUTH AFRICAN MUSEUM
Lin,
C D E
Fig. 2. Paridotea ungulata (Pallas).
A —Four posterior coxal plates; C—Uropodal rarhus; E—Pleotelsonic apex.
Paridotea nitida (Heller).
B—Four posterior coxal plates; D—Uropodal ramus; F-—Pleotelsonic apex.
Separation of P. nitida from P. apposita Barnard, 1965, recorded from
Gough Island, however, is more difficult. The pleotelsonic structure is identical
in these two species, the uropodal ramus very similar (possibly not as wide in
P. apposita as in P. nitida), while the seventh pereional coxal plate is perhaps
more acute and produced in P. apposita. These differences, however, are very
subtle, and more material from Gough Island may well prove the species to be
synonymous, the subtle differences being a reflection of the isolation of the
population.
Paridotea reticulata Barnard
Paridotea reticulata Barnard, 1914: 424, pl. 36D; 1940: 507; 1955: 6.
Previous records
Liideritzbucht, Port Nolloth, Lamberts Bay, Table Bay, False Bay.
Material
AMS. 223° std St Paul 32 699 1 ovig. 2
79 8656 1599 3juv.
100 435 592 1 ovig. 2 7 juv.
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 275
Remarks
The notch found on the sternum of the seventh pereional segment in
South African specimens is not present in specimens from the St Paul and
Amsterdam Islands. The spines of the disto-lateral corner of the pleotelson
are sometimes absent, possibly worn away.
Suborder ANTHURIDEA
Family Anthuridae
Eisothistos crateris sp. nov.
Fig. 3A-G
Description
Male: (head and mouthparts damaged). Head with anterior margin
between eyes evenly convex, about half length of pereional segment I, with
poorly defined median ridge. Pereional segments I-III subequal, each with
anterior ‘shoulder’ and medio-dorsal ridge not quite reaching anterior margin.
Pereional segments IV—VII posteriorly expanded, rounded, also with medio-
dorsal ridge. Segment VII somewhat shorter than preceding segments. Dorso-
lateral keels obvious on anterior three segments only. Anterior three pleonal
segments wider than long, laterally rounded, 4th and Sth segments not distinct,
possibly fused, much shorter and narrower than preceding segments.
Eyes well developed, ocelli large, distinct.
Antennular peduncle consisting of three stout segments, distal segment
bearing pad of elongate setae; flagellum 6-segmented.
Antennal peduncle consisting of two short proximal segments plus two
slender elongate segments; flagellum 6-segmented.
Pereiopod I dactylus with well-developed unguis; propodus two and a half
times longer than wide, ventral margin bearing single distal spine plus 10 small
serrate spines; carpus short, triangular.
Following pereiopods essentially similar to pereiopod I, but with carpi
slightly longer, not underriding the propodi.
Pleopod 2 with elongate rami, exopod carrying six slender setae, endopod
with four setae and sabre-shaped stylet on inner margin reaching well beyond
apex of ramus, apically acute.
Uropods and telson indurated. Uropodal exopod tripartite, consisting of
inner rounded basal process armed with three to four small spines, median
spike-like elongate portion and outer spine-like process; endopod just reaching
apex of telson, apically acute, margins dentate.
Telson evenly rounded, distally wider than proximally, latero-distal
margins serrate; strong flattened medio-dorsal keel present, bearing three or
four small spines distally.
VERGE RTE ; ‘
a iabld wee dana
ra
A—Holotype in
5
=
*
F
E>
E
E
FP.
F
3
ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 3. Eisothistos crateris sp. nov.
dorsal view; B—Telson and uropod; C—Antennule; D—Antenna;
E—Pereiopod I; F—Pereiopod VII; G—Pleopod 2.
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 277
Material
Holotype: St Paul 90 1.6. Paris Museum Is. 1001
Remarks
Of the three species of Eisothistos described, E. vermiformis Haswell and
E. atlanticus Vanh6ffen are easily separated from the present species, either
by general body shape, or on the nature of the telson. There is a marked resem-
blance to E. antarcticus Vanhéffen 1914. The telson of the Antarctic species
differs from the present species in having the medio-dorsal ridge armed with
numerous spines (instead of four obscure spines), and in the outer process of
the uropodal exopod, which is blunt and dentate (rather than spiniform).
These differences could perhaps be due to variation within the same species.
Another obvious difference is in the pleonal structure. The present specimen
has the anterior three pleonal segments large and laterally rounded and the
posterior segments obscure, while in E. antarcticus the anterior five pleonal
segments are subequal.
The species is named crateris as it was collected from the crater of St Paul
Island which is open to the sea.
Panathura amstelodami sp. nov.
Figs 4A-H, 5A-F
Description
Male: Anterior margin of head with median point; eye spots lateral.
Pereional segment I twice length of head, segment II slightly shorter than I,
segments III-VI subequal, VIIth somewhat shorter. Pleonal segments distinct.
Antennule with 5-segmented peduncle, two basal segments as broad as long,
3rd and 4th segments subequal, wider than long, 5th segment more elongate,
flagellum 4-segmented. Antenna with 3-segmented peduncle, 3-segmented
flagellum. Mandibular palp 3-segmented, Ist and 3rd segments subequal, each
about half length of middle segment; third segment bearing row of 7 spines,
proximal 4 spines short; incisor portion of mandible consisting of three strong
teeth plus thin plate with dentate margin.
Maxilla slender, bearing 6 spines.
Maxillipedal palp 4 or 5-segmented, 3rd segment largest, endite extending
on inner margin to distal level of 3rd palp segment, distally tapered.
Pereiopod I propodus with strong proximal triangular tooth forming a
‘thumb’, palm with two smaller proximal and one distal tooth; unguis of
dactylus large and well-defined.
Pereiopod II more slender and longer than I, palm with strong proximal
‘thumb’ and small tooth at base, larger distal tooth; carpus with two spines on
ventral margin. Pereiopods III—VII similar, propodus twice length of carpus,
both segments bearing ventral fringed spines; carpus hardly underriding
propodus.
278 ANNALS OF THE SOUTH AFRICAN MUSEUM
3
GC
D>
QO
X23
yy
———_>
=——— >»
——_
——— 5
————
———
Fig. 4. Panathura amstelodami sp. nov.
A-—Holotype in dorsal view; B—Antenna; C—Antennule; D—Uropodal exopod;
E—Uropodal basis and endopod; F—Mandible; G—Maxilla; H—Maxilliped.
———
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 279
Fig. 5. Panathura amstelodami sp. nov.
A—Pereiopod I; B—Pereiopod Il; C—Telson; D—Pereiopod VII; E—Pleopod 1;
F—Pleopod 2.
280 ANNALS OF THE SOUTH AFRICAN MUSEUM
Pleopod | exopod broad, operculiform, fringed with 13-14 plumose setae;
endopod less than half length of exopod, very narrow, triangular, with single
terminal plumose seta.
Pleopod 2 exopod and endopod equal in length, former distally truncate,
latter distally rounded.
Uropodal exopod leaf-shaped, bearing numerous marginal setae, outer
margin with four to five serrations in hyaline border; endopod with distal
segment longer than proximal, with hyaline border non-serrate.
Telson widest in distal half, with broad hyaline border, latter finely
dentate distally, apical region bearing numerous simple setae.
Material
Holotype AMS a4 13 total length 5,0 mm Paris Museum Is. 1002
Paratypes AMS 119 392 3,0mm 3,2mm 4,4mm SAM-A14994
Paratypes AMS 142b 2992 3,2mm 3,8 mm Paris Museum Is. 1003
Remarks
Barnard (19255) defines Panathura as similar to Apanthura, but possessing
a 6-segmented maxilliped and with the palm of pereiopod I straight. The
distinctive maxilliped, together with the distinct pleonal segmeuts, operculiform
Ist pleopods, and the 5th segment of pereiopods IV—VII underriding the 6th
segment, complete the generic definition. The present material agrees with all
these features, with the possible exception of the pereiopodal carpi, which only
just underride the propodi. The present species differs from P. serricauda,
with which it was collected, on several counts. The telson and uropods in
Barnard’s species are indurated and obviously serrated, and the uropodal
exopod is a broad structure. In P. amstelodami the telson and uropods are
not indurated, and possess a thin hyaline border, the exopod is a leaf-shaped
structure, and although slightly serrate on the outer margin does not approach
the almost dentate condition in P. serricauda. Further differences may be seen
in the pereiopodal structure, as well as in the Ist pleopod with its reduced
endopod. P. formosa Menzies & Frankenberg (1966), recorded from deep
water off Georgia, U.S.A. possesses a very distinctive pleotelsonic structure,
while the endite is not as developed as in the present species.
Panathura serricauda (Barnard)
Fig. 6A-D
Apanthura serricauda Barnard, 1920: 339, pl. 15 (figs 11-12).
Panathura serricauda: Barnard, 1940: 490, 497; 1955: 5.
Previous records
Liideritzbucht, Saldanha Bay, Table Bay, False Bay.
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 28]
C
Fig. 6. Panathura serricauda (Barnard).
A—Animal in dorsal view; B—Maxilliped; C—four distal
segments of pereiopod I; D—Telson.
Material
AMS a4 7 ovig. 22 + 20 specimens St Paul 90 2 specimens
a5 1 specimen
a6 1 specimen
Suborder FLABELLIFERA
Family Cirolanidae
Cirolana rugicauda Heller
Cirolana rugicauda Heller, 1861: 497; 1865: 142, pl. 12 (fig. 13). Brocchi, 1877: 99. Hansen,
1890: 358. Vanhoffen, 1914: 503, fig. 40. Barnard, 1940: 397, fig. 8.
Previous records
St Paul Island, Port Nolloth, St Helena Bay.
282 ANNALS OF THE SOUTH AFRICAN MUSEUM
Material
AMS St Paul
al 1 juv. 6c 233 399 7 juv.
a2 1 juv. Ta 1¢ 3 29 5 juv.
a3 1 juv. 7b 933 1099 6 juv.
a4 19 gg 33 99 sev. juv. 8a 10g¢ 2099 1 ovig. 2 sev. juv.
a5 6 QP 8b 255 499 6 juv.
a6 sev. juv. 8c 7 juv.
a7 a? 5 juv. 14 13 1 juv.
a8 43d 499 22a 1 juv.
a9 2 juv. 24b 1¢ 6 2° sev. juv.
al0 13 8 9° 30a 3d¢ 899 sev. juv.
b3 1 juv. 30b 4 99 8 juv.
Cl ibesy 2 99 2 juv. 90 17363 1099
C3 139 3 juv. 93 433 1199 4 juv.
P46 19
6b 13 4 99 2 juv.
8 13 1 ovig. 2 sev. juv.
10 19
24a DEG 1? sev. juv.
2.5.1969 1°
27.3.1970/b 333 699
Family Sphaeromatidae
Cymodocella sapmeri sp. nov.
Figs 7A-G, 8A-E, 9A-E
Description
Body two and one third times longer than wide, widest at VIIth pereional
segment; head and pereional segments I-VI smooth; posterior margin of
VIIth segment slightly nodose; pleon segment | usually concealed by pereion;
last pleonal segment bearing irregular row of ten to twelve small conical
tubercles. Pleotelson very convex, bearing numerous tubercles, some rounded,
most conical. Apex of pleotelson forming dorsally flexed tube, the opening of
which has tiny spike or papilla protruding into it. Coxal plates of pereional
segments bearing pile of short hairs.
Epistome A-shaped, with slender rami.
Antennules shorter than antennae, with 3-segmented peduncle longer than
flagellum. Antennal peduncle 5-segmented, distal segment longest; flagellum
of about twenty-two segments, longer than peduncle.
Mandibular palp 3-segmented, terminal segment with thirteen pectinate
setae, middle segment with eight pectinate setae; incisor process of four teeth,
lacinia mobilis of three teeth; setal row of eight penicilis; molar process broad,
with dentate margin.
Inner ramus of first maxilla bearing four elongate fringe setae, plus shorter
simple seta; outer ramus tipped with eight dentate spines, degree of dentition
varying from fairly smooth to many large denticles.
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 283
1,0
k | i 4 fh i ity,
Fig. 7. Cymodocella sapmeri sp. nov.
A—Holotype in dorsal view; B—Pleotelson in lateral view; C—Left mandible;
D—Second maxilla; E—First maxilla; F—Maxilliped; G—Epistome.
284 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 8. Cymodocella sapmeri sp. nov.
A—Antenna; B—Antennule; C—Pereiopod I; D—Pereiopod VII; E—Penes.
Second maxilla, inner ramus with about ten fringed spines, both lobes of
outer ramus tipped with four or five pectinate curved spines.
Maxillipedal palp 5-segmented, second segment longest; segments two,
three and four somewhat lobed, lobes bearing clumps of simple setae; endite
bearing about ten fringed spines, and single coupling hook.
Pereiopod I with propodus, carpus, merus, and distal part of ischium
bearing pile of fine short setules; propodus bearing five fringed spines; carpus
very short; dactylus tipped with distal curved spine with shorter blunt spine at
its base. Pereiopods increasing slightly in length posteriorly.
Pereiopod VII with carpus bearing six fringed spines; ischium with two
strong spines on dorsal margin.
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 285
SS
a>
ie
Fig. 9. Cymodocella sapmeri sp. nov.
A—Pleopod 1; B—Pleopod 2; C—Pleopod 3; D—Pleopod 4; E—Pleopod 5.
286 ANNALS OF THE SOUTH AFRICAN MUSEUM
Rami of penis moderately elongate, tapering slightly.
Pleopod | exopod broadly oval, longer than triangular endopod.
Pleopod 2 exopod oval, much smaller and shorter than triangular endopod;
latter with stylet on median margin, stylet slightly longer than endopod, distally
dilated, apically narrowly rounded.
Pleopod 3 with oval exopod shorter than triangular endopod.
Pleopods 4 and 5 with both rami membranous, pleated.
Both rami of uropod elongate-oval, of equal length, distally rounded,
margins slightly serrulate.
Pleotelson of female less obviously tuberculate than that of male.
Material
Holotype AMS C3 14. Paris Museum Is. 1004
Allotype AMS C2 1 ovig. 2 Paris Museum Is. 1005
Paratypes St Paul DS/e 1353 922 6 ovig. 292 Paris Museum Is. 1006
Paratypes AMS Cl 5d 1222 14 ovig. 22 SAM-A14995
AMS St Paul
a2 5 292 1 juv. D/Sa 3 99 lovig.2 2 juv.
a3 [9 yy. D5/b 1 ovig. 2
a4 3 99 14 433 1099 5 juv.
a5 ey eile 16 1d Use.
a6 3 99°" ovis. 9 2uv- 19 2 ovig. 2°
bl ir? 20 Ig) 12 Wowieso2
b2/2 6 22 6 juv. 22b eis:
ie 533 799 10 ovig. 99 17 juv. 28 792 lovig.2 11 juv.
C3 43d 12 3ovig.29 7 juv. 90 633 1622 3 ovig. 22
P29 1 ovig. 2
P30 399
P33 1 ovig. 2
P34 1 ovig. 2
P36 2 99
3 13
6b 236 2 juv.
24a ZG 222) Wovigs?
147 2 juv.
28.3.1970/b 13 12
Remarks
Of the eleven species of Cymodocella described, the present species only
resembles C. nipponica Nishimura, from Japan, to a limited degree. That
Japanese species possesses numerous rounded tubercles on the pleotelson,
while the present species possesses more numerous conical tubercles. Apart from
this tenuous similarity, C. sapmeri is quite distinct from all other described
species. This species is named for S.A.P.M.E.R., the lobster-fishing company
operating around the St Paul and Amsterdam Islands (see introduction), and
for the ship used by the Company.
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 287
1940: 419.
Previous records
St Paul Island.
Material
P16 13
P30 3 3d
P38 13g
2.519699 1g
13.1970 1¢
27.3.1970/b 3¢
28.3.1970/b 43
Dynamenella brunnea Vanhoffen
19
2 99
499
Fig. 10A-B
Dynamenella brunnea Vanhoffen, 1914: 516, fig. 49. non Dynamenella huttoni: Barnard,
2 juv.
sev. juv.
sev. juv.
sev. juv.
1 juv.
3 juv.
2 juv.
sev. juv.
2 juv.
sev. juv.
4 juv.
1 juv.
sev. juv.
1 juv.
sev. juv.
sev. juv.
sev. juv.
sev. juv.
1 juv.
1 juv.
2 juv.
sev. juv.
sev. juv.
7 juv.
sev. juv.
10 juv.
St Paul DS5S/a
D5/c
Tb
8a
14
20
26
28
266. 292° i fuy-
11 99 sev. juv.
13
2 juv.
13 sev. juv.
Ee 5 juv.
Me
as¢
288 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 10. Dynamenella brunnea VanhOffen.
A—Pleotelson; B—Epistome; C—Epistome of D. huttoni.
Remarks
Examination of Vanhdéffen’s type material in both the Berlin Museum and
the British Museum, together with the very large number of specimens in the
present collection, shows that this is indeed a valid species. Separation from
D. huttoni which resembles (and with which Barnard synonymized it) is best
done by reference to three features: pleotelsonic structure, epistome and head
structure, and colour pattern. The dorsal surface of the pleotelson in D. brunnea
always has some indication of tuberculation. In juveniles of less than 2 mm
length, the two main submedian tubercles can already be seen, while in larger
specimens these two large tubercles are supplemented by several smaller tuber-
cles. D. huttoni by contrast invariably possesses a smooth pleotelson. The
median ventrally-directed lobe of the sinuous frontal margin of the head in
D. brunnea is relatively narrower than in D. huttoni; a subtle difference in the
shape of the epistomes is also constant. The colour pattern, though variable in
both species, is more constant in D. brunnea, where a rhomboidal pale patch is
frequently seen mid-dorsally on pereional segments II to IV. While D. huttoni
often has pale dorsal patches, these almost never are as regular as in the former
species.
Dynamenella dioxus Barnard
Dynamenella dioxus Barnard, 1914: 419; 1940: 418, 505. Day, Field & Penrith, 1970: 48.
Previous records
Liideritzbucht, Port Nolloth, Lamberts Bay, Table Bay, False Bay.
Material
AMS a4 Digg | wks St Paul D/Sa 1¢
74 3gg 42992
142a 1¢ 19
142b 7335 1899 5 juv.
147 1¢ 12
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 289
Parisocladus perforatus (H. M. Edwards)
Sphaeroma perforata Edwards, 1840: 211. Heller, 1861: 496; 1865: 139, pl. 12 (fig. 9).
Spheroma perforata: Brocchi, 1877: 97.
Dynamenella perforata: Hansen, 1905: 117, 126.
Cycloidura perforata: Stebbing, 1910: 431. Vanhoffen, 1914: 511, figs 45-46.
Parisocladus perforatus: Barnard, 1914: 402, pl. 32H; 1940: 418, 505. Penrith & Kensley,
1970a: 228; 1970b: 259. Day, Field & Penrith, 1970: 48.
Previous records
Rocky Point (S.W.A.), Mowe Bay (S.W.A.), Swakopmund, Liideritzbucht,
Port Nolloth, Lamberts Bay, Dyers Island, Table Bay, False Bay, Port Alfred,
East London, St Paul Island, Amsterdam Island.
Material
AMS al
a3 63d
a4 433
a5
a7
a9 1¢
b2/la
b2/1b
b2/2 36
b3 es
33
Oy Oy OY Cy
Os OJ OY
2.5.1969
21.3;1910/b) 2
28.3.1970/b
499
3 99
5 29
2 99
37 99
6 2
21 99
5 99
sev. juv.
sev. juv.
sev. juv.
sev. juv.
sev. juv.
2 juv.
2 juv.
1 juv.
sev. juv.
6 juv.
6 juv.
1 juv.
6 juv.
6 juv.
sev. juv.
sev. juv.
sev. juv.
1 juv.
7 juv.
1 juv.
3 juv.
3 juv.
sev. juv.
sev. juv.
2 juv.
sev. juv.
sev. juv.
3 juv.
4 juv.
Si Panl) DS/a" “1d
D5/b
Ds5/e 55d
6a
6c 12 33
Ta 13 33
8a 233
8b 13
14 6 33
15 2
16 7
18 1
19 3
20
22a
36 29
8 99
3 99
3 99
1 juv.
1 juv.
1 juv.
sev. juv.
sev. juv.
4 juv.
sev. juv.
sev. juv.
sev. juv.
3 juv.
sev. juv.
sev. juv.
2 juv.
2 juv.
1 juv.
sev. juv.
4 juv.
sev. juv.
9 juv.
sev. juv.
290 ANNALS OF THE SOUTH AFRICAN MUSEUM
Family Limnoriidae
Limnoria (Limnoria) quadripunctata Holthuis
Limnoria quadripunctata Holthuis, 1949: 167. Menzies & Mohr, 1952: 81. Menzies, 1957:
127. Schultz, 1969: 143.
Previous records
North Sea coast of Holland, California coast, Valparaiso (Chile).
Material
AMS St Paul
a4 1d B19 13
Sa 12 1 ovig. 2 13 juv. Tla 635 922 9 ovig. 29
Sb 12 4 juv. 1970 366 4 ovig. 22
16.1.1971 236 2 ovig. 22 6 juv.
Remarks
These specimens were all found in the holdfasts of the giant brown algae
Macrocystis pyrifera and Laminaria pallida from the upper infralittoral zone,
with the exception of station B19 on St Paul Island, where the isopods were
found in clusters of red algae from a depth of 30 metres.
Family Cymothoidae
Lironeca raynaudii (H. M. Edwards)
Livoneca raynaudii: Barnard, 1920: 358; 1940: 501; 1955: 6.
Previous records
Table Bay, Durban, New Zealand, Tasmania, New South Wales, Japan.
Material
St Paul 82 1g
3.1.1971 co
Family Aegidae
Aega ‘antillensis’ Schiddte & Meinert
Aega antillensis: Richardson, 1905: 170. Barnard, 1925a: 389. Schultz, 1969: 190.
Previous records
Natal, West Indies, Japan.
Material
St Paul 971 1 from fishing-boat
at 1 on Thyrsites atun
Remarks
Slight differences in the frontal laminae, telson, etc., suggest that A. antil-
lensis s.s., and specimens from South Africa, and others from St Paul Island
are not all the same species, although all keyed out to this species.
Aega monilis Barnard
Aega monilis Barnard, 1914: 365, pl. 31C; 1940: 500.
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 291
Previous records
Table Bay, off Cape Peninsula, off East London.
Material
St Paul 85 1 ovig. 2
Remarks
It seems probable from the brief description provided by Brocchi (1877:
100) of Rocinela major, from St Paul Island, that this was a specimen of Aega
monilis. Examination of Brocchi’s type, however, is necessary to establish the
identity of R. major
Suborder ASELLOTA
Family Stenetriidae
Stenetrium crassimanus Barnard
Stenetrium crassimanus Barnard, 1914: 217; 1940: 510. Wolff, 1962: 23.
Previous records
False Bay (Cape), Natal.
Material
St Paul 18 164
91 23¢
Stenetrium saldanha Barnard
Fig. 11A-F
Stenetrium saldanha Barnard, 1920: 403. Wolff, 1962: 24, 29.
Previous records
Saldanha Bay, False Bay, Still Bay.
Material
AMS D1 1¢ 19 St Paul 18 19 1 ovig. 2
142b 1¢ 12 91 1¢ re
Remarks
Wolff’s key (1962: 22), taken from Barnard’s description of S. saldanha,
places this species in the group which lacks any process at the antero-lateral
corner of the first antennal peduncle segment. This segment, however is produced
into a triangular process which is sometimes difficult to see.
Family Antiasidae
Antias dimorphus Menzies
Antias dimorphus Menzies, 1962: 63, fig. 16.
292 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 11. Stenetrium saldanha Barnard .
A—Pereiopod I; B— head in dorsal view; C—Pleotelson; D—Operculum 9;
E—Pleopod 1 ¢; F—Pleopod 2 3.
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 293
Previous records
Southern Chile, Kerguelen Island.
Material
AMS al 1566 16292 14 ovig. 29 St Paul D3 1¢
a2 5 3d 2 ovig. 22 8a 42g5 92922 24 ovig. 2°
a3 53d 422 3 ovig. 29 8b 33d 1 ovig. 2
C2 93d 299 5 ovig. 29 8c 16dd 1692 5 ovig. 99
Ce 2a. ve 3 ovig. 22 14 lo 2 ovig. 99
6b 1g 90 2 ovig. 22
93 2633 999 10 ovig. 99
Remarks
Amongst Vanhéffen’s material of Antias marmoratus collected by the
Siidpolar Expedition at Kerguelen Island (Berlin Museum 17699) are four
specimens of Antias dimorphus showing the enlarged first pereional segment.
The remaining specimens of A. ‘marmoratus’ from Kerguelen at St Paul Island
collected by the Siidpolar Expedition are probably A. hofsteni Nordenstam.
As Vanhdffen’s material is a mixture of two species, each from a different
location, the name A. marmoratus will be omitted from the faunal list of the
two islands, but remains on the list of species for Kerguelen Island.
Antias hispidus Vanhéffen
Fig. 12A-B
Antias hispidus Vanhoffen, 1914: 533, fig. 60. Stephensen, 1927: 356, fig. 24. Nordenstam,
1933: 201, fig. 47. Menzies & Miller, 1955: 385.
Previous records
St Paul Island, Auckland Island, Falkland Island, Graham region
(Antarctica).
-
Fig. 12. Antias hispidus Vanh6ffen.
A—Pleopod 2 $; B—Pleopod 1 ¢.
294 ANNALS OF THE SOUTH AFRICAN MUSEUM
Material
AMS a6 1d ile St Paul 8a 733 792 6 ovig. 29
a4 633 12 3 ovig. 2° 8b 1 ovig. 2
b3 DiSGe ey 14 1d
D12 2 16 2922 $1 ovig. 9
1735 1S da. lis
93 3g3 2929 4 ovig. 99
Antias hofsteni Nordenstam
Fig. 13
Antias hofsteni Nordenstam, 1933: 205. Menzies & Miller, 1955: 385. Menzies, 1962: 60.
Previous records
South Georgia.
Material
AMS © 17.1.1971 1g 1 ovig. 2 St Paul 29.1.1971 192
D19 13 B7 1 ovig. 2
119 1¢
142b
Fig. 13. Antias hofsteni Nordenstam.
Uropod.
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 295
Remarks
In the general body structure and proportions, and in the appendages,
the present material agrees completely with Nordenstam’s description. The
uropod, which was lacking in all the Antarctic material previously collected,
is figured.
Family Janiridae
Caecianiropsis ectiformis (Vanhéffen)
Fig. 14A—C
Austroniscus ectiformis Vanhoffen, 1914: 553, fig. 80.
Caecianiropsis ectiformis: Menzies & Pettit, 1956: 446.
Previous records
Observatory Bay, Kerguelen Island.
Material
St Paul 90 12 _ Total length 1,5 mm
Remarks
Vanh6ffen’s figure does not show the first pleonal segment. This segment,
although difficult to see, is present in both the Kerguelen and St Paul specimens.
Taniroides gen. nov.
Diagnosis
Janirid possessing eyes, slight rostral point, scale on antennal peduncle.
Pereional segments more or less equal in length and width. Coxal plates dorsally
visible on all segments. Pleon longer than wide.
Pereiopod I similar in male and female; propodus distally expanded, palm
straight; dactylus biunguiculate.
Pereiopods II-VI triunguiculate.
Uropod with well-developed basis, exopod half length and width of
endopod.
Pleopod | in male narrow, Y-shaped, proximal halves of rami contiguous,
distal halves divergent, narrow.
Discussion
In the structure of the antennae and mouthparts, the rostral projection,
the prehensile first pereiopod, the triunguiculate dactyli of the remaining
pereiopods, this species could be placed into the genus Janira, as Barnard
(1920) did. The Y-shaped first pleopod of the male, however, which resembles
no other janirid, demands the creation of a new genus. The type species of the
genus is Janiroides angusta (Barnard, 1920), and was originally described from
a single male from False Bay, Cape. Some of the appendages of this specimen
have been refigured.
296 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 14. Caecianiropsis ectiformis (Vanhoffen).
A-—$ in dorsal view; B—Mandible; C— Maxilliped.
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 297
Taniroides angusta (Barnard)
Figs 1SA-F, 16A—D
Janira angusta Barnard, 1920: 404, pl. 17 (figs 1-3); 1940: 511. Wolff, 1962: 41.
Previous records
False Bay, Cape 1 ¢.
Material
AMS 74 3 99 15mm 1.8mm 2,0mm
Supplementary description
Eyes with nine or ten ocelli. Coxal plates visible on all pereional segments,
those on segments I-IV situated at antero-lateral corners, those on segments
V-VII on postero-lateral corners; coxal plates of segments I and II distally
acute, remaining plates rounded.
Pereiopod I similar in male and female, with propodus distally broad,
palm armed with five or six short blunt sensory setae; dactylus apically
biunguiculate.
Operculum in female broader than long, with distal margin slightly concave.
Pleopod | in male narrow, rami proximally contiguous, distally divergent,
apically tapered, tipped with several setae.
Tanisera gen. nov.
Diagnosis
Janirid possessing eyes, no distinct rostral point; scale on antennal
peduncle; antennule well developed. Pereional segments more or less equal in
length and width. Coxal plates dorsally visible on all segments. Maxillipedal
palp 5-segmented, three proximal segments expanded. Pereiopods similar,
ambulatory, dactyli all biunguiculate. Uropodal basis short, rami separate,
exopod slightly shorter and narrower than endopod. First pleopod of male
very broad, expanded, rami fused proximally. Pleotelson marginally serrate,
bearing dorso-lateral ridge. The type species of the genus is Janisera trepidus
sp. nov.
Remarks
The presence of eyes, well-developed antennules, uropods, and molar
process of the mandible, together with the enlarged segments of the maxilli-
pedal palp, all the pereiopods being ambulatory, and the body parallel-sided,
suggest the group of genera Jaera, Janira, Janilirata, Ianiropsis. The first pereio-
pods of the male are not more elongate than the following pereiopods as in
Taniropsis, or prehensile as in Janira, neither are the uropods produced well
beyond the body margin as in Janilirata, nor does the species possess indented
lateral margins of the posterior pereional segments. This species is charac-
terized by the broad first pleopod of the male, the pleotelson bearing a lateral
ridge, and all the pereiopods armed with two dactylar spines.
298
ANNALS OF THE SOUTH AFRICAN MUSEUM
ones
~ ’ RK poe
SD
Cx
T=
Sp,
NaS gare DUS
soles
Fig. 15. Janiroides angusta (Barnard).
A— in dorsal view; B—Mandible; C—First maxilla; D—Second maxilla; E—Mazxilliped;
F—First pereiopod 9°.
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 299
Fig. 16. Ianiroides angusta (Barnard).
A—Pereiopod VII; B—Pleopod 1 $; C—Pleopod 2 ¢; D—Operculum Q.
300 ANNALS OF THE SOUTH AFRICAN MUSEUM
Tanisera trepidus sp. nov.
Figs 17A—G, 18A-—D
Description
Male: Body elongate, parallel-sided, bearing numerous short setae. Head
anteriorly trilobed, rostral process evenly rounded; eyes tiny, dorsal, situated
in posterior half of head. Coxal plates visible on all pereional segments. Pleo-
telson slightly longer than wide; lateral margins bearing about five serrations,
posterior margin with broadly rounded median lobe; single lateral ridge on
distal half, just median to lateral margin, ending distally in spine. Antennule
about half length of antennae, consisting of large basal segment, second segment
about half length and width of basal segment, flagellum of four segments.
Antenna consisting of 5-segmented peduncle, second segment bearing well-
developed scale; first and second segments equal in length to third segment,
latter two-thirds length of fourth segment; flagellum of twelve segments.
Mandible bearing 3-segmented palp, distal segment curved, armed with seven
serrate spines, middle segment with four slender serrate spines; incisor process
of five teeth, setal row of six serrate setae well separated from distally truncate
molar process.
First maxilla, inner ramus bearing two stout setae plus several very fine
setae, outer ramus with at least twelve serrate spines.
Second maxilla, outer ramus slender, bearing three elongate simple setae;
outer lobe of inner ramus slender, bearing four elongate simple setae, inner
ramus stout, carrying numerous simple setae.
Maxillipedal palp 5-segmented, two distal segments slender, three proximal
segments expanded; endite bearing several fringed setae distally, two coupling
hooks medially. Pereiopods similar, basal segment longest, all dactyli tipped
with two curved spines. Pleopod 1 broad, two basal sections together forming
almost complete sphere, distal area between median line and outer spine
broadly convex, carrying about twenty alternately long and short setae.
Pleopod 2 bearing eight elongate simple setae distally.
Uropods with base almost hidden by distal margin of pleotelson, outer
ramus slightly longer and broader than inner, both carrying numerous setae.
Female: Similar in all head and pereional appendages to male.
Operculum carrying numerous close-set setae on distal margin, latter
somewhat concave medially.
Material
Holotype AMS 16.1.1971 1d 2,;0mm _ Paris Museum Is. 1007
Paratypes AMS 133 2 gS 1,1 mm 1,7 mm 2 99 1,3 mm 1,9 mm Paris
Museum Is. 1008
Paratypes AMS 17.1.1971 13 1,8mm 299 1,3mm 1,8mm SAM-14996
AMS 39 2 22 1,9 mm 2,0mm
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 301
0,5
Fig. 17. Janisera trepidus sp. nov.
A—Holotype in dorsal view; B—Antennule; C—Antenna; D-—First maxilla; E—Second
maxilla; F—Mandible; G—Pereiopod I.
302 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 18. Janisera trepidus sp. nov.
A-—Makxilliped; B—Pleopod 1 ¢; C—Pleopod 2 ¢; D—Operculum 9.
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 303
w\ \
\
N i
\ eS
\
\\ i
\
SS
i
AY
ANY
aN
QS
ASS
Fig. 19. Janira capensis Barnard.
A-—d in dorsal view; B—First maxilla; C—Second maxilla; D—Mandible; E—Maxilliped.
304 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 20. Janira capensis Barnard.
A—Antenna; B—Pleopod 1 ¢; C—Pleopod 2 ¢; D—Pleopod 3 ¢.
Janira capensis Barnard
Figs 19A-E, 20A—D
Janira capensis Barnard, 1914: 220, pl. 20b. non Iathrippa longicauda Chilton, Menzies, 1962:
iz
Previous records
Liideritzbucht, Saldanha Bay, Table Bay, False Bay.
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN
Material
AMS _ a7 St Paul 6c 233 299
24a 1¢ Ta 33g 299
B7 36g 12 Ijuv.
7b 1233 3o0vig.22 4299
18 1333 2ovig.92 11 29
20 433 = Hh.
22a Sods lovig.? 2 oO
22c 73s Tovig.99 10 99 4 juv.
23 533 lovig.? 8922 8 juv.
30a 1¢ 1°
30b 2d¢ 1°
32 Lig
91 7133 2ovig.22 399
Remarks
305
Nordenstam (1933), in his description of J. longicauda, figures the pleo-
telsonic margins entire, the carpus of pereiopod I without a distal curved spine,
and mentions the distinct rostrum for the species. Menzies (1962, fig. 51f-g)
also shows a well-developed rostrum, which is lacking in the present material.
Thus J. longicauda is easily distinguishable from Barnard’s valid species.
Taniropsis palpalis Barnard
Fig. 21
Ianiropsis palpalis Barnard, 1914: 222, pl. 21A. Wolff, 1962: 251.
Previous records
Liideritzbucht, Table Bay, False Bay, Port Elizabeth, East London.
Material
AMS St Paul
A6 12 B7 DaGe iee 1 ovig.
B9 1° B19 93s 699
bl 655 322 3 ovig. 2° D3 43g 192
b3 20 3S 1192 7 ovig. 2° D5/a 1¢g 399 $1 ovig.
D1 235 4992 D5/c 3gd 292 4 ovig.
D7 1.9 D6 203d 492 5 ovig.
DI 103g 109° 11 ovig. 22 D8 ib res
D12 11gs 8992 7 ovig. 99 Dil ily
Jil 1g 2:99 3 40 $$ 2299 15 ovig.
ell 43d Ta 10g¢ 792 11 ovig.
P4 1g 7b 83d 692 7 ovig.
P19 1¢ 222 1 ovig. 2 14 2 ovig.
P28 12 16 2d60-i 1 ovig.
P34 19 18 144g3 392 3 ovig.
P45 1 ovig. 2 19 SSS Le 1 ovig.
P83 13 20 2033 999 13 ovig.
27.3.1970/b Lig 3 99 22a 83d 292 6 ovig.
28.3.1970/b 23g 299 22b 63d 2992 3 ovig.
12.12.1970 26d 12 1 ovig. 2 22¢ 12¢g¢ 792 1 ovig.
17.1.1971 392 $1 ovig. 2 23 53d 292 3 ovig.
16.1.1971 43S 12 23a 1? 12 2 ovig.
4 i? 24b 2 ovig.
306 ANNALS OF THE SOUTH AFRICAN MUSEUM
Material
9 13 30a 83S 522 3 ovig. 22
11 1¢3 ie 30b 265d 1299 = 8 ovig. 99
14 22 33 34992 4 ovig. 29 35 2133 1599 11 ovig. 99
28 52 33 5292 25 ovig. 22 67 13
39 173d 6292 4 ovig. 2° Tla 6335 32922 3 ovig. 99
4la 30 dS 22992 11 ovig. 29 90 533 3 ovig. 29
41b 13 392 2 ovig. 22 1970 hee: 1 ovig. 2
48 28 S$ 2999 4 ovig. 29 91 DiSGr Ae 1 ovig. 2
60 835 422 2 ovig. 29 1.1971 235 392 4 ovig. 99
64a 29 3S 999 12 ovig. 99 29.1.1971 36g¢ 1192 11 ovig. 99
64b 23d 322 2 ovig. 9°
73 53d 7922 3 ovig. 92
74 83s 4929
83 23d 1 ovig. 2
94 66 53 3022 31 ovig. 22
96 42 3g 2092 11 ovig. 29°
101 S35 S22 5 ovig. 29
103 183g 999 4 ovig. 99
111 1 ovig. 2
119 1633 692 6 ovig. 29
132 2S) eee
133 4gs 12 1 ovig. 2
142b 21 gd 1092 2 ovig. 22
147 26a 229
148 4335 499
166 58 SS 4092 34 ovig. 22
173 715 $3 34992 44 ovig. 99
Family Jaeropsidae
Jaeropsis beuroisi Kensley
Jaeropsis beuroisi Kensley, 1975: 374, figs 7-8.
Previous records
St Paul and Amsterdam Islands.
Material
AMS St Paul
D12 23g 18 1 ovig. 2 B7 12
39 19 1 ovig. 2 B19 13 rs
4la 13 eS D6 Sigs 292. 1 ovig. 2
41b 1¢ 3 13 PLS;
44 19 7b 13 $$ 1399 10 ovig. 92
64a 13 1 ovig. 2 18 56d 3:99 A ovigsge
74 433 20 ig 19 1 ovig. 2
94 23s 22c 123d 292 9 ovig. 99
119 l1lgd 82922 2 ovig. 22 Tla ASS AS 1 ovig. 2
142b 2353 392 2 ovig. 22 90 18 gg 2099
147 is 3 ee 29.12.1970 1 juv.
173 655 622 4 ovig. 22 19.1.1971 1 he
Jaeropsis paulensis Vanhoffen
Jaeropsis paulensis Vanhéffen, 1914: 531, fig. 59a. Barnard, 1965: 201, fig. 2b. Kensley, 1975:
371, figs 5-6.
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 307
Fig. 21. Janiropsis palpalis Barnard.
3 in dorsal view.
Previous records
St Paul Island, Gough Island.
Material
AMS St Paul
a4 93d 8 ovig. 22 8a 143g5 5922 1 ovig. 2
a8 2 22 8b 3¢¢ 129 2 ovig. 22
a9 1g 8c 933 392 2 ovig. 99
b3 335 622 3 ovig. 22 93 13d 292 4 ovig. 292
14 233 292 1 ovig. 9
27.3.1970/b 1¢
308 ANNALS OF THE SOUTH AFRICAN MUSEUM
Family Munnidae
Munnogonium subtilis sp. nov.
Fig. 22A-I
Description
Female: Body pear-shaped, widest at second and third pereional segments.
Head with anterior margin between antennules straight; eyestalks elongate.
Pereional segments I-IV broad, antero-lateral corners of segment I rounded,
of segment II quadrate, segments III and IV notched; segments V to VII
narrower than preceding segments, with coxal plates visible. Pleotelson as
long as broad, distally broadly rounded, lateral margins as far as insertion of
uropoda dentate (about 12 teeth). Antennule with 2-segmented peduncle, basal
segment shorter, more curved and wider than second segment; flagellum
4-segmented. Antennae missing.
Mandible with narrow toothed incisor process, narrow lacinia mobilis,
followed by four elongate setae; molar process elongate, distally slightly
expanded, truncate; palp missing.
Maxilliped with two distal segments of palp much narrower than three
proximal segments; endite bearing about 6 setae (three simple, three plumose),
plus two coupling hooks. Pereiopod I dactylus bearing elongate terminal
curved spine plus smaller spine; propodus with two sensory spines on ventral
margin, carpus somewhat shorter and broader than propodus, also with two
sensory spines on ventral margin; basis elongate, equal in length to merus and
ischium.
Operculum distally narrowed to rounded apex bearing four stout setae.
Uropoda short, biramous, inner ramus half length and width of outer.
Material
Holotype AMS D9 1 ovig. 2 total length 1,8 mm Paris Museum Is. 1009
Remarks
Bowman & Schultz (1974) recently revised the genus Munnogonium George
& Stromberg. They separated the members of the genus from the closely related
species of Austrosignum Nordenstam by the lack of a mandibular palp in species
of Munnogonium. The resemblance is most marked in general body shape and
proportion between the new species and 4A. Jatifrons Menzies (1962), but that
species has a palp on the mandible. A. globifrons Menzies was placed in Munno-
gonium by Bowman and Schultz, but does not resemble the new species as
closely in general body shape as does A. /atifrons. The new species also resembles
Paramunna kerguelensis Vanhoffen, but it is not known if a mandibular palp
is present or absent in this species.
Genus Coulmannia Hodgson
Hodgson, 1910: 52. Vanhéffen, 1914: 580. Nordenstam, 1933: 225. Menzies, 1962: 173.
Wolff, 1962: 62.
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 309
|
|
Fig. 22. Munnogonium subtilis sp. nov.
A—Holotype in dorsal view; B—Antennule; C—First maxilla; DD—Second maxilla;
E—Mandible; F—Maxilliped; G—Pereiopod I; H—Uropoda; I1—Operculum.
310 ANNALS OF THE SOUTH AFRICAN MUSEUM
Coulmannia unicornis sp. nov.
Figs 23A-F, 24A-H
Description
Male: Head steeply rounded anteriorly. Pereion widest at IIIrd segment.
Pereional segment I laterally bulbous, rounded, bearing strong medio-dorsal
‘horn’, segments II-IV each with single digitiform lateral extension; segments
V-VII posteriorly directed, laterally rounded. Pleotelson anteriorly narrow,
cylindrical, posteriorly bulbous, with five serrations on each side, apically
bluntly rounded.
Eyestalks reaching to proximal half of second antennular segment, with
four ocelli. Antennule with two subequal peduncular segments, flagellum of
four segments. Antenna with 5-segmented peduncle, two distal segments
elongate, subequal; flagellum of six segments.
Mandible lacking palp, with incisor process bearing five teeth; setal row
of three setae; molar process large, cylindrical, distally truncate, with blunt
irregular teeth on grinding surface.
Maxillipedal palp 5-segmented, three basal segments broad, two distal
segments more slender, all segments bearing setae; endite with strong conical
tooth at medio-distal corner, seven or eight setae, single coupling hook present.
Pereiopod I subchelate, carpus with emarginate fringed palm, demarked by
strong conical tooth; dactylus with long unguis; propodus bearing two stout
setae with sensory tips.
Pereiopods II-VII similar, longer and more slender than pereiopod I;
dactyli with long unguis; propodi and carpi elongate, meri short, ischium and
bases subequal elongate. Pleopod 1 proximally fused, distally separate, distal
lobes triangular, with outer basal corners bearing several short setae.
Uropod biramous, inserted without peduncle beneath ridge on bulbous
pleotelson; endopod half length of exopod, both tipped with elongate setae.
Female: Pereional segments II-IV broader than in male; pereional segment
I not armed with ‘horn’ as in male.
Pereiopod I shorter than following legs, but not subchelate.
Operculum longer than broad, distally tapering to bluntly rounded tip,
fringed at widest part with short setae.
Material
Holotype AMS 17.1.1971. 14 2,0mm_ Paris Museum Is. 1010
Allotype AMS 17.1.1971. 1 ovig.2 1,4mm_ Paris Museum Is. 1011
Remarks
The absence of a mandibular palp is the most important character of this
genus, along with the strong apically truncate molar process of the mandible.
Hodgson created the genus for two species, viz. C. australis from Coulman
Island in Victoria Land, South Georgia, and Graham Land, and C. frigida,
described from a single specimen also from the Antarctic. The differences
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 311
Fig. 23. Coulmannia unicornis sp. nov.
A—Holotype in dorsal view; B—Head and anterior segments in lateral view; C— Mandible;
D—Second maxilla; E—Maxilliped; F—Antennule and eyestalk.
312 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 24. Coulmannia unicornis sp. nov.
A-—Antenna; B—Pereiopod I; C—Pereiopod VII; D—Uropoda; E—Operculum 9;
F—Pleopod 1 ¢; G—Pleopod 2 §; H—Pleopod 3 ¢.
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 313
between these two species and C. unicornis are readily apparent. The most
obvious is in the structure of the lateral pereional extensions— bifid in C. australis,
single in C. unicornis and C. frigida. C. australis carries a single median dorsal
spine on each of the pereional segments while in the present species, only
the first pereional segment bears a strong dorsal ‘horn’ (hence the specific
name). Regarding the appendages, there is general agreement between those of
C. australis, well illustrated by Nordenstam (1933) and the present species.
Subtle differences are apparent, particularly in the maxilliped, and first pereiopod
of the male.
Echinomunna uroventralis sp. nov.
Figs 25A-E, 26A—D
Description
Female: Body longer than wide, spinose. Head bearing three spines
anteriorly, median spine at higher level than lateral spines. Eyes lateral. Pereional
segments each bearing strong lateral spine, with two coxal spines visible in
dorsal view. Pereional segment I with two submedian dorsal spines, segments
II-IV with seven dorsal spines, segments V—VII with three dorsal spines. Pleon
fused, longer than wide, with two strong backwardly-directed lateral spines at
widest point, medio-distally with two long diverging spines.
Antennule with 2-segmented peduncle, segments subequal in length, basal
segment wider than distal segment; flagellum 4-segmented, two proximal and
distal segment subequal, short, third segment nine times longer than wide, very
slender. Antennae in all specimens with flagella missing; peduncle of four short
segments, second and third segments each with two strong spines.
Right mandible, incisor process of five strong teeth, followed by five strong
fringed setae, molar process strong, distally truncate; palp 3-segmented, basal
segment two-thirds length of middle segment, distal segment curved, half
length of middle segment, bearing two or three fringed setae, plus numerous
fine setules. Left mandible with incisor process of five strong teeth, narrow
lacinia mobilis carrying five teeth, four stout fringed setae in setal row.
Maxillipedal endite broad, with nine or ten short setae on medio-distal
margin, three coupling hooks on median margin, palp 6-segmented, with
first to third segments wider than fourth to sixth segments, but not as marked
as in Echinomunna s.s., first segment one-third length of second segment, third
segment somewhat shorter than second, segments four and five subequal in
length, terminal segment tiny.
Pereiopod I considerably shorter than following pereiopods; dactylus
curved, with well-marked unguis; propodus with convex palm bearing delicate
fringed membrane plus several stout setae with sensory tips; carpus distally
broader than proximally, shorter than propodus, subequal to merus in length;
ischium and basis elongate, subequal.
314 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 25. Echinomunna uroventralis sp. nov.
A—Holotype in dorsal view; B—Maxilliped; C—Antennule; D—Uropod;
E—Operculum 92
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 315
Pereiopods II-VII slender, elongate, dactyli bearing two terminal claws;
propodi and carpi slender, elongate; meri, ischia, and bases much shorter,
coxae carrying two or three spinose processes, visible dorsally beneath lateral
pereional spine.
Operculum longer than wide, distally tapering to broadly-rounded apex.
Uropods ventral to posterior pleonal spines, uniramous, tipped with
several setae, three-and-a-half times longer than wide.
Material
Holotype AMS 28 lovig.2 Total length 1,8mm Paris Museum Is. 1012
Paratypes AMS 28 2 ovig. 22 Total length 1,7 mm Paris Museum Is. 1013
Remarks
The slender ambulatory pereiopods, lateral eyes, the pleon longer than
broad, segments one to three of the maxillipedal palp broader than segments
four to six, the uropod lacking a peduncle, the strong mandibular molar process,
apically truncate, all place this species in the family Munnidae. Nevertheless,
some differences are apparent when considering the various diagnoses for the
family (e.g. Menzies 1962: 172; Wolff 1962: 59-60). The first three segments
of the maxillipedal palp are not as wide as the endite, and a second pleonal
segment is not visible.
A generic position for this species cannot be arrived at with any confidence.
From Menzies’s key (1962), using the following characters, one arrives at the
choice of either Echinomunna or Acanthomunna: coxal plates visible in dorsal
view, mandibular palp 3-segmented, coxal plates visible on pereional segments
two to seven, body strongly spinose. The uropods of the present species are
neither lateral as in Echinomunna nor dorsal as in Acanthomunna. The relatively
massive, pedunculate, biramous uropods of Acanthomunna would seem to rule
out this genus. Using Wolff’s key (1962), the following characters place the
species in the genus Echinomunna: molar process subcylindrical and strong,
coxal plates two to seven visible in dorsal view, body strongly spinose.
The present material agrees with Vanhéffen’s description of Echinomunna
horrida in the spinose body, the position of the eyes, the construction of the
antennule with one long flagellar segment plus several short segments, in the
structure of the maxilliped and the first pereiopod. The main differences between
E. horrida and E. uroventralis lie in the number of dorsal pereional spines,
and especially in the pleonal structure with the ventrally inserted uropods.
The Antarctic species possesses five proximal spines, while the uropods are
inserted laterally, and the pleon is distally rounded-truncate. In E. uroventralis
there are two strong lateral spines, plus two submedian distal spines with the
uropods inserted beneath them.
The present species is thus placed in the genus Echinomunna with some
reservations, and with the necessity to enlarge the definition of the genus to
include uropods which are inserted either laterally or ventrally.
316 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 26. Echinomunna uroventralis sp. nov.
A—Pereiopod I; B—Pereiopod VII; C—Right mandible; D—Left mandible.
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 317
Munna (Uromunna) nana Nordenstam
Munna nana Nordenstam, 1933: 222, figs. 56-57.
Munna nana forma ‘‘a” Menzies, 1962: 42 fig 5.
Previous records
Chile, Falkland Islands.
Material
AMS a4 2 29 St Paul B19 12 1 ovig. 2
Cl 1 ovig. 2 D36 1d 1 ovig. 2
28 292 4 ovig. 99 6c 236
4la 13 2 ovig. 22 Ta 3d5 222 2 ovig. 99
48 1 ovig. 2 8c 1 ovig. 2
64a 13 19 13 2 ovig. 22
94 19° 22a 19
96 2 ovig 29 22c 233 3992 3 ovig. 99
103 tI 23a 1 ovig. 2
142b 3 3d 1 ovig. 2 30a 1 ovig. 2
173 19 30b Ae. 1 ovig. 2
77a 1d 19
77b 1 ovig. 2
90 53d 1092 2 ovig. 22
Remarks
Slight differences between the present material and Menzies’s forma ‘a’
as well as the forma typica are apparent. The last segment of the antennal
peduncle is not twice the length of the penultimate segment, while the superior
dactylar spine of the pereiopods is smooth.
Suborder ONISCOIDEA
Family Oniscidae
Subfamily Scyphacinae
Deto echinata Guérin
Deto echinata: Budde-Lund, 1885: 234; 1906: 85, pl. 4 (figs 37-38). Panning, 1924: 185,
figs 4-8. Barnard, 1932: 221, fig. 12. Vandel, 1945: 261. Green, 1974: 240.
Deto armata Budde-Lund, 1906: 85, pl. 4 (figs 26-36). Panning, 1924: 191, fig. 10.
Previous records
Rocky Point (S.W.A.), Liideritzbucht, Lamberts Bay, Olifants River
Mouth, Dyers Island, Dassen Island, Table Bay, False Bay, Hermanus, Knysna,
St Paul Island.
Material
AMS a6 7133 4992
15 32 dd 26 $2
27.3.1970/a 2 Oo
28.3.1970/a 24535 18 99
318 ANNALS OF THE SOUTH AFRICAN MUSEUM
Remarks
Panning (1924) synonymized D. armata with D. acinosa, which he regarded
as a species separate from D. echinata. His figure 10, however, is of an immature
male from St Paul Island, not yet showing the development of dorsal spines on
pereion and pleon. Barnard (1932) regards D. acinosa (and with it D. armata)
as synonymous with D. echinata, but qualifies this by stating that the small
strongly granulate form of D. echinata might be regarded as the form acinosa.
That this group of isopods is extremely variable is without doubt. It is
interesting, however, to note the following: of about 100 mature males of
D. echinata from South African localities examined, none showed spinose
processes on the pleon, and that both mature males and females from South
Africa frequently reach a total length of more than 20 mm. Of the 64 adult
males from St Paul and Amsterdam Islands examined, none were larger than
17,5 mm; while 42 specimens possessed a pair of spines on the third pleonal
segment, 14 showed a pair of spines on both pleonal segments three and four,
while 8 specimens lacked pleonal spines completely. Further, these pereional
and pleonal spines never showed the markedly incurved condition of the South
African forms. No differences in the structure of the male genital apparatus
could be seen between St Paul-Amsterdam Island specimens and South African
specimens. It would seem that the St Paul-Amsterdam Island population
should be regarded as part of the D. echinata group, but that this island popula-
tion, isolated as it is, is beginning to show morphological signs of diverging
from the mainland African stock.
Family Oniscidae
Porcellio scaber Latreille
Porcellio scaber: Budde-Lund, 1906: 88. Barnard, 1932: 252, fig. 21 (references).
Porcellio paulensis Heller, 1865: 136, pl. 12 (fig. 5). Brocchi, 1877: 97. André, 1932: 177, 180.
Previous records
Cape Province, St Helena Island, Tristan da Cunha, St Paul and Amsterdam
Islands.
Material
AMS 5a 1 ovig. @ 2 juv.
6a 6 22
6b 1 juv.
ZOOGEOGRAPHICAL DISCUSSION
The total number of species of isopods from the St Paul and Amsterdam
Islands, including past collections plus the present collection, is 43. To get the
effective list of species on which zoogeographical conclusions may be based,
the two undetermined species, viz. Munna sp., and the damaged tanaid, and
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 319
the four inadequately described species of Brocchi (1877) the types of which
have been lost, viz. Spheroma (sic) tuberculata, Cymodoce picta, Cymothoa
gadorum, and Rocinela major, must be removed. This leaves an effective total
of thirty-seven species. These thirty-seven species may be divided into various
categories to give some idea of the relationships of the fauna. When the thorough-
ness with which the present collection was made is considered, it is unlikely
that any major components of the isopod fauna have been overlooked, thus
zoogeographical conclusions may be made with a fair degree of confidence.
The following lists reflect the various categories into which the fauna has been
divided, and the percentage of the total number of species they constitute.
Species endemic to St Paul and Amsterdam Islands—10 species—27 %
Munnogonium subtilis
Coulmannia unicornis
Cymodocella sapmeri
Dynamenella brunnea
Echinomunna uroventralis
Eisothistos crateris
Tanisera trepidus
Jaeropsis beuroisi
Panathura amstelodami
Paridotea nitida
Species with Antarctic, Subantarctic, and South American Affinities—7 species—
18,9%
Antias dimorphus
Antias hispidus
Antias hofsteni
Caecianiropsis ectiformis
Cleantis granulosa
Jaeropsis paulensis
Munna nana
Species with widespread distribution—7 species— 18,9 %
Aega antillensis
Anatanais gracilis
Idotea metallica
Leptochelia savignyi
Limnoria quadripunctata
Lironeca raynaudii
Porcellio scaber
320 ANNALS OF THE SOUTH AFRICAN MUSEUM
Species found only in southern Africa and St Paul and Amsterdam Islands—
13 species—35,1% (with southern African distribution)
Aega monilis Table Bay, East London
Cirolana rugicauda Port Nolloth, St Helena Bay
Deto echinata Rocky Point, S.W.A. to Knysna
Dynamenella dioxus Liideritzbucht to False Bay
Taniroides angusta False Bay
Taniropsis palpalis Liideritzbucht to East London
Janira capensis Liideritzbucht to False Bay
Leptochelia barnardi Table Bay, False Bay
Panathura serricauda Liideritzbucht to False Bay
Paridotea reticulata Liideritzbucht to False Bay
Parisocladus perforatus Rocky Point, S.W.A. to East London
Stenetrium crassimanus False Bay to Natal
Stenetrium saldanha Saldanha Bay, False Bay, Still Bay
Endemism
The degree of certainty with which the endemism of an area can be described
is obviously related to the degree to which surrounding areas have been sampled.
As four of the endemics in the present list range in depth from 30 to 120 metres
—a depth range not often well sampled, these cannot be regarded as endemics
with any certainty. Nevertheless, the figure of 27 per cent agrees well with that
for the fish of St Paul and Amsterdam Islands given by Briggs (1974) of 28 per
cent.
Southern African|St Paul and Amsterdam species
Twelve of the thirteen species in this category are typical of the cold west
coast of South Africa, several being known only from Liideritzbucht to False
Bay. Three species extend to East London or Durban on the east coast; of
these, Aega monilis is a fish parasite, while the other two are asellote isopods
with a predominantly subtidal distribution, and may be considered as within
Stephenson’s (1947) southern warm-temperate province stretching (in senso
stricto) from Cape Agulhas to Algoa Bay.
Knox (1960) regards St Paul and Amsterdam as a separate cold-temperate
province of the austral sea, not especially related to southern Africa. Briggs
(1974: 151), however, considering seven of the ten non-endemic species of fish
of these islands which also occur in southern Africa, regards St Paul and
Amsterdam as more probably related to the ‘Cape of Good Hope. . . within
the southern Africa Warm-Temperate Region’. Stephenson, with a detailed
knowledge of the intertidal of southern Africa, regarded the west coast of
South Africa from about Cape Point to Tropical West Africa as a cold-temperate
province. As twelve of the thirteen isopod species common to South Africa and
St Paul-Amsterdam may be regarded as typical cold-temperate west coast
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 321
inhabitants, Briggs’s view of a warm-temperate fauna is misleading, as is
Knox’s view of an unrelated cold-temperate fauna.
A more accurate view, supported by the isopods and the fish, is that the
St Paul and Amsterdam Islands have a cold-temperate fauna, with a marked
affinity to the cold-temperate west coast fauna of South Africa, but that a small
warm-temperate component related to the warm-temperate south coast fauna
of South Africa is also present. An example of this latter component is Stenetrium
crassimanus, known from False Bay to Natal. Comparison of the sea-surface
temperatures of the two areas gives further weight to this view. St Paul and
Amsterdam have an average summer temperature of 17,5°C, and average
winter temperature of 12,5°C (Wyrtki 1971; Briggs 1974), while on the west
coast of South Africa the annual inshore temperature ranges from 10°-16°C
(Division of Sea Fisheries Report 33).
The presence of a large number of South African species amongst the
isopod fauna of St Paul and Amsterdam as well as other organisms common to
both areas, such as the portunid crab Ovalipes trimaculatus (Arnaud, Beurois
& Noel 1972) and various algae including Splachnidium rugosum and the kelp
Macrocystis pyrifera (Briggs 1974), may easily be explained by invoking the
effect of the West Wind Drift, as noted by Briggs (1974: 150).
ACKNOWLEDGEMENTS
My sincere thanks are due to the following scientists and institutions for
the help and hospitality shown me: Dr H.-E. Gruner of the Natural History
Museum of the Humboldt University, East Berlin; Dr R. J. Lincoln of the
British Museum (Natural History); and Dr G. Pretzmann of the Natural
History Museum, Vienna.
I am grateful to Mr D. C. Lee of the South Australian Museum for the
loan of Idoteid material.
My grateful thanks are due to Dr P. Arnaud, and Dr J. Beurois, and their
colleagues of the Station Marine D’Endoume et Centre D’Oceanographie,
Marseille, for making this collection available to me, and for providing informa-
tion in the form of data, as well as reprints of early works.
I am very grateful to Dr T. E. Bowman of the Smithsonian Institution,
Washington, D.C., and Dr G. A. Schultz of the Jersey City State College, for
their critical reading of the manuscript, and for their many useful comments.
I wish to thank the Trustees of the South African Museum, and the Council
for Scientific and Industrial Research, for a travel grant allowing me to visit
several European museums.
REFERENCES
ANpRE, M. 1932. Crustacés recueillis par M. E. Aubert de la Rue aux Iles Kerguelen, Saint-
Paul et de la Nouvelle-Amsterdam.— Bull. Mus. natn. Hist. nat., Paris (2) 4: 174-181.
ARNAUD, P. M., Beurois, J. & Noe, P. 1972. Portunidae et Grapsidae des Tles Saint Paul
et Amsterdam, Océan Indien (Decapoda, Brachyura).— Beaufortia 20: 7-14.
322 ANNALS OF THE SOUTH AFRICAN MUSEUM
BARNARD, K. H. 1914. Contributions to the crustacean fauna of South Africa. I. Additions
to the Marine Isopoda.— Ann. S. Afr. Mus. 10: 197-230.
BARNARD, K. H. 1920. Contributions to the crustacean fauna of South Africa. VI. Further
additions to the list of Marine Isopoda.— Ann. S. Afr. Mus. 17: 319-438.
BARNARD, K. H. 1925a. Contributions to the crustacean fauna of South Africa. IX. Further
additions to the list of Isopoda.— Ann. S. Afr. Mus. 20: 381-412.
BARNARD, K. H. 19254. A revision of the family Anthuridae (Crustacea Isopoda), with remarks
on certain morphological peculiarities.—J. Linn. Soc. Lond. (zool.) 36: 109-160.
BARNARD, K. H. 1932. Contributions to the crustacean fauna of South Africa. XI. Terrestrial
Isopoda.— Ann. S. Afr. Mus. 30: 179-388.
BARNARD, K. H. 1940. Contributions to the crustacean fauna of South Africa. XII. Further
additions to the Tanaidacea, Isopoda, and Amphipoda together with keys for the identi-
fication of the hitherto recorded marine and fresh-water species.—Amnn. S. Afr. Mus.
32: 381-543.
BARNARD, K. H. 1955. Additions to the fauna list of South African Crustacea and Pycno-
gonida.— Ann. S. Afr. Mus. 43: 1-107.
BARNARD, K. H. 1965. Isopoda and Amphipoda collected by the Gough Island Scientific
Survey.— Ann. S. Afr. Mus. 48: 195-210.
Bowman, T. E. & ScHuLtz, G. A. 1974. The isopod crustacean genus Munnogonium George
and Stromberg, 1968 (Munnidae, Asellota).—Proc. biol. Soc. Wash. 87: 265-272.
BricGs, J. C. 1974. Marine Zoogeography. New York: McGraw-Hill Book Company.
Broccul, G. 1877. Liste des crustacés isopodes provenant de l’ile Saint-Paul et déposés dans
les galeries du Muséum d’Histoire naturelle par MM Vélain et de Lisle.— Bull. Soc.
Dhilomath. Paris (6) 11: 97-101.
Brown, A. C. 1957. A revision of the genus Leptochelia (Crustacea: Tanaidacea) in southern
African waters.— Ann. Mag. nat. Hist. (12) 10: 401-408.
BubDDE-LUND, G. 1885. Crustacea. Isopoda terrestria per familias et genera et species descripta.
Hauniae.
BubbE-LunpD, G. 1906. Die Landisopoden.— Dt. Siidpol.-Exped. 9: 69-92.
Day, J. H., Frecp, J. G. & PeNritH, M. J. 1970. The benthic fauna and fishes of False Bay,
South Africa.— Trans. R. Soc. S. Afr. 39: 1-108.
Epwarps, H. M. 1840. Histoire Naturelle des Crustacés. Paris: Libraire Encyclopédique de
Roret.
GrEEN, A. J. A. 1974. Oniscoidea (Terrestrial Isopoda). In: WILLIAMS, W. D., ed. Biogeography
and Ecology in Tasmania: 229-249. The Hague: Junk.
Hate, H. M. 1924. Notes on Australian Crustacea. III.—Trans. R. Soc. S. Austr. 48: 209-225.
HANSEN, H. J. 1890. Cirolanidae et familiae nonnullae propinquae Musei Hauniensis. —
K. danske Vidensk. Selsk. Skr. 5: 239-426.
HANSEN, H. J. 1905. On the propagation, structure, and classification of the family Sphaeromi-
dae.—Q. JI microsc. Sci. 49: 69-135.
HELLER, C. 1861. Vorlaufiger Bericht tiber die wahrend der Weltumseglung der k. k. Fregatte
Novara gesammelten Crustaceen.— Verh. zool.-bot. Ges. Wien. 11: 495-498.
HELLER, C. 1865. Crustaceen. In: Reise der Oesterreichischen Fregatte Novara um die Erde.
Zoologischer Theil. 2 (8): 1-280. Vienna.
Hottuuts, L. B. 1949. The Isopoda and Tanaidacea of the Netherlands, including the descrip-
tion of a new species of Limnoria.— Zool. Meded., Leiden 30: 163-190.
Hopcson, T. V. 1910. Crustacea IX. Isopoda. In: National Antarctic Expedition. Natural
History. 5: 1-77. London: Trustees of the British Museum.
KENSLEY, B. F. 1975. Five species of Jaeropsis from the southern Indian Ocean (Crustacea,
Isopoda, Asellota).—Ann. S. Afr. Mus. 67: 367-380.
Knox, G. A. 1960. Littoral ecology and biogeography of the southern oceans.—Proc. R. Soc.
(B) 152: 577-624.
Menzies, R. J. 1957. The marine borer family Limnoriidae (Crustacea, Isopoda). Part 1.
Northern and Central America: Systematics, distribution and Ecology.—Bull. mar.
Sci. Gulf Caribb. 7: 101-200. ‘
Menzies, R. J. 1962. The Zoogeography, ecology, and systematics of the Chilean marine
isopods.— Acta Univ. lund. (2) 57: 1-162.
Menzies, R. J. & FRANKENBERG, D. 1966. Handbook on the common isopod Crustacea of
Georgia. Athens, Georgia: University of Georgia Press.
ISOPODAN AND TANAIDACEAN CRUSTACEA FROM SOUTHERN INDIAN OCEAN 323
MENzIES, R. J. & MILuer, M. A. 1955. Marine Asellote isopods of the genus Antias with the
description of a new species from New Zealand.—Trans. R. Soc. N.Z. 83: 383-389.
Menzies, R. J. & Mour, J. L. 1952. The occurrence of the wood-boring crustacean Limnoria
and the Nebaliacea in Morro Bay, California.—Wasmann J. Biol. 10:.81-86.
Menzies, R. J. & Pettit, J. 1956. A new genus and species of marine asellote isopod Caeciani-
ropsis psammophila, from California.—Proc. U.S. natn. Mus. 106: 441-446.
NorpenstaM, A. 1933. Marine Isopoda of the families Serolidae, Idotheidae, Pseudidotheidae,
Arcturidae, Parasellidae, and Stenetriidae, mainly from the South Atlantic. — Further
zool. Results Swed. Antarct. Exped. 3: 1-284.
PANNING, A. 1924. Isopoda.— Beitr. Kennt. land-u. Siisswasserfauna Dt.-SudwAfr. 2: 169-201.
PENRITH, M.-L. & KENSLEY, B. F. 1970a. The constitution of the intertidal fauna of rocky
shores of South West Africa. Part 1. Liideritzbucht.— Cimbebasia (A) 1: 191-239.
PENRITH, M.-L. & KENSLEY, B. F. 19706. The constitution of the fauna of rocky intertidal
shores of South West Africa. Part 2. Rocky Point.—Cimbebasia (A) 1: 243-268.
RICHARDSON, H. 1905. A monograph of the Isopods of North America.— Bull. U.S. natn.
Mus. 54: 1—727.
ScHULTz, G. A. 1969. How to know the marine isopod crustaceans. Dubuque, Iowa: W. C.
Brown.
SHEPPARD, E. M. 1957. Isopod Crustacea Part II. The Suborder Valvifera. Families: Ido-
teidae, Pseudidoteidae and Xenarcturidae Fam. N.—‘Discovery’ Rep. 29: 143-189.
STEBBING, T. R. R. 1910. General catalogue of South African Crustacea.— Ann. S. Afr. Mus.
6: 281-593. -
STEPHENSEN, K. 1927. Crustacea from the Campbell and Auckland Islands. — Vidensk. Meddr.
dansk naturh. Foren. 83: 289-390.
STEPHENSON, T. A. 1947. The constitution of the intertidal fauna and flora of South Africa. III.
— Ann. Natal Mus. 11: 207-324.
VANDEL, A. 1945. La répartition géographique des Oniscoidea (Crustacea Isopodes terrestres).
— Bull. biol. Fr. Belg. 79: 221-272.
VANHOFFEN, E. 1914. Die Isopoden der Deutschen siidpolar-Expedition 1901-1903.— Dr.
Siidpol.-Exped. 15: 447-598.
Wo rr, T. 1962. The systematics and biology of the bathyal and abyssal Isopoda Asellota. —
Galathea Rep. 6: 1-320.
WyrTkI, K., ed. 1971. Oceanographic atlas of the International Indian Ocean Expedition.
Washington: National Science Foundation.
i
ecettel et
1 2 UP a ten nae So
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ioe Awa aime nee ay Pm:
_ dn hes iy, > ©
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_ 3-8 5
6. SYSTEMATIC papers must conform with the International code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
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Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
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Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (figs 8a—b).
Nuculo leegillierti Philippi, 1861: 87.
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BRIAN KENSLEY
ISOPODAN AND TANAIDACEAN CRUSTACEA
FROM THE ST PAUL AND AMSTERDAM ISLANDS,
SOUTHERN INDIAN OCEAN
JOLUME 69 PART 12 SEPTEMBER 1976 ISSN 0303-2515
p-A f- ape wy |
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BuULLouGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FIsCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques.—J. Conch., Paris 88: 100-140.
FiscHER, P.-H., DuvAL, M. & RarFy, A. 1933. Etudes sur les échanges respiratoires des littorines.— Archs
Zool. exp. gén. 74: 627-634.
Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon. —
Ann. Mag. nat. Hist. (13) 2: 309-320. :
Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean. —
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
TuHrELe, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. — Denkschr. med.-naturw. Ges. Jena 16: 269-270. / ae
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 69 Band
September 1976 September
Part ...:42.. . Desi
PLIOCENE GIRAFFOIDEA (MAMMALIA, ARTIODACTYLA)
FROM THE CAPE PROVINCE
By
JOHN M. HARRIS
Cape Town Kaapstad
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PLIOCENE GIRAFFOIDEA (MAMMALIA, ARTIODACTYLA) FROM
THE CAPE PROVINCE
By
JOHN M. HARRIS
Department of Palaeontology, National Museum of Kenya, Nairobi
(With 9 figures and 10 tables)
[MS. accepted 8 April 1976]
CONTENTS
PAGE
Introduction : : : : : 329
Systematics : , , : F £4) 3326
Sivatherium hendeyi sp. nov. : Grek
Giraffa cf. G. jumae : : ; — 347
Discussion . : : ‘ : : F 8352
Acknowledgements . : : ‘ rae Rees
References . é : 3 : . 353
Abbreviations . : j ae o, 353
ABSTRACT
One sivathere species and one giraffine species, both represented by ossicones, teeth and
postcranial elements, have been recovered from ‘E’ Quarry at Langebaanweg. The sivathere
remains appear more primitive than those of the common African sivathere, S. maurusium,
and warrant the creation of a new species. The giraffine species is less abundantly represented
and is very similar to Giraffa jumae from east African Pliocene and Pleistocene localities.
Other faunal evidence from ‘E’ Quarry suggests an early Pliocene age for the Varswater
Formation. The Sivatherium species supports such an interpretation, suggesting that the
Langebaanweg giraffoids are the earliest representatives yet reported of their respective genera.
INTRODUCTION
In recent years a number of Pliocene mammalian assemblages have been
recorded from east and north Africa, but from South Africa only the local
faunas from ‘E’ Quarry, Langebaanweg, are of undoubted Pliocene age. The
younger material from Baard’s Quarry at Langebaanweg is not considered in
this paper. Hence all subsequent references to ‘Langebaanweg’ refer solely to
‘E’ Quarry.
Two local faunas are recognized in the Varswater Formation at Langebaan-
weg but the earlier, from the Quartzose Sand Member, is of similar faunal com-
position to the later local fauna from the superjacent Pelletal Phosphorite
Member. Summaries of the local faunas and geology of the Langebaanweg
locality have been published (Hendey 1974, 1976a; Tankard 1975) as have
detailed descriptions of some of the faunal elements (Maglio & Hendey 1970;
Hendey 1972; Hendey & Repenning 1972; Hooijer 1972). The fossiliferous sedi-
ments cannot be dated by radiometric methods and, though the fossiliferous
325
Ann. S. Afr. Mus. 69 (12), 1976: 325-353, 9 figs, 10 tables.
326 ANNALS OF THE SOUTH AFRICAN MUSEUM
horizons are associated with a marine transgression, chronological evidence from
marine invertebrate faunas has so far proved inconclusive (Hendey, pers. comm.).
The Langebaanweg local faunas include a number of exotic taxa unrepre-
sented elsewhere in Africa (Hendey 1972; Hendey & Repenning 1972; Hendey
19766), but other elements of the fauna may, when studied in detail, afford the
best means of gauging the age of the Langebaanweg deposits relative to other
fossil mammal localities. The Langebaanweg rhinoceros (Hooijer 1972) suggests
chronological equivalence with east African sites dated at or more than 4 m.y.
Elephantid remains from ‘E’ Quarry suggest equivalence with Mursi and Kanapoi
(Maglio 1973: 73; Maglio & Hendey 1970). Available evidence from the Lange-
baanweg Suinae suggests an age of between 4 and 5 m.y. (Harris & White, in
preparation). The Langebaanweg giraffoids tend to support these interpretations
and the Sivatherium appears more primitive than other African occurrences of
this genus. It will be interesting to see in due course the relative ages suggested
by other mammalian groups.
All specimens referred to in the text or tables with the prefix L are from ‘E’
Quarry, Langebaanweg. The full prefix for such specimens is SAM-—PQ-L.
All measurements given in Tables 1-10 are in millimetres.
SYSTEMATICS
Superfamily GIRAFFOIDEA Simpson, 1931
Following his study of the Miocene giraffoids from east and north Africa,
Hamilton (1973) extracted the sivatheres from the family Giraffidae and placed
them into a new family Sivatheriidae. His interpretation that the family Giraffidae
included only two subfamilies—the Paleotraginae and Giraffinae—is followed
here. Two giraffoid species are recognized from Langebaanweg—a sivatheriid
represented by nearly 200 specimens and a giraffine represented by over 80
specimens.
Family Sivatheriidae Hamilton, 1973
Six sivatheriid genera are recognized from Eurasia and Africa and may be
readily distinguished by their ossicone morphology. The frontal bones of Proliby-
therium, a genus restricted to the early Miocene of north Africa, support large
aliform ossicones that have anterior and posterior palmations (Hamilton 1973).
Helladotherium, known from the early Pliocene of Eurasia (and questionably
recorded from north Africa), apparently lacks ossicones and may indeed be a
female of Hydaspitherium or Bramatherium (Matthew 1929; Colbert 1935).
Hydaspitherium, from the middle Pliocene of Asia, is a gigantic sivatheriid with
two ossicones fused at their base into one solid mass on the frontoparietal region
(Colbert 1935). Birgerbohlinea, from the early Pliocene of Spain, possesses two
rounded vertical ossicones that are also strongly connected at their base (Crusa-
font 1952). Bramatherium from the middle Pliocene of Asia has four ossicones
PLIOCENE GIRAFFOIDEA (MAMMALIA, ARTIODACTYLA) FROM THE CAPE PROVINCE 327
—a large pair growing upward from the frontoparietal region and a smaller pair
extending laterally from the parietals (Colbert 1935). Male specimens of Sivathe-
rium giganteum from the late Pliocene and Pleistocene of Asia also possess two
pairs of ossicones—a small pair arising from the frontal region and a larger pair
extending nearly vertically from the parietal region. It is clear from both ossicone
and dental morphology that the Pliocene and Pleistocene sivatheriids from Africa
are more closely related to Sivatherium than to other Eurasian genera and their
consequent allocation to this genus seems justified.
Sivatherium Falconer & Cautley, 1835
A diagnosis of this genus is given in Harris (1976). Previously recorded
specimens of African sivatheriids exhibit a number of differences from the Asian
genotype Sivatherium giganteum. Only one relatively complete cranium is known
from Africa. This specimen, from the Pleistocene of northern Kenya (Harris
1976), differs from S. giganteum in having a longer facial region and a narrower
and less deep cranial region. The posterior ossicones of recorded African siva-
theriids are orientated differently from the Asian species and the anterior
ossicones, if interpreted as such, are sited more posteriorly and are less promi-
nently developed (Harris 1974). The teeth of Asian and African Sivatherium
specimens are, however, similar in morphology. Nevertheless the cranial differ-
ences, and particularly those of the ossicones, are sufficient to warrant taxonomic
separation from S. giganteum.
Owing to the fragmentary nature of much of the material, a number of
difficulties are encountered in attempting to identify African sivatheriids to
species level. In their monograph on the fossil giraffoids of Africa, Singer & Boné
(1960) recognized six different taxa:
Libytherium maurusium
Sivatherium olduvaiense
S. olduvaiense haughtoni
S. olduvaiense vanhoepeni
S. olduvaiense subsp. indet.
Sivatherium cingulatum
As subsequently noted (Churcher 1974; Harris 1974), the type mandible of
L. maurusium was found by Arambourg (1960) to have been wrongly restored.
Arambourg’s new restoration clearly showed that L. maurusium and S. olduvai-
ense were conspecific and that the common African sivathere should be called
Sivatherium maurusium.
With regard to the other taxa recognized by Singer & Boné (1960) it is
evident that the type (and only) upper molar of S. cingulatum, the type (and only)
lower molar fragment of S. olduvaiense haughtoni, the four incomplete teeth
from Tierfontein assigned to S. olduvaiense subsp. indet. and an incomplete
lower molar from Florisbad assigned to S. olduvaiense are all larger than the
range of variation exhibited by other specimens assigned to S. maurusium.
328 ANNALS OF THE SOUTH AFRICAN MUSEUM
Tierfontein, Florisbad and Cornelia are all middle to late Pleistocene sites and
the provenance of the other large specimens is poorly substantiated other than
that they come from the ‘Vaal River Gravels’. It is entirely possible that these
large teeth from the Orange Free State and the Cape Province are from late
South African representatives of S. maurusium, especially as the ossicones from
Tierfontein may unequivocally be assigned to this species. If so, then nearly all
the African sivatheriids reported to date can be assigned, or at least referred, to
a single species.
South African material that is more directly comparable to east and north
African specimens of S. maurusium has been collected from Elandsfontein
(Singer & Boné 1960). The Elandsfontein material includes deciduous and
permanent teeth, posterior ossicones and metapodials. A few deciduous and
permanent teeth are known from Makapansgat but, in the absence of ossicones
from that site, cannot be identified to species. A single unerupted Sivatherium
molar is also known from Swartkrans (Churcher 1974).
There is currently no evidence to support more than one species of Sivathe-
rium from Langebaanweg. If an early Pliocene age is corroborated by subsequent
investigation of other fossil mammals from that locality the Langebaanweg
Sivatherium would appear to be the earliest record of the genus. The sivathere
from this locality possesses conical anterior ossicones that are present also in
the Asian species but absent in S. maurusium. The posterior ossicones of the
Langebaanweg form are not, however, palmate and near-vertically orientated as
in S. giganteum, but extend backwards and outwards from the cranium as in
the common morph of S. maurusium (Type A of Harris 1974). Several posterior
ossicones are known from Langebaanweg and all may be distinguished from
those of S. maurusium by their lack of ornamentation in terms of knobs and
flanges.
African Sivatherium ossicones are quite variable in morphology although
all apparently conform to one of three basic shapes (Harris 1974). The ossicones
are present only in male specimens and the degree of ornamentation may well
be a function of the age of the individual (cf. secondary bone apposition in
extant male giraffes). It is possible therefore that the Langebaanweg sivathere
ossicones, like those of the Langebaanweg giraffines, are all from immature
individuals. On balance, however, the erection of a new species for the Lange-
baanweg sivathere appears warranted on morphological grounds.
Sivatherium hendeyi sp. nov.
Figs 1-7
Sivatherium olduvaiense (Hopwood): Singer & Boné, 1960: 544—-S.
Libytherium olduvaiense (Hopwood): Hendey, 1970: 98.
Holotype
SAM-PQ-L12730, virtually complete left ossicone from the Quartzose
Sand Member of the Varswater Formation in ‘E’ Quarry, Langebaanweg.
329
CAPE PROVINCE
M THE
ARTIODACTYLA) FRO
’
MAMMALIA
GIRAFFOIDEA (
PLIOCENE
*MOIA IOLIONUY “ “MIA [IPI “VY
‘(adAyesed ‘SgZ7IET-Od-WYS) 2UuO0d0Isso JOLIO}UR Io] Mapuay UnayIoAIS “| “B14
330 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 2. Sivatherium hendeyi left posterior ossicone (SAM-—PQ-L12730, holotype).
A. Ventral view. B. Dorsal view.
ee
PLIOCENE GIRAFFOIDEA (MAMMALIA, ARTIODACTYLA) FROM THE CAPE
Fig. 3. Sivatherium hendeyi incomplete posterior ossiocones.
A
B.
e
D
. SAM-PQ-L7243, right posterior ossicone, dorsal view.
SAM-PQ-L7244, left posterior ossicone, ventral view.
. SAM-PQ-L7244, left posterior ossicone, dorsal view.
SAM-PQ-L7243, right posterior ossicone, ventral view.
PROVINCE
332 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 4. Giraffoid dentitions from ‘E’ Quarry, Langebaanweg.
A. SAM-PQ-L10343, Giraffa cf. G. jumae upper dentition (LDP?—M?), occlusal view.
B. SAM-PQ-L31137, Sivatherium hendeyi lower dentition (LP,—,), occlusal view. C. SAM-—
PQ-L31137, Sivatherium hendeyi lower dentition (LP,_.,), lateral view. D. SAM—PQ-L31137,
Sivatherium hendeyi lower dentition (LM,, LM;), lateral view.
$b 4d 4 4 4 4 4
SIAlDI~ OM TON) —F— NM ST ML olm| oi a o&
Fig. 5. Giraffoid postcranial elements from Langebaanweg.
A. Distal humeri of Giraffa cf. G. jumae (top) and Sivatherium hendeyi (bottom), anterior view.
B. Left astragali of Giraffa cf. G. jumae (left) and Sivatherium hendeyi (right), anterior view.
334
ANNALS OF THE SOUTH AFRICAN MUSEUM
a a aes i= Lf 4
AOmS =
a
B
Fig. 6. Giraffoid naviculocuboids from Langebaanweg.
A. Left naviculocuboids of Sivatherium hendeyi (left) and Giraffa cf. G jumae (right), proximal
view. B. left naviculocuboids of Sivatherium hendeyi (left) and Giraffa cf. G. jumae (right),
distal view.
PLIOCENE GIRAFFOIDEA (MAMMALIA, ARTIODACTYLA) FROM THE CAPE PROVINCE 335
Fig. 7. Giraffoid metacarpals.
A. Left metacarpal of Sivatherium hendeyi from Langebaanweg, anterior view. B. Proximal
right metacarpal of Giraffa cf. G. jumae from Langebaanweg, anterior view. C. Right meta-
carpal of Sivatherium maurusium from Elandsfontein, anterior view.
336 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 8. Ossicones of Giraffa cf. G. jumae from Langebaanweg.
A. SAM-PQ-L20490, Giraffa cf. G. jumae left ossicone, medial view. B. SAM—PQ-L20940,
Giraffa cf. G. jumae left ossiocone, lateral view. C. SAM—PQ-L20940, Giraffa cf. G. jumae right
ossicone, lateral view. D. SAM—PQ-L20490, Gircffa cf. G. jumae right ossicene, medial view.
Ww
we
~~
PLIOCENE GIRAFFOIDEA (MAMMALIA, ARTIODACTYLA) FROM THE CAPE PROVINCE
Fig. 9. Teeth of Giraffa cf. G. jumae from Langebaanweg.
A. SAM-PQ-L30126, Giraffa cf. G. jumae upper premolars (RP*~*), occlusal view. B. SAM-—
PQ-L31138, Giraffa cf. G. jumae partial right mandible (RP,;—M,), occlusal view. C. SAM-PQ-
L31138, Giraffa cf. G. jumae partial right mandible (M,—,), occlusal view.
338 ANNALS OF THE SOUTH AFRICAN MUSEUM
Paratype
SAM-PQ-L31285, right and left anterior ossicones from the Pelletal Phos-
phorite Member of the Varswater Formation in ‘E’ Quarry, Langebaanweg.
Diagnosis
Species of Sivatherium of similar size and dental morphology to S. giganteum
and S. maurusium. Posterior ossicones short, extending laterally and backwards
from the cranium and unornamented by knobs and flanges or palmate digita-
tions. Anterior ossicones conical as in S. giganteum. Metacarpals longer than
in other species of Sivatherium.
Etymology
The species is named after Q. B. Hendey, who has been responsible for
collecting most of the ‘E’ Quarry Langebaanweg fauna.
Cranial material
Anterior ossicones
A single pair of anterior ossicones (L31285) has been recovered from the
Pelletal Phosphorite Member at Langebaanweg. The left anterior ossicone lacks
its distal tip but the base of the ossicone is complete and, on the medial side,
extends as far as the intrafrontal suture. The right ossicone is broken distally
and the lateral portion of the base is also missing.
The anterior ossicones are conical structures, as in S. giganteum, and are
slightly compressed mediolaterally (the lateral side being the flatter). They are
inclined backwards at an angle of 50° to the cranial vault at the intrafrontal
suture. They also diverge outwards from each other. A large basal sinus is
present and the adjoining frontal bone is cancellous. The anterior and lateral
surfaces of the anterior ossicones are relatively smooth but a number of irregu-
lar longitudinal ridges and furrows are present on the posterior surface though
less prominent than the grooves on the posterior ossicones.
The anterior ossicones from Langebaanweg would appear to be the first
unequivocal specime 1s from an African sivathere. Anterior ossicones have pre-
viously been reported from Tierfontein and Elandsfontein (Singer & Boné 1960,
pls 29, 42) but the Tierfontein specimen is almost certainly part of a posterior
ossicone while the Elandsfontein specimen is unfortunately no longer available
for examination. The similarity of the conical anterior ossicones to those of
Sivatherium giganteum confirm the relationship of the Pliocene African siva-
theres with Sivatherium from Asia. Anterior ossicones are not present in the
relatively complete sivathere skull from Koobi Fora, Kenya (Harris 1976), and
the apparent absence of anterior ossicones from other later Pliocene and Pleisto-
cene accumulations suggests that they are greatly reduced or lost in S. maurusium.
PLIOCENE GIRAFFOIDEA (MAMMALIA, ARTIODACTYLA) FROM THE CAPE PROVINCE 339
Posterior ossicones
No complete posterior ossicone is known from Langebaanweg although
L12730 is virtually complete and apparently lacks only a few centimetres of
bone from its junction with the cranial vault. Other less complete specimens
include L2542 (right ossicone fragment), L3690 (dorsal fragment of proximal
right ossicone), L4083 (ventral fragment proximal right ossicone), L7243 (distal
right ossicone), L7244 (distal left ossicone), L7245 (posterior fragment of proxi-
mal right ossicone), L7246 (fragment of proximal ossicone) and L6247 (fragment
of proximal left ossicone).
The posterior ossicone morphology is closest to Type A of Harris (1974),
extending outwards and backwards from the cranial vault and perhaps slightly
upwards at the distal tip. The ossicones exhibit only a faint degree of torsion
(anticlockwise in the right ossicone). Near the base each ossicone is almost
triangular in transverse section with a flat dorsal surface and a strongly convex
ventral surface. The distal third is more rounded and is almost circular in
transverse section. The posterior and ventral surfaces are smooth for almost their
entire length. The anterior and dorsal surfaces are sculpted by deep and wide
longitudinal grooves that through torsion extend on to the ventral surface in the
distal portion of the ossicone. The intervening ridges are rugose on the anterior
surface of the proximal portion of the ossicones and again at intervals on the
distal portion. These rugosities are interpreted to foreshadow the flange and
discrete knobs that are a distinctive feature of specimens of S. maurusium. The
longitudinal grooves of the dorsal surface of the ossicone are less deep and less
numerous than those of the anterior surface. Each ossicone terminates in a
distinct bulbous knob.
The low degree of torsion, small size, small basal sinus, virtual absence of
grooves on the ventral surface and lack of flanges and knobs on the posterior
ossicones when taken in conjunction with the discrete conical anterior ossicones
serve to distinguish S. hendeyi from S. maurusium. It is likely, none the less, that
the latter evolved from the former.
Dentition
Although a couple of reasonably complete lower dentitions are known, most
of the Langebaanweg sivathere teeth comprise isolated specimens. The more
complete specimens are listed in the tables of dental measurements (Tables 2-4).
The teeth of S. hendeyi are morphologically indistinguishable from those of
S. maurusium and are of similar size to the S. maurusium teeth from the near-by
but later locality of Elandsfontein. The Elandsfontein and Langebaanweg siva-
there teeth are larger than those from Makapansgat but smaller than comparable
specimens from the Vaal River localities.
Postcranial material
A large number of giraffoid postcranial elements have been recovered from
the Varswater Formation. The apparent presence of only one giraffine and one
340 ANNALS OF THE SOUTH AFRICAN MUSEUM
sivatheriid species and the occurrence of associated skeletal elements permit
ready identification of Sivatherium versus Giraffa postcranial elements. Accession
numbers of Sivatherium limb bones are given in the tables of measurements.
Observed differences between the Langebaanweg sivatheriid and giraffine skeletal
elements are listed below.
Scapula
It is difficult if not impossible to separate scapulae of Sivatherium from those
of Giraffa solely on fragments of the glenoid. Specimens larger than those of
extant G. camelopardalis almost certainly belong to Sivatherium but smaller
specimens may represent either Giraffa or immature Sivatherium individuals.
The posterior border of the blade is distinctly more rounded in Sivatherium, but
this feature can be used only for the more complete specimens which, by virtue
of the relative fragility of the blade, are rare in the fossil record.
Humerus
No complete or proximal giraffoid humeri are known from Langebaanweg
but several distal epiphyses, some with distal portions of the shaft, have been
collected. Sivatherium humeri tend to be larger in overall size and with shorter
but relatively more massive diaphyses than those of giraffines. The distal portion
of the shaft widens to meet the lateral epicondyle more rapidly than in Giraffa.
The coronoid fossa is smaller and less deep in Sivatherium. The posterior articular
surface of the lateral condyle is smaller in Sivatherium and the area for attach-
ment of the extensor digitalis communis is larger.
Ulna
The semilunar notch of the Sivatherium ulna is proportionately larger and
wider than in Giraffa. It is divided into two unequal halves by a low ridge but,
unlike in Giraffa, the lateral portion is larger than the medial. The semilunar
notch extends for the entire width of the ulna in Sivatherium but not in Giraffa.
Distal to the semilunar notch the lateral facet for articulation with the radius
is much shorter in Giraffa.
Radius
The radius of Sivatherium is proportionately shorter and less straight than
that of Giraffa although more massive. The notch dividing the medial from the
lateral facet in the proximal epiphysis extends for nearly half the craniocaudal
length of the epiphysis in Sivatherium but is much shorter in Giraffa. The distal
epiphysis is of similar morphology to that of Giraffa but is wider laterally. On
the caudal edge of the distal epiphysis the articular facets extend farther proxi-
mally but the styloid process of the ulna extends less far posteriorly in Sivathe-
rium than in Giraffa.
PLIOCENE GIRAFFOIDEA (MAMMALIA, ARTIODACTYLA) FROM THE CAPE PROVINCE 341
Metacarpals
Sivatherium metacarpals closely resemble those of Giraffa but are shorter,
wider and with larger epiphyses. Most specimens from Langebaanweg comprise
either proximal or distal epiphyses. The distal epiphysis of the sivathere meta-
carpal may be distinguished from that of Giraffa by its larger size and propor-
tionately wider epiphyseal region. Proximal epiphyses may be separated by the
same factors but, in addition, the lateral facet of the proximal epiphysis is
normally less widely separated from the medial facet at its posterior edge in
Giraffa. The shaft of the sivathere metacarpal is proportionately less deep
craniocaudally.
Tibia
Sivatherium tibiae are shorter but proportionately more massive than those
of giraffine species. The proximal epiphysis is of similar width to that of Giraffa
but is relatively longer craniocaudally. The cnemal crest, extending distally from
the proximal epiphysis, is stout and extends nearly half-way down the shaft. In
Giraffa the cnemal crest is less prominent and is confined to the proximal third
of the shaft. Moreover the central and distal portions of the shaft are more
craniocaudally compressed in Giraffa. In the distal epiphysis the articular facets
for the fibula are more horizontally aligned in Sivatherium, those of giraffine
tibiae being inclined more steeply towards the lateral edge of the epiphysis.
Fibula
Sivatherium fibulae are rather smaller in overall size than those of Giraffa
and the tibial facets are less wide, less elongate and have greater relief. The
anterior tibial facet is convex rather than flat as in Giraffa and is inclined laterally
at a less steep angle. The process on the dorsal edge of the sivathere fibula is
both less tall and less massive than that of Giraffa. The posterior tibial facet of
the Giraffa fibula extends farther forwards towards the lateral edge. The calcaneal
facet on the ventral surface of the fibula is both more elongate anteroposteriorly
and more concave in Sivatherium.
Calcaneum
Sivathere and giraffine calcanea are difficult to separate. Even with the aid
of associated skeletal material of known genus, only one minor morphological
feature serves to distinguish the two. Both Sivatherium and Giraffa calcanea have
an elongate anteroventral facet on the inferior surface for articulation with the
naviculocuboid. In Giraffa the posteromedial edge of the naviculocuboid facet
extends dorsally on to the medial side of the anterior process of the calcaneum.
The additional facet is apparently absent from Sivatherium calcanea.
Astragalus
Sivathere and giraffine astragali are very similar in morphology. Sivathere
astragali tend to be larger in fully adult specimens. The only morphological
342 ANNALS OF THE SOUTH AFRICAN MUSEUM
feature serving to differentiate the two genera is that Sivatherium lacks the
lateral notch that in Giraffa is present between the tibial trochlea and naviculo-
cuboid trochlea. The validity of this distinguishing feature is questionable, for
the lateral notch tends to be at least partly infilled in mature male specimens of
G. camelopardalis.
Naviculocuboid
The Sivatherium naviculocuboid tends to be larger than that of Giraffa. On
the dorsal (proximal) surface the articular facet for the calcaneum often extends
farther posteriorly in Giraffa. The two articular facets for the astragalus are more
markedly separated at their posterior edge in Sivatherium and the posterior
edge of the lateral astragalus facet is not raised dorsally as in Giraffa. On the
ventral (distal) surface the metatarsal facet is proportionately larger than that
of Giraffa but giraffine naviculocuboids possess an additional transversely orien-
tated facet for articulation with the posterior edge of the metatarsal and which
is absent in Sivatherium.
Metatarsal
The Sivatherium metatarsal is shorter, wider and more massive than that of
Giraffa. The proximal epiphyses differ slightly to reflect the different morphology
of the distal naviculocuboid and, in addition, are larger in Sivatherium.
Remarks
Certain trends are evident in the evolution of African sivatheriid ossicones.
These include reduction (and perhaps posterior migration) of the anterior ossi-
cones, elongation and increase in torsion of the posterior ossicones, development
of secondary bone apposition on the anterior edges of the posterior ossicones
and increase in size of the basal sinus concomitant with increase in overall size.
The common morph of the posterior ossicones of S. maurusium (Type A
of Harris 1974) could clearly have evolved from the condition seen in S. hendeyi.
So far different morphs of S. maurusium ossicones (Types B and C of Harris
1974) have been recorded only from Bed II at Olduvai, but each morph is repre-
sented by several individuals. It is possible that the different ossicone shapes seen
at Olduvai are indicative only that the ossicones of S. maurusium were variable
and they thus have no taxonomic significance. It is, however, also possible that
the African sivatheriids were as diverse as those from Asia and, as in some bovid
tribes, specific differences are more immediately evident from the cornual protu-
berances than from other parts of the body. This question can be resolved only
after the collection of further and more complete cranial material from different
localities.
Another apparent evolutionary trend is seen in the metacarpals, those of
S. hendeyi being longer than those of S. maurusium and S. giganteum. Metacar-
pals of S. maurusium appear to decrease in length through time. Shortening of
the metatarsals is less marked and no pronounced trends have been observed
in other elements of the skeleton or in the teeth.
PLIOCENE GIRAFFOIDEA (MAMMALIA, ARTIODACTYLA) FROM THE CAPE PROVINCE 343
TABLE |
Measurements of Sivatherium hendeyi ossicones
L12730 (posterior) 31285 (anterior)
Length - . er ow 450+ 195+
Basal diameter ap. - 120 94
Basal diameterdvy .. 92
Basal diameter tr : : 71
TABLE 2
Measurements of Sivatherium hendeyi deciduous teeth
L2047 L2048 L2049 L2358 L13051 L30475 L2540 L10179 L10809 L13315
DP? ap 35,9’
tr 28,7’
DP® ap 36,7 33,9
tr 31,2 31,6
DP*ap 38,5* S755 --
tr 39,3 39,3 40,0
35,8 32,6
tr 19,7 20,4
TABLE 3
Measurements of Sivatherium hendeyi upper teeth
L1865 L1873 L2046 L11352A L12083
aap : . 32,9
TABLE 4
Measurements of Sivatherium hendeyi lower teeth
L1839 L1876 L3468 L5468 L7247 L7248 L7249 L9161 1L9163 L30875 L31137
>, ap 27,5
tr 17,4
?3 ap 37,9 38,9 39,7
tr 28,5 27,1 26,3
P, ap 43,5 40,7 40,6 42,5 44,5
tr SLs) 382 Sas Ors 31,9
M, ap 45,5 44,3 46,1 47,2 44,4
tr 32,4 344 30,6 35,0 33,8
M, ap 50,6 52,2
tr 36,7 36,7
M,;ap_ 74,5’ 71,5 64,2
tr 38,1’ 34,7 33,1
ANNALS OF THE SOUTH AFRICAN MUSEUM
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PLIOCENE GIRAFFOIDEA (MAMMALIA, ARTIODACTYLA) FROM THE CAPE PROVINCE 347
Family Giraffidae Gray, 1821
According to Hamilton (1973) this family contains only paleotragines and
giraffines. Zarafa from the early Miocene of north Africa is the earliest repre-
sentative of the family and was assigned to the Paleotraginae although not on
the direct lineage to later giraffines or paleotragines (Hamilton 1973, fig. 13).
Giraffines appear to have been of Asian origin but the presence of Giraffa in the
early Pliocene of South Africa may require some modification of the evolutionary
sequence of Honnanotherium, Bohlinia, Giraffa, suggested by Bohlin (1935) and
accepted by Hamilton (1973).
Subfamily Giraffinae Zittel, 1893
Three extinct giraffine species have been recognized in the late Pliocene and
early Pleistocene localities of east Africa (Harris 1976). The smallest species,
Giraffa pygmaeus, is known from ossicones and dentitions but postcranial
remains have so far proved rare. The other two species, G. gracilis and G. jumae,
are more completely represented in the fossil record although no complete crania
have yet been found of Giraffa gracilis. The two larger extinct Giraffa species
may be readily identified on the basis of ossicone morphology and orientation.
The teeth and postcranial elements of all giraffine species are, however, very
similar in morphology. Both the teeth and postcranial elements of G. jumae tend
to be larger than those of G. gracilis, but owing to size variation and sexual
dimorphism neither the teeth nor the skeletal elements of these two species can
always be separated with certainty. The same is true of the teeth of G. pygmaeus
and G. gracilis from Pliocene localities. It is unfortunate that so few associated
giraffine cranial and postcranial remains have been retrieved to date but, taking
into account the fragmentary nature of the Plio-Pleistocene giraffine remains
from east Africa, the taxonomic criteria adopted by Harris (1976) appear to be
valid.
Girafja cf. G. jumae Leakey, 1965
Figs 4-9
Diagnosis
A diagnosis revising that of Leakey (1965) is given in Harris (1976).
Cranial remains
Ossicones
Only three giraffine ossicones have been retrieved from Langebaanweg. The
best preserved specimens comprise a left and right ossicone from a single
individual (SAM-—PQ-L20940) collected from the Quartzose Sand Member. The
individual was immature and neither ossicone had been fused to the cranium.
The two ossicones are similar in morphology to, but rather smaller than, those
of immature extant male giraffes. The lateral edge of each ossicone base is flat
348 ANNALS OF THE SOUTH AFRICAN MUSEUM
but the medial edge is expanded towards its fellow. Each ossicone tapers
gradually above its base but a distinct terminal knob is present at the distal
extremity. The posterior surface of the ossicone is slightly concave but the
anterior surface is virtually flat, both as in G. jumae. The surface of each ossicone
is cut by a number of irregular longitudinal grooves that are most pronounced
on the anterior and median surfaces. As might be expected in an immature indivi-
dual, there is little evidence of secondary bone apposition.
The third ossicone (SAM—PQ-L30960) is somewhat larger but is incomplete
distally. The rugose nature of the bone within the basal cavity suggests that this
specimen, too, was derived from an immature individual.
Dentition
A few nearly complete lower tooth rows are known but the majority of
Langebaanweg giraffine teeth were isolated specimens. The teeth are, in general,
a little larger than those of the extant Giraffa camelopardalis and than Eurasian
fossil giraffine species (G. priscilla, G. attica, G. punjabensis, G. sivalensis).
Though larger than the teeth from the type specimen of G. jumae from Rawe
(Leakey 1965) they are of similar size to teeth of Late Pliocene and early Pleisto-
cene specimens of G. jumae from other east African localities (Harris 1976, and
in preparation).
Postcranial remains
The Langebaanweg giraffine postcranial elements have typical giraffine
morphology. They are of similar size and proportions to those of extant male
giraffes, larger than specimens of G. jumae from the Omo Basin (Coppens et al.,
in preparation) but of similar size to the largest G. jumae specimens from the
Ethiopian locality of Hadar (Taieb et al. 1974; Harris, in preparation). Differ-
ences in postcranial anatomy between the Langebaanweg giraffine and Sivathe-
rium hendeyi have been discussed previously in this paper.
Remarks
In the absence of mature ossicones, whose orientation on the cranium can
be determined, it is difficult to make a positive identification of the Langebaan-
weg giraffine. The known specimens are too large for Giraffa pygmaeus and the
presence of a distinct terminal knob suggests that they do not belong to G. gracilis.
They are slightly smaller than, but otherwise quite similar to, immature ossicones
of Giraffa camelopardalis but could equally well belong to G. jumae (immature
specimens of which are yet unknown) or even a new species. None of the three
east African fossil giraffine species is clearly ancestral to Giraffa camelopardalis
and the possibility that the extant giraffe was of South African origin, and
perhaps descended from the Langebaanweg form, cannot be entirely ruled out.
The acquisition of mature giraffine ossicones from Langebaanweg may in future
solve the problem. Meanwhile, because the teeth are rather larger than those of
PLIOCENE GIRAFFOIDEA (MAMMALIA, ARTIODACTYLA) FROM THE CAPE PROVINCE 349
extant giraffes and, on balance, the postcranial elements are rather more robust
than the latter, the Langebaanweg giraffine is provisionally assigned to Giraffa
cf. G. jumae.
TABLE 6
Measurements of Giraffa cf. G. jumae ossicones
L20940 (1) 20940 (r) L30690
Length : : 2 177 177
Basal diameter ap. 86 92 112
Basal diameter tr. 59 59
TABLE 7
Measurements of Giraffa cf. G. jumae deciduous teeth
L6606 L13510 L13043 (1) 113043 (r) L10938 L10939 ~—=L10800
DP? ap 23,3 2a 24,3
tr 19,4 19,7 17,4
DP* ap 26,1 2 | 26,8 9 ft | 28,8
tr 21,8 20,3 22,6 23:7 24,4 26,5
DP* ap 2155 28,2
tr 27,4 28,3
L30057 L2543 L6022 L11617 L2541
DP, ap 19,6
tr 10,9
DP, ap 25,8
tr 14,4
DP, ap 36,2 38,2 a2. 33,9*
tr 19,1 18,6 ule 16,5” 18,6’
TABLE 8
Measurements of Giraffa cf. G. jumae upper teeth
L30126 L22021 L31138 L10343 (1) L10343 (r)
Pap. ~% : 22,3
tr : : 26,5
P* ap 25,0
tr 27,4
Pap 25,6
tr ; ; 28,2
M' ap . : 31,3 29,2
tr , : 93,1 S537)
M?’ap.. : 33,0 33,4 31,6
tr ; : Sis 34,8 36,1
M*ap.. 5 35,2
tr : ‘ 36,8
TABLE 9
Measurements of Giraffa cf. G. jumae lower teeth
L22021 L5767 L31138 L20987 L4773
P, ap . ; 23,0
tr ; . 14,4
Psap. | 3 : 27,0 23,6
tr ; : 20,1 19,7
Pe ap. ; 28,5 24,4 26,1*
tr ‘ F 22,7 22,1 20,6
Miap.. , 27,5 33,1 27,5
tr : : 22,2 235 23,7 24,4
M.ap.. . B13 32,9 30,0
tr ain 25,3 2355 24,6
M,ap.. 2 46,9
tr : é 25,0
ANNALS OF THE SOUTH AFRICAN MUSEUM
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352 ANNALS OF THE SOUTH AFRICAN MUSEUM
DISCUSSION
The giraffoids from Langebaanweg are almost certainly the earliest repre-
sentatives of their respective genera yet reported from Africa although confirma-
tion of their age must await detailed investigation of other elements of the fauna.
If an early Pliocene age is substantiated the two Langebaanweg giraffoids are
of similar if not earlier age than their nearest Asian relatives and the question
of Asian versus African origin of the giraffines and later sivatheriids will need
to be reinvestigated.
Hamilton (1973) assigned Prolibytherium to the Sivatheriidae and suggested
that sivatheriids and giraffids were independently derived from Oligocene
Dremotheriidae. Prolibytherium shares many morphological features with
dremotheriids and primitive cervids and may, perhaps, be better assigned to one
of these families. The evidence for sivatheriid affinity is not conclusive and, as
Hamilton himself points out (1973, fig. 13), if placed in the Sivatheriidae
Prolibytherium can be only distantly related to other known members of the
family.
As is evident from Sivatherium hendeyi, a foreshortened anterior limb is not
a primitive feature of Sivatherium but a character acquired during the evolution
of this genus. Moreover, the posterior ossicones of S. hendeyi are relatively short,
narrow and unornamented, suggesting that the palmate posterior ossicones of
S. giganteum and the more complex examples of S. maurusium are advanced
features. The disposition of the ossicones of S. Aendeyi is not dissimilar to that
of Giraffokeryx (Colbert 1933). It is not implausible that Sivatherium and related
genera were relatively late (upper Miocene?) offshoots from paleotragine stock.
If so, relegation of the family Sivatheriidae to a subfamily of the Giraffidae
might appear warranted.
Abundant giraffine material is known from Hadar in Ethiopia (Harris, in
preparation) and Laetolil in Tanzania (Harris, in preparation). The earlier
horizons from Hadar are dated in excess of 3 m.y. (D. C. Johanson, pers. comm.)
and the Laetolil faunas are believed to be between 3,25 and 3,75 m.y. (M. D.
Leakey, pers. comm.). Occurring at both localities are representatives of the
three species known from east African Pleistocene localities. Giraffines have not
yet been reported from Lothagam and Kanapoi but a few undescribed and
incomplete specimens are known from Lukeino which have been dated at
between 5,4 and 6,7 m.y. (Pickford 1975). The subfamily Paleotraginae is cur-
rently being revised (W. R. Hamilton, pers. comm.) and perhaps in the near
future further east African material will be available to help place the Langebaan-
weg giraffines in their proper context.
ACKNOWLEDGEMENTS
The Langebaanweg giraffoid material was studied at the South African
Museum at the kind invitation of Dr Q. B. Hendey. I am grateful to Dr Hendey
and his staff for making available the material and facilities for its study. I am
PLIOCENE GIRAFFOIDEA (MAMMALIA, ARTIODACTYLA) FROM THE CAPE PROVINCE 353
grateful also to Drs Hendey, A. W. Gentry and W. R. Hamilton for discussion
of aspects of the manuscript. The project was undertaken while the author was
in receipt of a travel grant from the L. S. B. Leakey Foundation.
REFERENCES
ARAMBOURG, C. 1960. Precisions nouvelles sur Libytherium maurusium Pomel, Giraffide du
Villafranchien d’Afrique. Bull. Soc. geol. Fr. (7) 2: 888-894.
BOHLIN, B. 1935. Some remarks on fossil Giraffidae. Bull. geol. Soc. China 14: 83-89.
CHURCHER, C. S. 1974. Sivatherium maurusium (Pomel) from the Swartkrans Australopithecine
Site, Transvaal (Mammalia: Giraffidae). Ann. Transy. Mus. 29: 65-69.
COLBERT, E. 1933. A skull and mandible of Giraffokeryx punjabiensis Pilgrim. Am. Mus. Novit.
632: 1-14.
CoLserT, E. 1935. Siwalik mammals in the American Museum of Natural History. Trans. Am.
Phil. Soc. n.s. 26: 1-401.
CRUSAFONT, M. 1952. Los Jirafidos Fosiles de Espafia. Mem. Commun. Inst. geol. Barcelona 8:
9-239.
HAMILTON, W. R. 1973. The lower Miocene ruminants of Gebel Zelten, Libya. Bull. Brit. Mus.
nat. Hist. (Geol.) 21: 75-150.
Harris, J. M. 1974. Orientation and variability of the ossicones of African Sivatheriinae
(Mammalia: Giraffidae). Ann. S. Afr. Mus. 65: 189-198.
Harris, J. M. 1976. Pleistocene Giraffidae (Mammalia; Artiodactyla) from East Rudolf,
Kenya. Fossil Vert. Afr. 4: 283-332.
HENDEY, Q. B. 1972. A Pliocene ursid from South Africa. Ann. S. Afr. Mus. 59: 115-132.
HENDEY, Q. B. 1974. The Late Cenozoic Carnivora of the South-Western Cape Province. Ann.
S. Afr. Mus. 63: 1-369.
HENDEY, Q. B. 1976a. The Pliocene fossil occurrences in ‘E’ Quarry, Langebaanweg, South
Africa. Ann. S. Afr. Mus. 69: 215-247.
HENDEY, Q. B. 19765. Fossil Peccary from the Pliocene of South Africa. Science 192: 787-789.
HENDEY, Q. B. & REPENNING, C. A. 1972. A Pliocene phocid from South Africa. Ann. S. Afr.
Mus. 59: 71-98.
Hoover, D. A. 1972. A late Pliocene Rhinoceros from Langebaanweg, Cape Province. Ann.
S. Afr. Mus. 59: 151-191.
LEAKEY, L. S. B. 1965. Olduvai Gorge 1951-1961 1. Cambridge: University Press.
Mac1tio, V. J. 1973. Origin and evolution of the Elephantidae. Trans. Am. phil. Soc. n.s. 63:
3-149.
MacLio, V. J. & HENDEY, Q. B. 1970. New evidence relating to the supposed stegolophodont
ancestry of the Elephantidae. S. Afr. archaeol. Bull. 25: 85-87.
MATTHEW, W. D. 1929. Critical observations of Siwalik mammals (exclusive of Proboscidea).
Bull. Am. Mus. nat. Hist. 56: 437-560.
PICKFORD, M. 1975. Late Miocene sediments and fossils from the Northern Kenya Rift Valley.
Nature, Lond. 256: 279-284.
SinGER, R. & Bong, E. L. 1960. Modern giraffes and the fossil giraffids of Africa. Ann. S. Afr.
Mus. 45: 375-548.
TaieB, M., JOHANSON, D. C., CoppeNs, Y., BONNEFILLE, R. & KALB, J. 1974. Decouverte
d’Hominides dans les series Plio-Pleistocenes d’ Hadar (Bassin de l’Awash, Ethiopie). C. r.
hebd. Séanc. Acad. Sci., Paris 279: 735-738.
TANKARD, A. J. 1975. Varswater Formation of the Langebaanweg-Saldanha area, Cape
Province. Trans. geol. Soc. S. Afr. 77: 265-283.
ABBREVIATIONS
ap anteroposterior Lat lateral
dv dorsoventral prox proximal
tr transverse dist distal
estimated measurement med medial
* approximate measurement nayvcbd naviculocuboid
Max maximum tr trochlea (astragalus measurements only)
+ ip bette) igetmtktas 3a
Penbes putin
— as et Fe ,*
i 7 a) . 7
-
a
ee
_
* A
#
—
Yy
“og ne~' as 9 9 he
® ¢ *« ~ 4 =
9
Sn ba ees ere au
= " é
j
% Oy —
— |
— ie =
a 4
’
~~
Sires Pre
o Sy Perey <a
a= ee.
7 _ en a ay ¢
caieee jad Paar
acniaagp ai. Ane.
ott hh dene
re ee ed
> : _ pith
oF) <004 | wae
) OP ecw
teh re
Pe
i ate fd Ve
’ ®
— «
= 3
q
TT ~
? ‘
6. SYSTEMATIC papers must conform with the International code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
Nov., syn. nov., etc.
An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
_ a scientific name must be separated from the scientific name by a colon.
Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (figs 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma separates consecutive numbers
Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.
In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:
Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach,
Port Elizabeth (33.51S, 25.39E), collected by A. Smith, 15 January 1973.
Note standard form of writing South African Museum registration numbers, date and geographical positions.
7. SPECIAL HOUSE RULES
Capital initial letters
(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. ‘... the Figure depicting C. namacolus ...’; *. . .in C. namacolus(Fig. 10)...’
(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene
(c) Scientific names, but not their vernacular derivatives
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Punctuation should be loose, omitting all not strictly necessary
Reference to the author should be expressed in the third person :
Roman numerals should be converted to arabic, except when forming part of the title of a
book or article, such as
‘Revision of the Crustacea. Part VIII. The Amphipoda.’ F mt
Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively. ;
Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of Bio-
logical Abstracts.
JOHN M. HARRIS
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