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ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
VOLUME 67 BAND
THE TRUSTEES OF THE DIE TRUSTEES VAN DIE
SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM
CAPE TOWN KAAPSTAD
1976
@ SET, PRINTED AND BOUND IN THE REPUBLIC OF SOUTH AFRICA BY
THE RUSTICA PRESS (PTY.) LTD., WYNBERG, CAPE
852
ANNALS OF THE ANNALE VAN DIE
SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM
VOLUME 67 BAND 67
NEW GENERIC NAMES PROPOSED IN THIS VOLUME
Page
Agulanthura Kensley, 1975... ee = te ae a = oe de TD
Austroarcturus Kensley, 1975 ne Bs = a A ~ eS Bs 40
Chelydontops Cluver, 1975 _.. us te a ne? AG ~ aa fh 8
Flexanulus Debrenne, 1975 .. — ‘s ae aes a me cm ye 4385
Statanulocyathus Debrenne, 1975... ae he = suk ne eg eae 72
LIST OF CONTENTS
CLUVER, M. A.
A new dicynodont reptile from the Tapinocephalus zone (Karoo System, Beaufort
Series) of South Africa, with evidence of the jaw adductor musculature. (Pub-
lished June 1975.)
DEBRENNE, F.
Archaeocyatha provenant de blocs erratiques des tillites de Dwyka (Afrique du
Sud). (Published October 1975.) ee aa ais he ts
GARDINER, B. G. see JUBB, R. A.
GARDINER, B. G. & JUBB, R. A.
A new palaeoniscid from the Lower Beaufort Series of South Africa. (Published
December 1975.)
Gow, C. & MILLARD, N. A. H.
Two new species of campanularian hydroids from South Africa. (Published March
1975.)
GRIFFITHS, C. L.
The Amphipoda of southern Africa, Part 5. The Gammaridea and Caprellidea of
the Cape Province west of Cape Agulhas. (Published August 1975.) ..
JuBB, R. A. see GARDINER, B. G.
JupB, R. A. & GARDINER, B. G.
A preliminary catalogue of identifiable fossil fish material from southern Africa.
(Published October 1975.)
KENSLEY, B.
Taxonomic status of the pygocephalomorphic Crustacea from the Dwyka ‘White
Band’ (Permo-Carboniferous) of South Africa. (Published June 1975.)
KENSLEY, B.
Marine Isopoda from the continental shelf of South Africa. (Published July 1975.)
KENSLEY, B.
Five species of Jaeropsis from the southern Indian Ocean (Crustacea, Isopoda,
Asellota). (Published October 1975.)
KILBURN, R. N. & TANKARD, A. J.
Pleistocene molluscs from the west and south coasts of the Cape Province, South
Africa. (Published October 1975.)
MANNING, R. B.
A new species of Meiosquilla (Crustacea, Stomatopoda) from South Africa.
(Published October 1975.)
MILLARD, N. A. H. see Gow, C.
Nasu, D. S.
The morphology and relationships of a crocodilian, Orthosuchus stormbergi, from
the Upper Triassic of Lesotho. (Published October 1975)
TANKARD, A. J. see KILBURN, R. N.
Page
331
441
91
25
35
367
183
363
220
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NE 67 PART 1 MARCH 1975 ISSN 0303-2515
4 07. Lg
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.
Koun, A. J. 1960b. Spawning behaviour, egg masses and larval development in Conus from the Indian
Ocean.— Sull. Bingham oceanogr. Coll. 17 (4): 1-51.
THIELE, J. 1910. Moilusca: 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 67 ~~ Band
March 1975 Maart
Part 1 Deel
TWO NEW SPECIES OF
CAMPANULARIAN HYDROIDS
FROM SOUTH AFRICA
By
C. GOW
&
N. A. H. MILLARD
Cape Town Kaapstad
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TWO NEW SPECIES OF CAMPANULARIAN HYDROIDS FROM
SOUTH AFRICA
By
C. Gow
Bernard Price Institute for Palaeontological Research, University of the
Witwatersrand, Johannesburg
&
N. A. H. MILLARD
South African Museum, Cape Town
(With 2 figures)
[MS accepted 22 July 1974]
ABSTRACT
Two new hydroid species of the family Campanulariidae are described from ‘False Bay,
South Africa, namely Campanularia pecten and Campanularia roberti. Both are unusual in
the form of the gonotheca which is shaped like a bivalve shell.
CONTENTS
PAGE
Introduction . f 1
Description of material 1
Discussion . 5
References 6
INTRODUCTION
These two small species were discovered by the first author during the
preparation of a project for an honours degree at the University of Cape Town
in 1968. Both colonies were fertile and were retained alive in the laboratory
until the release of the products of the gonothecae. Microscopic examinations
were made of fresh material and permanent stained mounts, a useful technique
being to slip an empty hydrotheca onto a fine insect pin and rotate it into
different positions.
DESCRIPTION OF MATERIAL
Campanularia pecten sp. nov.
Fig. 1
Hoiotype: St James, False Bay, on the sea-grass Caulerpa filiformis (Suhr.),
just below L.W.S. Collected 29 March 1968. Catalogue number: SAM H1659.
Description
Hydrorhiza creeping and reticular, giving rise to solitary hydrothecae and
gonothecae. Hydrothecal pedicel smooth, constricted at base, with a single
1
Ann. S. Afr. Mus. 67 (1), 1975; 1-6, 2 figs.
pi ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 1. Campanularia pecten sp. nov.
a, colony on Caulerpa; b, hydrotheca and pedicel; c, hydrothecae, the centre one in end-on
view; d-g, sonophores in various stages of development; h, empty gonotheca, side view;
i, medusoid, showing from the centre outwards: opening to subumbrellar cavity, exumbrellar
aperture, ring of statocysts, gonads on radial canals.
TWO NEW SPECIES OF CAMPANULARIAN HYDROIDS FROM SOUTH AFRICA 3)
terminal spherule and with thickened perisarc. Irregular nodes sometimes present
due to regeneration.
Hydrotheca deep-campanulate, slightly compressed, with toothed margin.
Marginal teeth generally nine in number, rarely ten, usually narrower than the
bays between them, with bluntly rounded apices. Perisarc generally strongly
thickened on two opposite sides imparting a bilateral symmetry and an oval
cross-section. An annular perisarcal thickening present near base demarcating
a spherical basal chamber.
Gonotheca borne directly on hydrorhiza on a short, smooth pedicel,
scallop-shaped, rounded in broad view, opening around the circumference like
a bivalve shell and with one valve considerably flatter than the other, sometimes
with concentric ridges on the outer surface of the deeper valve. Gonophore
(only male present) eumedusoid, with four radial canals with the gonads
distributed along their length, with eight statocysts, without hypostome or
marginal tentacles, with a short free-living existence.
Measurements (mm)
Pedicel, length : : , Seog MS 0,52-1,74
maximum diameter. . : ; ; 0,10-0,20
Hydrotheca, depth A ; 0,51-0,69
diameter at mouth (broad view) oo ie eet 0,24-0,41
diameter at mouth (narrow view) . . .. . 0,30-0,33
Gonotheca, length including pedicel ties, ce te 0,93-1,09
maximum diameter. : 0,68-1,20
Remarks
In the early stages of development the gonotheca is oval in broad view,
becoming circular later. At this early stage the four branching radial canals
are clearly visible, but they become obscured later by the developing gonads.
Three eumedusae were released from the gonothecae in the laboratory and
were retained alive for 24 hours, during which period they reached a diameter
of 0,9-1,0 mm. They performed periodical spasms of contraction. The gonads
at this stage consisted of a number of small spherical aggregations along the
radial canals and were presumably partly spent. The exumbrellar surface of
the medusoid appeared to be pierced by an asymmetrical elliptical aperture.
Campanularia roberti* sp. nov.
Fig. 2
Holotype: Partridge Point, False Bay, on the weed Sargassum longifolium
(Turn.) attached to Ecklonia maxima (Osbeck). Collected 24 March 1968.
Catalogue number: SAM H1660.
* Named after Robert W. Day, who collected the material.
a ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 2. Campanularia roberti sp. nov.
a, hydrotheca and pedicel; b, hydrothecae, left in broad view, right in end-on view;
c-f, gonophores in various stages of development, f showing the branching radial canals;
g, mature gonotheca with gonophore containing planulae squeezed out of opening; h, upper
view of gonotheca showing concentric ridges.
TWO NEW SPECIES OF CAMPANULARIAN HYDROIDS FROM SOUTH AFRICA 5)
Description
Hydrorhiza creeping and reticular, giving rise to solitary hydrothecae and
gonothecae. Hydrothecal pedicel smooth, with a single terminal spherule, with
thickened perisarc. Regeneration nodes present in some.
Hydrotheca deep-campanulate, slightly compressed, with toothed margin.
Marginal teeth generally nine in number, rarely ten, triangular with bluntly
rounded apices. Perisarc generally strongly thickened on two opposite sides
and more so on one side than the other, imparting an oval cross-section and
an asymmetrical appearance when viewed from the broad side. An annular
perisarcal thickening present near base demarcating a spherical basal chamber.
Gonotheca borne directly on hydrorhiza on a short, smooth pedicel,
scallop-shaped, rounded in broad view, opening around the circumference
like a bivalve shell, recumbent and held with the flat lower surface against
the weed; upper valve curved above, thicker than lower valve, with distinct
concentric ridges when mature. Gonophore (only female present) in the form
of a fixed sporosac, with four branching radial canals but no other medusoid
structures, containing over 30 large eggs between the diverticuli of: the radial
canals. Eggs fertilized and developing into planulae within the gonotheca.
Measurements (mm)
Pedicel length oS Doll ae i a Mr lia ea ea 0,40-2,66
maximum diameter. : f ; 0,13-0,24
Hydrotheca, depth : . : : ; : 0,64-0,88
diameter at mouth (broad view) tar S Se eee ee 0,38-0,55
diameter at mouth (narrow view) . . .. . 0,30—0,46
Gonotheca, length including pedicel . ; 0,75—1,50
maximum diameter. : : : ; : 0,69-1,28
DISCUSSION
These two species are clearly closely related. Both belong to the hydranth
genus Orthopyxis L. Agassiz, 1862, and Campanularia pecten belongs to the
medusa genus Agastra Hartlaub, 1897. These two Benera are now commonly
included in Campanularia Lamarck, 1816 (Naumoy 1960; Vervoort 1972;
Millard & Bouillon 1974).
It is possible that C. pecten is the male and C. roberti the female of a single
species, for it is not unknown among the hydroids for sexual dimorphism to
occur and for the gonophore of one sex to reach a less advanced stage of
development than the other. However, there are certain differences in the
trophosome which distinguish the two and which support the retention of two
species at least until further material is forthcoming. Thus, the hydrotheca of
C. roberti is characteristically asymmetrical, with thicker perisarc on one
side than the other, whereas C. pecten has the perisarc symmetrically thickened
at the two narrow ends. Further, the hydrothecae of C. roberti are slightly
6 ANNALS OF THE SOUTH AFRICAN MUSEUM
larger than those of C. pecten, although there is an overlap between them.
Finally the marginal thecal teeth of C. roberti are in general slightly wider
than those of C. pecten.
REFERENCES
MILLARD, N. A. H. & BouILLon, J. 1974. A collection of hydroids from Mocambique, East
Africa. — Ann. S. Afr. Mus. 65: 1-40.
Naumov, D. V. 1960. Hydroids and Hydromedusae of the USSR.—Opred. Faune SSSR
70: 1-626. (In Russian. Translated by the Israel Program for Scientific Translations,
Jerusalem, 1969.)
VERVOORT, W. 1972. Hydroids from the Theta, Vema and Yelcho cruises of the Lamont-
Doherty Geological Observatory.— Zool. Verh. Leiden 120: 1-247.
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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A. GOW & N. A. H. MILLARD
TWO NEW SPECIES OF
CAMPANULARIAN HYDROIDS
FROM SOUTH AFRICA
pio
JOLt ME 67 PART 2 JUNE 1975 ISSN 0303-2515
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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. & RaFry, 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. 1960b. 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.
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ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 67 #2Band
June 1975 Junie
Part Zz Deel
A NEW DICYNODONT REPTILE FROM THE
TAPINOCEPHALUS ZONE (KAROO SYSTEM,
BEAUFORT SERIES) OF SOUTH AFRICA,
With EVIDENCE OF THE JAW, ADDUCTOR
MUSCULATURE
By
MICHAEL A. CLUVER
Cape Town Kaapstad
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A NEW DICYNODONT REPTILE FROM THE TAPINOCEPHALUS
ZONE (KAROO SYSTEM, BEAUFORT SERIES) OF SOUTH AFRICA,
WITH EVIDENCE OF THE JAW ADDUCTOR MUSCULATURE
By
MICHAEL A. CLUVER
South African Museum, Cape Town
(With 7 figures)
[MS accepted 15 August 1974]
ABSTRACT
A new dicynodont, Chelydontops altidentalis, from the farm Die Cypher, Beaufort West,
South Africa, is described from an incomplete skull and jaw in the South African Museum.
Important characteristics of the new form are the wide intertemporal region, the vaulted
anterior part of the secondary palate, the well-developed rows of teeth on the palate and
dentary, and a low but distinct coronoid process of the lower jaw. Upper and lower tooth
rows are bounded laterally by bony platforms, whose function is discussed in terms of jaw
musculature and mastication. The coroncid process on the lower jaw indicates that the medial
external adductor muscle of the lower jaw inserted on the rear of the dentary, as in the advanced
pelycosaur Dimetrodon. It is postulated that the dorsal surface of the dentary, lateral to the
tooth row, and the platform on the maxilla, lateral to the palatal tooth row, indicate the
presence in life of a reptilian cheek, such as recently proposed for certain ornithischian
dinosaurs. Similar but relatively larger palatal and dentary platforms are found in the
chronologically younger Endothiodon (including Esoterodon, Emydochampsa and Endogom-
phodon as junior synonyms) and Pachytegos, and it is proposed that the subfamily Endothi-
odontinae be enlarged to include Chelydontops and Prodicynodon. In terms of jaw function,
these genera appear to be distinct from other dicynodont groups.
CONTENTS
PAGE
Introduction . : : : F : s 7
Systematics : ; ‘ p : 8
Description of the material : A 9
Jaw musculature and function in Giehuoniops rie
Affinities of Chelydontops . : ‘ : ae 19
Summary : 2 : ‘ : : See
Acknowledgements . : : : , ei igeh el
References : : : : ‘ : = gemee
Abbreviations . : : : : : oS ae
INTRODUCTION
In his treatment of the South African Tapinocephalus zone Dicynodontia,
Boonstra (1948) briefly mentioned an unprepared skull (SAM-11558) which
he considered might eventually prove to be a new endothiodont. The specimen,
from the farm Die Cypher, Beaufort West, has recently been fully prepared
and clearly represents a new type of dicynodont, with Endothiodon and possibly
Prodicynodon its closest allies. A second specimen, a skull and jaw (SAM-—
12259) from Beukesplaas, Fraserburg, is generally similar to the Die Cypher
7
Ann. S. Afr. Mus. 67 (2), 1975: 7-23, 7 figs.
8 ANNALS OF THE SOUTH AFRICAN MUSEUM
skull, but differs in some respects; lateral crushing of this specimen has made a
close comparison with SAM-11558 difficult, and it has been used chiefly to
provide supplementary information in areas lacking in the type.
The dicynodonts of the Tapinocephalus zone were last reviewed by Boonstra
(1948) and Toerien (1953), but in the light of new classificatory criteria (Hotton
& Cluver, in preparation), the group is once more under investigation by the
present author. The specimens described and discussed below are, however,
systematically sufficiently far removed from their contemporaries to warrant a
separate, independent account.
SYSTEMATICS
As yet no satisfactory dicynodont classification exists. However, the
following scheme, modified from Haughton & Brink (1954) and Romer (1966),
may serve to indicate the systematic position of the form under consideration.
Class : Reptilia
Order : Therapsida
Suborder : Anomodontia
Infraorder : Dicynodontia
Family : Endothiodontidae
Subfamily : Endothiodontinae
Chelydontops gen. nov.
Diagnosis
Skull with wide intertemporal region and broadly exposed parietals. Pineal
opening large and situated on high boss. Maxilla rising high in snout, septo-
maxilla entirely within nostril. Caniniform process rudimentary, vestigial
caniniform tooth may be present. No anterior palatal ridges, anterior pre-
maxillary part of palate deeply vaulted. Palatine large, meeting premaxilla.
Posterior, palatal portion of maxilla raised to form prominent ledge, with
medial crest bearing nine robust teeth. Choanal vault wide and deeply recessed,
vomers forming narrow anterior septum. Long interpterygoidal vacuity. Medial
edge of dentary bearing row of ten teeth. Dentary extended posterodorsally
as a clear coronoid process.
Chelydontops altidentalis gen. et sp. nov.
Diagnosis: As for genus.
Type: Snout, occiput, partial lower jaw and cervical vertebrae in South African
Museum, Cape Town (Cat. No. 11558).
Locality: Die Cypher, Beaufort West, Cape Province.
Horizon: Tapinocephalus zone, Beaufort Series.
Collector: L. D. Boonstra.
A NEW DICYNODONT REPTILE FROM THE TAPINOCEPHALUS ZONE 9
DESCRIPTION OF THE MATERIAL
Type specimen (SAM-11558)
Skull
The skull is represented by the major portion of the snout and the occipital
segment up to the anterior border of the pineal foramen (Figs 1, 2, 3A). In
the skull roof the frontals, postorbital bars and zygomatic arches are lacking,
while in the ventral midline there is a gap in the pterygoid-basisphenoid central
stem at the level of the internal carotid artery foramina. There is thus no contact
between the front and rear halves of the skull, but as both portions agree
completely in size, in colour and in texture of bone and matrix, and were
(apparently) in close association when collected, there seems no reason to
doubt that they represent parts of a single individual.
pm.
lac. for
for mag.
Sq.
Denn.
——— EES |
Fig. 1. Chelydontops altidentalis gen. et sp. nov. Skull
reconstructed in dorsal view, missing areas shown in
outline. One-half natural size.
In the snout the premaxilla is smoothly curved and slopes far back between
the nasals. Behind the nostril the maxilla extends up to meet the nasal, and the
septomaxilla is confined to within the nostril. The nasals form low bosses
above the nostrils. Although neither side of the palatal rim is completely
10 ANNALS OF THE SOUTH AFRICAN MUSEUM
preserved, it is evident that no distinct caniniform process was present. On
both sides the base of a small marginal tooth is visible.
The lacrimal is a large bone in the antero-ventral corner of the orbit; a short
anterior process extends forwards on to the side of the snout to meet the nasal
in front and the maxilla below (Fig. 3A). The snout is not preserved above
this level, and the relationships of the frontals and pre-frontals could not be
determined. However, the base of the zygomatic arch can be seen clearly in
section on both sides. A large maxillary antrum is enclosed between the jugal
and lacrimal, and these two bones are in turn supported anteriorly and ventrally
by the maxilla and ectopterygoid. As in most dicynodonts, a large opening
between the inner edges of the lacrimals, jugals and palatines leads forward
from the orbital region into the bony nasal chamber.
The palate is marked by several distinctive features (Figs 2, 4). The palatal
rim is low and carries a weakly developed tooth in its rear, maxillary part.
There are no anterior palatal ridges on the premaxilla, but medially this bone
carries a prominent posterior ridge which meets the forked anterior edge of
the vomer. The anterior part of the palate and palatal rim is incomplete, but
it can nevertheless be seen that the premaxilla is deeply recessed in the anterior
midline so that, when complete, the secondary palate would have featured a
lat.pal.for
5 ane
EEE] ipt.vac.
q.ram.
Fig. 2. Chelydontops altidentalis gen. et sp. nov. Skull
reconstructed in ventral view, missing areas shown in
outline. One-half natural size.
A NEW DICYNODONT REPTILE FROM THE TAPINOCEPHALUS ZONE 11
high and narrow anterior vault. Much of the secondary palate is made up by
the premaxilla which, extending back to meet the anterior tip of the palatine,
excludes the maxilla from the choana. In palatal view, both premaxilla and
palatine are raised above the level of the adjoining maxilla, while laterally there
is a raised platform-like area on the maxilla medial to the posterior part of the
palatal rim (Fig. 2, max. sh.). This maxillary shelf, together with the palatal rim,
extends backwards to terminate posteriorly as a sharply defined crest below
the base of the zygomatic arch. The medial edge of the raised area bears an
irregular row of nine teeth, of which a few are newly erupted. The fully developed
teeth are relatively powerful, and the row extends back 17 mm from the
premaxilla—maxilla suture (where the base of the leading tooth is at least partly
surrounded by premaxilla) to the posterior limit of the maxillary shelf.
The palatal portion of the palatine is large and raised above the surface
of the surrounding premaxilla and maxilla. Posteriorly it lies high in the side
wall of the choana, medial to the ectopterygoid. This latter bone is strongly
po.
Sq.
pm:
cor. proc.
den.
Fig. 3. Chelydontops altidentalis gen. et. sp. nov. A. Skull reconstructed
in lateral view, missing areas shown in outline. B. Lower jaw fragment
in lateral view and (C) in dorsal view. All one-half natural size.
12 ANNALS OF THE SOUTH AFRICAN MUSEUM
developed, and consists of a lateral sheet, which lies beneath the base of the
zygomatic arch, and a posterior portion, which helps form part of the lateral
wall of the choana and appears to clasp the anterior edge of the pterygoid.
This section of the choanal side wall, composed of palatine, ectopterygoid and
pterygoid, is deep and the choana, seen as a whole, is a very wide and deeply
vaulted structure.
Behind its junction with the premaxilla, the vomer divides the internal
nares as a high, median septum, but postero-dorsally it bifurcates to enclose
the anterior part of the large (but incompletely preserved) interpterygoidal
vacuity.
In the occipital section the intertemporal part of the skull roof is preserved
up to the anterior margin of the pineal foramen. The postorbitals are well
separated by the parietals (Fig. 1) and lie mainly along the lateral margins
of the temporal roof. The broadly exposed parietals form a prominent boss
Fig. 4. Stereophotograph of palate of Chelydontops altidentalis gen. et sp. nov.
around the pineal foramen. The occiput is high, with the tripartite condyle
set at a relatively low level. A pair of deep basioccipital tubera, separated by a
median cleft, is present, and on the left side a slender stapes is partially preserved.
The squamosal, supporting the quadratojugal and quadrate ventrally, is of the
usual triradiate, dicynodont type.
A NEW DICYNODONT REPTILE FROM THE TAPINOCEPHALUS ZONE 13
Lower jaw
The middle portion (Fig. 3B, C) of each dentary is preserved and on the
left side the articular is in natural articulation with the quadrate. In each jaw
ten well-developed teeth are present, lying in an irregular row on the inside of
the dorsal edge of the dentary. A row of posterior serrations can be seen in
several of these teeth. Lateral to the tooth row the dorsal dentary surface is a
flat shelf (Fig. 3C, den. sh.), gently concave in lateral view. At the level of the
anterior dentary teeth, the inside edge of the dorsal dentary shelf is raised to
form the beginning of a high crest, which would have continued on to the missing
symphyseal region. Posteriorly the shelf extends past the end of the tooth row
and terminates on the anterior face of a small coronoid process, unique in
dicynodonts. From the tip of this coronoid process the dentary slopes back and
down to meet the surangular.
A fairly distinct lateral dentary shelf (Fig. 3B, lat. den. sh.), for insertion
of the lateral portion of the external adductor muscle, is present and overlies
a long and narrow mandibular fenestra. The articular of the left side, in contact
with the quadrate, appears to be of the normal dicynodont pattern.
SAM-12259
Lateral compression in this specimen has obscured some details of the
intertemporal region and palate, but there are strong resemblances with the
type specimen in the nature of the palatal rim, the maxillary part of the secondary
palate and the dentary.
The palatal rim is complete, and only a very modest caniniform process
is present. Somewhat behind the process is a marginal tooth, at the level of the
anterior choanal embayments. Medial to this the maxillary is thickened to form
a distinct platform, similar to that of the type specimen, and carries a number
of robust teeth along its inner border. Two large teeth and one smaller one
are present on the right-hand side, while on the left side there are three or
possibly four teeth; crushing and dislodging of the teeth of both upper and lower
jaws on this side has made aa exact assessment of the tooth number difficult.
den. sh.
art.
ang. 2cm.
Fig. 5. Chelydontops?sp. SAM-12259. Mandible
in lateral view. One-half natural size.
14 ANNALS OF THE SOUTH AFRICAN MUSEUM
As in the type, the palatal rim and maxillary platform lateral to the teeth are
extended back to a clearly marked termination below the base of the zygomatic
arch.
As far as can be seen, the anterior part of the palate slopes up fairly steeply
in front of the median premaxillary ridge, but lateral compression has made the
extent of this vaulting uncertain. More posteriorly, the pterygoids and
ectopterygoids of each side form a deep side wall to the choana, but the shape
of the vomer and the exact forward extent of the palatine is not clear.
The lower jaw (Fig. 5) is well preserved, and closely resembles what
remains of the type jaw. On the dorsal surface of the dentary there is a long,
wide and shallow trough, bounded by a distinct coronoid process posteriorly
and by sharp edges medially and laterally. The medial edge carries a row of
four teeth, which show a series of fine posterior serrations. In front of these
teeth the medial edge of the dentary is continued forwards for a short way as a
sharp crest, which merges anteriorly with the dorsal surface of symphysis.
At the symphysis the dorsal surfaces of the two jaw rami are separated
by a median cleft, but the front of the jaw is a fairly high and sharp blade,
notched on each side (Fig. 5, no.).
JAW MUSCULATURE AND FUNCTION IN CHELYDONTOPS
JAW MUSCULATURE
Crompton & Hotton (1967) have reconstructed the jaw musculature of the
dicynodonts Emydops and Lystrosaurus, and have analysed the probable range
of jaw movements during mastication. A few modifications of their interpreta-
tions have been suggested by Cluver (1971, 1974), these pertaining chiefly to
the crushing role played by the dentary during the final stages of the mastication
cycle, and to the insertion areas on the lower jaw of the jaw adductor muscle
fibres. Recently Barghusen (1968, 1972, 1973) has published a series of important
papers dealing with the adductor musculature and jaw mechanics in both
primitive and advanced synapsids, and many of his findings are relevant to
considerations of dicynodont jaw musculature.
Areas of origin of jaw adductor muscles
In dicynodonts the highly modified temporal region provides two large
areas of origin for the external adductor musculature. An inner division, the
medial external adductor muscle, arose from the lateral, posterior and medial
borders of the temporal fossa, these areas being formed by the squamosal and
postorbital (Fig. 6B). The outer surface of the postorbital, forming the dorsal,
inner border of the temporal fenestra, merges smoothly with the upper surface
of the squamosal, where this bone forms the posterior border of the fenestra.
This surface is continued laterally and forward on to the inside of the zygomatic
arch, and in this way an extensive area of origin is formed.
A NEW DYCINODONT REPTILE FROM THE TAPINOCEPHALUS ZONE 15
Laterally to the temporal fossa the squamosal extends back and outwards
as a characteristic external plate below the rear of the zygomatic arch. The
anterior surface of this smoothly moulded sheet is continued dorsally on to
the lateral face of the zygomatic arch, and includes the anterior face of the
quadratojugal ventrally. This substantial area served for the origin of the lateral
external adductor muscle (Fig. 6C), and among therapsids is found only in
dicynodonts.
add. ext. med.
leer
OT
| WHY
My) Wy,
.. biG
SSSx~
3 SS
SSF S
post. pt.
Fig. 6. Chelydontops altidentalis. Skull reconstructed in lateral view to show (A) pseudotemporalis
and posterior pterygoideus muscles, (B) medial external adductor and posterior adductor
muscles and (C) lateral external adductor muscle and cheek.
The degree of development of the intertemporal part of the skull roof
varies considerably in dicynodonts. In a genus such as Daptocephalus the
parietals are greatly reduced on the skull roof and the postorbitals approach
each other closely (Ewer 1961), while in cistecephalids (Keyser 1973; Cluver
1974) the parietals are broad elements and the postorbitals and temporal
fenestrae are widely separated. However, a constant feature in all dicynodonts
is a clearly demarcated recess beneath the lateral edge of the inter-temporal
16 ANNALS OF THE SOUTH AFRICAN MUSEUM
skull roof, bounded medially by a ventral flange of the parietal and, more
anteriorly, by the expanded dorsal part of the epipterygoid. This recess is
similar in position to the “medial depression’ described by Barghusen (1973:
829) in the undersurface of the skull roof of the advanced pelycosaur
Dimetrodon. Barghusen considered the depression to be the site of origin of the
pseudotemporalis muscle of the internal jaw adductor group (Fig. 6A, ps.),
which presumably also attached to the epipterygoid and parietal.
In addition, the area of origin of a posterior adductor muscle, also part
of the internal jaw adductor musculature, can be inferred. In recent reptiles
this muscle arises from the medial part of the quadrate bone, and Barghusen
(1973: 831) reconstructs it in Dimetrodon as arising from ‘the lateral face of the
well-developed anterior process of the quadrate which contacts the pterygoid
bone’. In a dicynodont such as Chelydontops the portion of the quadrate above
the condyle is a high and broad sheet lying up against the squamosal and the
posterior end of the quadrate ramus of the pterygoid, and is well situated to
serve as a site of origin of a muscle, such as the posterior adductor, running to
the inside of the lower jaw (Fig. 6B, add. post.).
In dicynodonts, the posterior pterygoideus muscle (sensu Barghusen 1973)
of the internal adductor group very likely arose from the ventro-lateral edge
of the pterygoid, where this forms the lateral boundary of the choana, and from
the pterygoid’s quadrate ramus (Fig. 6A, post. pt.). Such a posterior pterygoideus
muscle would include both the anterior and posterior pterygoideus muscles of
Crompton & Hotton (1967) and Cluver (1971).
Areas of insertion of jaw adductor muscles
The adductor musculature’s attachment areas on the lower jaw are more
clearly indicated in Chelydontops than in other dicynodonts. It seems most
likely that the lateral dentary shelf, present in varying degrees of prominence
above the mandibular fenestra in all dicynodonts (Crompton & Hotton 1967;
Cluver 1970, 1971, 1974) marks the site of insertion of the lateral external
adductor muscle (Fig. 6C). Such a purely lateral insertion area corresponds
well with the lateral position of the area of origin of this muscle in dicynodonts.
The insertion area of the medial external adductor muscle in Chelydontops
is of special significance. The low but distinct coronoid process on the rear end
of the dentary is strong evidence of a tendinous attachment, and it seems very
likely that the large medial division of the external adductor muscle (Fig. 6B)
inserted on to the lower jaw via a ‘bodenaponeurosis’, such as reconstructed by
Barghusen (1973) in Dimetrodon, on the basis of comparisons with recent rep-
tiles. This is in contrast with Crompton & Hotton’s (1967) restoration, according
to which the medial external adductor muscle inserts into a deep groove in the
dorsal edge of the dentary behind the symphyseal region.
The pseudotemporalis muscle appears to have had a fleshy attachment
on the inside of the lower jaw above the adductor (Meckelian) fossa. In
Chelydontops the surangular and rear part of the dentary are smoothly excavated
A NEW DICYNODONT REPTILE FROM THE TAPINOCEPHALUS ZONE 1
above this fossa and, in fact, it is likely that it was to accommodate a changed
orientation of the pseudotemporalis muscle that the dorsal part of the lower
jaw is bowed laterally in cistecephalids (Cluver 1974).
The adductor posterior muscle, arising from the anteromedial face of the
quadrate, has a clear insertion site in and around the adductor fossa, while the
pterygoideus posterior muscle (Fig. 6A) extended to the lower edge of the jaw
and inserted on the lateral face of the angular, at least partly beneath the
reflected lamina (Crompton & Hotton 1967; Cluver 1971; Barghusen 1973).
A posterior slip of this muscle probably inserted on the inner surface of the
retroarticular process.
With one exception, the jaw adductor musculature thus reconstructed in
Chelydontops is essentially comparable with that restored in Dimetrodon by
Barghusen (1968, 1973) and is readily derivable from a reptilian model. The
exception is the lateral division of the external adductor muscle, this representing
a basic dicynodont innovation. Moreover, it seems probable that the insertion
of the medial external adductor in Chelydontops is typical in its position
(although exaggerated in form) since the rear end of the dentary was the most
anterior area of insertion of the external adductor muscle in primitive synapsids
(Barghusen 1973).
In Chelydontops the slightly excavated dorsal surface of the dentary behind
the symphysis may be compared with the deep dorsal dentary groove seen in
some other dicynodonts (Crompton & Hotton 1967; Cluver 1970, 1971) but,
lying as it does in front of the coronoid process and directly lateral to the tooth
row, it seems unlikely that it served as an area of muscle attachment. In
Chelydontops this part of the dentary should more properly be considered in
conjunction with the raised area of maxilla lateral to the row of palatal teeth.
As shown below, it is unlikely that more than the most medial part of this
platform was directly involved in any chewing activity and, in effect, there is
a space between the upper and lower tooth rows and the side of the snout.
This structural arrangement is perhaps best explained in terms of the
development of a reptilian cheek, such as reconstructed by Galton (1973) in
certain ornithischian dinosaurs which show evidence of similar platforms lateral
to the tooth row. In Chelydontops, evidence suggesting the presence of a cheek
consists of the crest running back from the weak caniniform process to the rear
of the maxillary platform. This crest can be interpreted as marking the dorsal
line of attachment of the cheek, while its ventral attachment was most likely
along the sharp lateral edge of the dorsal dentary platform. A cheek of this sort
would have occupied more or less the same position as the ‘Mundplatt’
reconstructed by Crompton & Hotton (1967) in Emydops. In fact, modification
of the reptilian ‘Mundplatt’ offers a simple derivation for the cheek reconstructed
in Chelydontops (Fig. 6C).
The actual structure of the cheek reconstructed for Chelydontops is
uncertain, but the possibility that it was muscular does exist. Galton (1973)
suggests that musculature in his reconstructed ornithischian cheek might easily
18 ANNALS OF THE SOUTH AFRICAN MUSEUM
have been derived from either the reptilian levator angularis oris muscle, or
the pars superficialis of the reptilian external adductor jaw musculature—a
similar derivation is possible in Chelydontops. Crompton & Hotton (1967)
reconstructed a levator oris muscle arising from the infraorbital bar in Emydops
and inserting on the ‘Mundplatt’. As shown by Galton (1973) it is not necessary
to draw an analogy with mammals to postulate the presence of a muscular cheek
in reptiles. When a mammalian type muscular cheek, innervated by the facial
nerve, arose is uncertain; if such an event occurred in therapsids, it may have
been a late development in the most progressive of therapsid lines, i.e. the
cynodonts.
JAW FUNCTION
Cheek function in a reptile such as Chelydontops should be considered
in conjunction with the probable range of jaw movements during the masticatory
cycle. As shown by Crompton & Hotton (1967), the power stroke during
dicynodont mastication occurred during retraction of the lower jaw, and this
would certainly have been true in Chelydontops. While some slicing would
very likely have occurred between the lateral sides of the lower jaw beak and
the palatal rim, the batteries of powerful mandibular and palatal teeth suggest
that considerable slicing and cutting between these teeth took place towards
the end of the stroke. Both upper and lower tooth rows (best preserved in the
type specimen) are obliquely orientated, with the anterior teeth of each row.
lying medial to the posterior teeth. Upper and lower tooth rows are practically
the same distance apart, and it is apparent that strictly orthal retraction would
cease when the mandibular tooth rows meet the palatal tooth rows in face-to-face
contact. This arrangement would prevent the mandibular teeth from immediately
reaching the inflated palatal portions of the palatines, so obviously involved in
the mastication process in other dicynodonts (Crompton & Hotton 1967;
Cluver 1971).
A sideways displacement of the mandible on the quadrates, to permit the
upper and lower tooth rows to slide past each other, would result in a complex
series of tooth-to-tooth and tooth-to-beak contacts coming into play, during
which only one palatine at a time would be involved. However, movement of
this type would be possible only if the articulation between the quadrate and
lower jaw lacked the tightly interlocking articular facets seen in most dicynodonts
(Cluver 1974). Since, as far as can be seen, this is not the case in Chelydontops,
an alternative possibility involving two separate phases of masticatory activity
may be considered. Thus, apart from a shearing phase during which the lower
jaw is drawn back from a protracted position until the mandibular teeth meet
the palatal tooth rows, a purely up-and-down biting action, involving both
tooth rows and the palatines, could have taken place with the lower jaw in an
almost fully retracted position.
With only incomplete material available, the range of lower jaw movements
during mastication must remain uncertain, but it seems clear that the upper and
A NEW DICYNODONT REPTILE FROM THE TAPINOCEPHALUS ZONE 19
lower tooth rows played an important part in the process. Retention of, and
control over, partly chewed plant matter, which would tend to move to the side
of the mouth, could become a critical factor, and the need for a cutaneous or
muscular sheet in the corner of the mouth might very likely arise. A non-
contractile cheek would serve merely to retain masticated food until this could
be drawn back into the oral cavity by the tongue. A muscular cheek would be
capable of actively assisting in mastication by returning partly-chewed material
to the teeth. It is noteworthy that only in forms where the palatal and mandi-
bular teeth are strongly developed, such as Chelydontops and Endothiodon, is a
lateral maxillary shelf formed; to this extent, jaw function sets these genera
apart from other dicynodonts.
AFFINITIES OF CHELYDONTOPS
In several of its characters Chelydontops closely resembles members of
the sub-family Endothiodontinae, which, according to Cox (1964), includes
the genera Endothiodon (with Esoterodon, Emydochampsa and Endogomphodon
as junior synonyms) and Pachytegos, the latter from the Ruhuhu Valley of
Tanzania. Resemblances between Chelydontops and Endothiodon involve features
of the palate and mandible. Thus, the anterior part of the secondary palate in
Endothiodon is deeply vaulted as in Chelydontops, and there is no true caniniform
process. A ventral process of the anterior, premaxillary part of the palatal rim,
characteristic of Endothiodon, is seen also in SAM-12259—this portion of
the palatal rim is incomplete in the Chelydontops type specimen. Prominent
features in both the palate and mandible of Endothiodon are the wide and shallow
troughs lying lateral to the upper and lower tooth rows. In the palate (Fig. 7A)
this is especially marked, and the shelf of each side is bounded by a distinct
crest which is extended back and outwards below the zygomatic arch as in
Chelydontops. Cox (1964) mentions this feature, which may be regarded as an
expanded equivalent of the maxillary platform of Chelydontops and indication
of a fairly extensive cheek. The palatal part of the premaxilla in Endothiodon
also extends far back, but does not meet the palatine as it does in Chelydontops.
A point of importance is the rooting in Endothiodon of the first two palatal
teeth in the rear of the premaxilla. Unfortunately it was not possible to determine
with certainty whether the first tooth in Chelydontops is placed in the premaxilla
in the same way, but it is clear that at least part of the base of this tooth is
surrounded by the premaxilla.
Although these are for the most part important resemblances, Chelydontops
differs sharply from Endothiodon in the retention of a broad intertemporal
region, with widely exposed parietals, and also in the apparently smoothly
rounded snout, without the longitudinal ridges so characteristic of the dorsal
surface of the Endothiodon snout.
Besides Endothiodon, Chelydontops shares several features with the poorly-
known genus Prodicynodon (Broom 1912). The type specimen of Prodicynodon
20 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 7. Endothiodon uniseries BMNH-49414. Palatal
reconstruction of type specimen. B. Prodicynodon beaufort-
ensis AMNH-S5509. Palatal reconstruction of type
specimen. Both one-half natural size.
pearstonensis (AM-2551) is too crushed and incomplete to allow adequate
comparisons to be made, but recent preparation of the type specimen of
P. beaufortensis (AMNH-5509) shows that in addition to the pointed anterior
beak of the lower jaw, typical of Endothiodon (Cox 1964), this form has a row
of Endothiodon-like teeth bounded laterally by a wide, shallow trough (Fig. 7B).
A small marginal tooth lies on the palatal rim, in the same position as in
Chelydontops, and a caniniform process is absent. However, a weak ventral
process at the maxilla—premaxilla meeting on the palatal rim resembles that of
Endothiodon and Chelydontops. In the palate the premaxilla is vaulted in the
endothiodontinid manner to receive the anterior tip of the lower beak, while a
notch behind this anterior tip is reminiscent of the condition in SAM-12259.
A NEW DICYNODONT REPTILE FROM THE TAPINOCEPHALUS ZONE 21
Cox (1964) recognizes two genera in the subfamily Endothiodontinae, these
being Endothiodon itself, from the Endothiodon zone (Cistecephalus zone of
Kitching 1970) and Pachytegos from the Ruhuhu Beds of Tanzania. It now
seems necessary to include Chelydontops and Prodicynodon in this subfamily.
Prodicynodon pearstonensis is from Pearston, and Prodicynodon beaufortensis
is from Kuilspoort, Beaufort West; both are probably from Endothiodon zone
(Kitching’s Cistecephalus zone) strata. Chelydontops, from the Tapinocephalus
zone of Beaufort West, represents the earliest known member of this distinctive
group of dicynodonts.
SUMMARY
The skull and mandible of Chelydontops altidentalis from the Tapinocephalus
zone of the Beaufort Series show strong affinities with Endothiodon and
Prodicynodon, and, together with Pachytegos of Tanzania, these three genera
should be included in the subfamily Endothiodontinae. While several conserva-
tive characters and its low stratigraphic occurrence make Chelydontops a
primitive (but not necessarily ancestral) member of the subfamily, a functional
reconstruction of the skull helps interpret the characteristic skull structure of
the group as a whole. A clear coronoid process on the dentary allows a
reconstruction of the external jaw adductor muscles to be made with fair
confidence, while a shelf lateral to each upper tooth row is taken as indication
of the former presence of a reptilian cheek, such as has been recently proposed
for ornithischian dinosaurs. There is a possibility that fibres from either the
levator angularis oris or adductor externus lateralis muscles had invaded
the primarily non-contractile cheek, which was very likely derived from a
reptilian ‘Mundplatt’.
With the identification of Chelydontops altidentalis, the Endothiodon grade
of dicynodont development can be traced farther back into the Permian than
before, and since the functional innovations seen in Chelydontops are continued
and improved upon in the later genera, this grade can now be more closely
defined and better understood.
ACKNOWLEDGEMENTS
I am indebted to Drs A. J. Charig and Eugene S. Gaffney of the
Departments of Vertebrate Palaeontology of, respectively, the British Museum
(Natural History), London, and the Amezican Museum of Natural History,
New York, for permission to study and prepare specimens in their charge,
and to Mr Gilbert Stucker of the American Museum of Natural History for
the preparation of the type skull of Prodicynodon beaufortensis. A study grant
from the South African Council for Scientific and Industrial Research and the
Trustees of the South African Museum enabled me to visit the above institutions
during 1972/73.
22 ANNALS OF THE SOUTH AFRICAN MUSEUM
For the loan of the type specimen of Prodicynodon pearstonensis 1 am
indebted to the authorities of the Albany Museum, Grahamstown. The
preparation and photography of the type specimen of Chelydontops altidentalis
and the related specimen SAM-12259 were undertaken by, respectively,
Imogen M. Chesselet and Neville J. Eden, of the Department of Palaeontology,
South African Museum.
REFERENCES
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. 1972. The origin of the mammalian jaw apparatus. In: G. H. SCHUMACHER.
Morphology of the maxillo-mandibular apparatus: 26-32. Leipzig: VEB Georg Thieme.
BARGHUSEN, H. R. 1973. The adductor jaw musculature of Dimetrodon (Reptilia, Pely-
cosauria).—J. Paleont. 47: 823-834.
BoonsTrA, L. D. 1948. On the anomodont reptiles from the Tapinocephalus zone of the Karroo
System. Jn: ROYAL SOCIETY OF SOUTH AFRICA. Robert Broom commemorative volume:
57-64. Cape Town: Royal Society of South Africa. (Special publication.)
Broom, R. 1912. On some fossil reptiles from the Permian and Triassic beds of South Africa. —
Proc. zool. Soc. Lond. 1912: 859-876.
CLuver, M. A. 1970. The palate and mandible in some specimens of Dicynodon testudirostris
Broom & Haughton (Reptilia, Therapsida).— Ann. S. Afr. Mus. 56: 155-274.
CLuverR, M. A. 1971. The cranial morphology of the dicynodont genus Lystrosaurus.— Ann.
S. Afr. Mus. 56: 155-274.
CLuver, M. A. 1974. The skull and mandible of a new cistecephalid dicynodont.— Ann. S. Afr.
Mus. 64: 137-155.
Cox, C. B. 1964. On the palate, dentition and classification of the fossil reptile Endothiodon
and related genera.—Am. Mus. Novit. 2171: 1-25.
CROMPTON, A. W. & Hotton, N. 1967. Functional morphology of the masticatory apparatus
of two dicynodonts (Reptilia, Therapsida).—Postilla 109: 1-51.
Ewer, R. F. 1961. The anatomy of the anomodont Daptocephalus leoniceps (Owen).— Proc.
zool. Soc. Lond. 136: 375-402.
GALTON, P. M. 1973. The cheeks of ornithischian dinosaurs. — Lethaia 6: 67-89.
HAUGHTON, S. H. & Brink, A. S. 1954. A bibliographic list of Reptilia from the Karroo beds
of South Africa.—Palaeont. afr. 2: 1-187.
Keyser, A. W. 1973. A preliminary study of the type area of the Cistecephalus zone of the
Beaufort Series, and a revision of the anomodont family Cistecephalidae.— Mem. geol.
Surv. Rep. S. Afr. 62: 1-72.
KITCHING, J. W. 1972. A short review of the Beaufort zoning in South Africa. In: INTER-
NATIONAL UNION OF GEOLOGICAL SCIENCES. Commission on Stratigraphy. Subcommission
on Gondwana Stratigraphy and Palaeontology. Gondwana Symposium 2nd, South Africa,
1970. Proceedings and papers: 309-311. Pretoria: CSIR.
Romer, A. S. 1966. Vertebrate paleontology. Chicago: University of Chicago Press.
TOERIEN, M. J. 1953. The evolution of the palate in South African anomodonts, and its
classificatory significance.—Palaeont. afr. 1: 49-117.
ABBREVIATIONS
add. ext. lat. . lateral division of external adductor muscle
add. ext. med. . medial division of external adductor muscle
add. post. . . posterior adductor muscle
ang. . ; . angular
art; : . articular
bog: : . basioccipital
A NEW DICYNODONT REPTILE FROM THE TAPINOCEPHALUS ZONE
CAs « ‘
cor. proc. .
den...
den. sh.
ect.
eoc. .
for. mag.
ipt. vac.
sug...
lac.
lac. for.
lat. den. sh.
lat. pal. for.
max. ;
max. sh.
med. pm. r.
has...
cheek
coronoid process
dentary
dentary shelf
ectopterygoid
exoccipital
foramen magnum
interpterygoidal vacuity
jugal
lacrimal
lacrimal foramen
lateral dentary shelf
lateral palatal foramen
maxilla
maxillary shelf
median premaxillary ridge
nasal
notch
parietal
palatine
pineal foramen
premaxilla
postorbital
posterior pterygoideus muscle
pseudotemporalis muscle
pterygoid
quadrate
quadratojugal
quadrate ramus of pterygoid
surangular
squamosal
stapes
vomer
Albany Museum, Grahamstown
American Museum of Natural History, New York
British Museum (Natural History), London
South African Museum, Cape Town
23
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
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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:
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In describing new species, one specimen must be designated as the holotype; other speci-
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SAM-A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach,
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Note standard form of writing South African Museum registration numbers and of date.
7. SPECIAL HOUSE RULES
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‘Revision of the Crustacea. Part VIII. The Amphipoda.’
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MICHAEL A. CEMVER
A NEW DICYNODONT REPTILE FROM THE
TAPINOCEPHALUS ZONE (KAROO SYSTEM,
BEAUFORT SERIES) OF SOUTH AFRICA,
WITH EVIDENCE OF THE JAW ADDUCTOR
MUSCULATURE
|
VOLUME 67 PART 3 JUNE 1975 ISSN 0303-2515
~ ANNALS
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.
Koun, 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.
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 67 Band
June 1975 Junie
Part 3 Deel
TAXONOMIC STATUS OF THE
PYGOCEPHALOMORPHIC CRUSTACEA
FROM THE DWYKA ‘WHITE BAND’
(PERMO-CARBONIFEROUS) OF SOUTH AFRICA
By
BRIAN KENSLEY
Cape Town Kaapstad
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TAXONOMIC STATUS OF THE PYGOCEPHALOMORPHIC
CRUSTACEA FROM THE DWYKA ‘WHITE BAND’
(PERMO-CARBONIFEROUS) OF SOUTH AFRICA
By
BRIAN KENSLEY
South African Museum, Cape Town
(With 4 figures)
[MS accepted 14 October 1974]
ABSTRACT
The taxonomic status of Notocaris tapscotti Broom, a pygocephalomorphic crustacean
from the Dwyka ‘White Band’ of South Africa is discussed, a reconstruction of the external
morphology suggested and new collecting localities recorded.
CONTENTS
PAGE
Introduction . 2 : : 2 : aw? 25
Systematics . : : ; ‘ ; epee
Material and localities . : ; ; re: |
References : ’ ; : : : Peake Ps
INTRODUCTION
Since 1923, when Woods gave a brief description of a peracaridean crusta-
cean from Kimberley under the generic name Pygocephalus, some confusion has
arisen regarding the status of this organism. Additional localities of this
crustacean have since been found, this new material allowing the description of
the fossil to be supplemented. The purpose of this paper is to record these
localities and to attempt to resolve the taxonomic confusion.
The following works refer to Dwyka pygocephalomorphic crustaceans from
South Africa:
Rogers & Du Toit (1909: 193) mention the presence of crustacean fossils
‘probably Anthrapalaemon’ in the Upper Dwyka shales, while Haughton (1919)
records Anthrapalaemon from the Dwyka White Band at Orange River Station.
Woods (1923) recorded a Pygocephalus from the Dwyka White Band at
Kimberley, but did not provide a specific name.
Broom (1931) re-examined Woods’ material and gave the species the name
Notocaris tapscotti n. gen. et sp.
Du Toit (1954) and Haughton (1969) merely repeat the previous records of
Notocaris and Anthrapalaemon.
Fabre (1967) described Pygaspis ginsburgi from material collected at
Laingsburg, Cape, also from the Dwyka White Band.
25
Ann. S. Afr. Mus. 67 (3), 1975: 25-33, 4 figs.
26 ANNALS OF THE SOUTH AFRICAN MUSEUM
Secretan (1967) in a discussion of lines of evolution in the Archaeostraca,
deals with Pygaspis ginsburgi, its tagmatization and segmentation.
Brooks (1969) synonymized Pygaspis Beurlen (type species P. brasiliensis)
and Liocaris Beurlen with Paulocaris Clarke. Brooks incorrectly ascribed the
specific name tapscotti to Woods. Pinto (1971), however, re-examined the South
American species of Pygaspis, Liocaris, and Paulocaris, and concluded that the
three genera should be maintained. He also tentatively suggested that Pygaspis
(Beurlen 1934) was a synonym of Notocaris (Broom 1931). Savage (1971)
described trails from the Dwyka of Natal and related them to syncaridan and
peracaridan crustaceans, although not actually correlating the trails with any
Pygocephalomorpha.
McLachlan & Anderson (1973) review and contribute to the evidence for
marine conditions during Dwyka times in South Africa, but note that by the
time of the deposition of the White Band (Upper Carboniferous or Lower
Permian), with its associated crustaceans, palaeoniscid fish, and the small
aquatic reptile Mesosaurus, conditions were probably non-marine. (This
conclusion has by no means been fully proven.)
In their list of Dwyka invertebrate fossils (1973: 54) McLachlan &
Anderson conclude that the names Anthrapalaemon, Pygaspis ginsburgi and
Notocaris tapscotti probably refer to the same crustacean.
SYSTEMATICS
Superclass CRUSTACEA
Class MALACOSTRACA
Subclass EUMALACOSTRACA
Superorder EOCARIDA
Order pYGOCEPHALOMORPHA
Family Pygaspidae
Genus Notocaris Broom, 1931
Generic diagnosis
Carapace lacking mid-dorsal carina, cervical groove, and tubercles, but
possessing 1-3 antero-lateral spines, as well as a suprabranchial suture.
Abdomen reflexed, somewhat reduced.
Discussion
Pinto (1971) notes that it is necessary to compare ‘Pygaspis’ and Notocaris,
as these may well be synonymous. Comparison of Pygaspis brasiliensis and
Notocaris tapscotti shows that several differences exist in the nature of the
carapaces. In the former species the carapace is covered with tiny tubercles,
especially well developed at the cervical sulcus and dorsal carina. There are no
antero-lateral spines, but five or six spaced lateral spines, while a bifurcate
cervical sulcus curves posteriorly in the medio-dorsal region, forming a smooth
TAXONOMIC STATUS OF THE PYGOCEPHALOMORPHIC CRUSTACEA 27
carina which stretches almost to the posterior margin. In Notocaris there is a
well-developed antero-lateral spine (not always preserved or visible in many
specimens) as well as one or two lateral spines. The carapace, however, bears no
sign of tubercles, a cervical groove, or median carina, but signs of a longitudinal
suprabranchial suture can be detected on a few specimens.
Whether these characters are of generic or infrageneric significance amongst
the pygocephalomorphic crustaceans requires investigation, as indeed does the
whole generic complex within this group. Presence or absence of carapace
grooves, carinae, and spines amongst living crustaceans, e.g. the Caridean
decapods, is of definite value in generic separation. If this applies in the present
case, the South American Pygaspis and the South African Notocaris must be
maintained as valid genera.
Notocaris tapscotti Broom
(Figs 1-4)
cml,
ss &
Fig. 1. Dorsal/ventral reconstruction of Notocaris tapscotti.
28 ANNALS OF THE SOUTH AFRICAN MUSEUM
A
Fig. 2. A. Reconstruction of ventral view of female thorax, showing oostegites. B. Pleon in
lateral view.
Description
Exoskeleton very thin. Carapace almost circular (when compressed),
apparently attached to the body only in the anterior region. No sign of a
medio-dorsal carina. Rostrum short, triangular. Antero-lateral corner of
carapace bearing one small acute spine; occasionally a second or even a third
tiny spine may be seen posterior to the antero-lateral spine. Anterior margin of
carapace slightly concave. A faint longitudinal suprabranchial suture, stretching
from the region of the anterior margin to the posterior margin may be seen on
the carapace of some specimens. Posterior margin of carapace dorsally concave.
Abdomen usually under thorax, consisting of five segments plus telson. (A
reduced first segment, hidden by the posterior part of the thorax and carapace
may be present.) Pleura of abdominal segments apically acute, apices directed
posteriorly.
Antennule with 3-segmented peduncle, basal segment equal in length to
two distal segments together, peduncle extending beyond antennal scaphocerite.
Inner flagellum slightly shorter than outer, latter only slightly longer than
peduncle.
Antennal scaphocerite elongate-oval, reaching to end of second segment of
antennular peduncle. Single flagellum reaching posteriorly to level of posterior
carapace margin.
Eyestalks slender, situated dorsal to the antennae.
Epistome ventral, elongate-oval, separating the mandibles. Position and
structure of the mouthparts uncertain. An elongate slender mandibular palp
Fig. 3. Notocaris tapscotti. (All figures natural size.) A—D. Laingsburg. E. Ratelklip, Calvinia.
F_-]. Kimberley.
A. Specimen showing ventral sternites and eyestalks. B-D. The Laingsburg specimens
preserved in dark grey shale, appear to have a ‘smear’ of white material surrounding and
obscuring the fossil and its impressions. E. Specimen showing eyestalks. F—G. Specimens in
lateral view. Note pleonal pleura, peraeopodal exopodites, and antennae. H. Specimen
showing typically enrolled pleon. I. Carapace in lateral view. Note also antennal scaphocerites.
TAXONOMIC STATUS OF THE PYGOCEPHALOMORPHIC CRUSTACEA 29
Legend to Figure 3 on opposite page
ANNALS OF THE SOUTH AFRICAN MUSEUM
30
Legend to Figure 4 on opposite page.
TAXONOMIC STATUS OF THE PYGOCEPHALOMORPHIC CRUSTACEA 31
present, the distal portion protruding beyond the anterior carapace margin,
medial to the antennular peduncles.
Seven posterior appendages essentially similar, biramous. Exopod con-.
sisting of an elongate basal segment, two short distal segments, plus plume-like
flagellum. Endopod longer than exopod, consisting of short dactyl, slender
propodus one-third longer than carpus, three or four proximal segments
indistinct. In female, seven pairs of overlapping roughly circular oostegites,
those of opposite sides not meeting, the brood pouch thus remaining open.
Seven series of gills present around the bases of the peraeopods, exact structure
indistinct.
Pleopods seldom seen, biramous, consisting of short protopod and two
plume-like rami.
Uropods consisting of short basal segment plus two broad rami; outer
ramus shorter and narrower than inner.
Telson triangular, bearing three lateral spines plus an articulating median
spine extending beyond furcal lobes. Latter flattened, strongly curved. There is
still some uncertainty about the exact proportions of the uropodal rami.
MATERIAL AND LOCALITIES
Carapace length range
(including rostrum)
Kimberley, Cape . : : : . 28.45S, 24.46E 7-27 mm
Mmenpspure, Cape. . . . . 33.128, 20.51E 14-19 mm
Loeriesfontein, Cape . E - ,30:59S8, 19.29E 13-28 mm
Ratelklip, district Calvinia, Cape . 32.02S, 19.48E 13 mm
Orange River Station district Hope-
fown,Cape . . . . . _ 29.388, 24.15E (material not seen)
Discussion
The presence of two closely-related genera, viz. Notocaris and Pygaspis,
each represented by a single very similar species from the same stratum (i.e. the
White Band) of the Dwyka of the Great Karoo Basin (McLachlan & Anderson
1973) demands close scrutiny. Fabre (1967), in discussing the affinities of his new
species, P. ginsburgi, concludes that it is closely related to the South American
P. brasiliensis. Regarding Notocaris, Fabre mentions having examined some
material of N. tapscotti in the Albany Museum, Grahamstown, but did not see
any of the Kimberley material. He notes similarities in size, appendages, and
Fig. 4. Notocaris tapscotti (All figures natural size.) A-J. Loeriesfontein.
A. Carapace showing short triangular rostrum and acute anterolateral angles. B. Carapace
showing rostrum, antero-lateral spines, and supra-branchial suture. C. Note antennal peduncle
structure. D. Specimen with extended pleon, in dorsal view. E. Note peraeopodal exopodite
structure. F. Note antennal structure. G. Detached pleon above; possibly a single pleonal
segment with its attached pleopods below. H. Extended pleon. Note telsonic and uropodal
structure. I-J. Note antennal and antennular structure.
32 ANNALS OF THE SOUTH AFRICAN MUSEUM
general structure but without closer examination of Notocaris material, prefers
to place his material in the genus Pygaspis. |
Examination of type material of Notocaris tapscotti from Kimberley, the
type material of Pygaspis ginsburgi from Laingsburg, as well as additional
material from Loeriesfontein, Calvinia and Laingsburg, Cape, makes it obvious
that only one species is involved here. Fabre observes that Woods mentions
enrolled individuals (individus enroulés) and that this could not clearly be seen
in the Laingsburg specimens. In fact, Woods meant the flexure of the abdomen,
and from Fabre’s figures 2 and 3, it is obvious that his material also showed this
flexure, although the ventral view (fig. 3) is somewhat confusing regarding the
uropod and telsonic structure. No differences regarding general form, or
structure of the appendages, can be detected, which would suggest that not more
than one species is involved amongst the material from Kimberley, Laingsburg,
and Loeriesfontein. As Broom’s name has precedence, all the known pygo-
cephalomorphic crustaceans from the abovementioned localities must therefore
bear the name Notocaris tapscotti, while Pygaspis ginsburgi becomes a synonym
of this earlier name.
ACKNOWLEDGEMENTS
I am grateful to Messrs W. A. and E. Loubser of Loeriesfontein for
permission to collect on their farm, and for their hospitality, and to Dr M.
Cluver and Mr M. R. Cooper of the Department of Palaeontology of the South
African Museum, for assistance with collecting material of Notocaris.
My thanks are due to Dr J. N. Theron of the Geological Survey, and the
Director of the Alexander McGregor Memorial Museum, Kimberley, for the
loan of material.
My sincere thanks are due to Professor I. Damiani Pinto of the Universidade
Federal do Rio Grande do Sul, Brazil, and Dr Sylvie Secretan of the Institut de
Paléontologie, Paris, for reading the manuscript and for their many useful
comments and criticisms.
REFERENCES
Brooks, H. K. 1969. Eocarida. In: Moore, R. C., ed. Treatise on invertebrate palaeontology
Part R. Arthropoda 4. Volume 1: R332—-R345. Boulder: Geological Society of America;
Lawrence: University of Kansas.
Broom, R. 1931. On the Pygocephalus-like Crustacea of the South African Dwyka.—Proc.
zool. Soc. Lond. 1931: 571-573.
Du Torr, A. L. 1954. The geology of South Africa. Edinburgh & London: Oliver & Boyd.
Fasre, J. 1967. Un arthropode nouveau des ‘“‘Upper Dwyka shales” Pygaspis ginsburgi,
nov. sp.—Annls Paléont. (Invert.) 53: 121-141.
HAUGHTON, S. H. 1919. A review of the reptilian fauna of the Karroo System of South
Africa.— Trans. geol. Soc. S. Afr. 22: 1-25.
HAUGHTON, S. H. 1969. Geological history of southern Africa. Johannesburg: Geological
Society oi South Africa.
McLAcuLAN, I. R. & ANDERSON, A. 1973. A review of the evidence for marine conditions in
southern Africa during Dwyka times —Palaeont. afr. 15: 37-64,
TAXONOMIC STATUS OF THE PYGOCEPHALOMORPHIC CRUSTACEA 33
Pinto, I. D. 1971. Reconstitucao de Pygaspis Beurlin, 1934 (Crustacea —Pygocephalomorpha).
Sua Posicao sistematica, seu significado e de outros fosseis para o Gondwana.— An. Acad.
brasil. Ciénc. 43: 387-401.
Rocers, A. W. & Du Tort, A. L. 1909. An introduction to the geology of Cape Colony. London,
etc.: Longmans, Green.
SavacE, N. M. 1971. A varvite ichnocoenosis from the Dwyka Series of Natal.—Lethaia
Ae 247: 233.
SECRETAN, S. 1967. Proposition d’une nouvelle compréhension et d’une nouvelle subdivision
des Archaeostraca. — Annls Paléont. (Invert.) 53: 153-188.
Woops, H. 1923. Note on Pygocephalus from the Upper Dwyka shales of Kimberley. —Trans.
geol. Soc. S. Afr. 25: 41-42.
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6. SYSTEMATIC papers must conform with the International code of zoological nomenclature
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BRIAN KENSLEY
TAXONOMIC STATUS OF THE
PYGOCEPHALOMORPHIC CRUSTACEA
FROM THE DWYKA ‘WHITE BAND’
(PERMO-CARBONIFEROUS) OF SOUTH AFRICA
t OLUME 67 PART 4 JULY 1975 ; | ISSN 0303-2515
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Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.—
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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 67 Band
July 1975 Julie
Part 4 Deel
MARINE ISOPODA
FROM THE CONTINENTAL SHELF
OF SOUTH AFRICA
By
BRIAN KENSLEY
Cape Town Kaapstad
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MARINE ISOPODA FROM THE CONTINENTAL SHELF
OF SOUTH AFRICA
By
BRIAN KENSLEY
South African Museum, Cape Town
(With 27 figures and 3 tables)
[MS accepted 15 October 1974]
ABSTRACT
A collection of marine isopods taken from the continental shelf on the Agulhas Bank
in the regions of Still Bay, False Bay, Saldanha Bay, and the southern west coast, is dealt with.
Descriptions and figures of two new genera, viz. Agulanthura and Austroarcturus, as well as
fourteen new species are provided.
CONTENTS
PAGE
FMEFOUUCHON--! Ye is weet 1s oe~ os ° “ar ol3d
SPECIESHISts Vine Behe) ol Oe gs 2 ee 6
Systematic discussion
WalvitGha:. sl ace eo aY os. ot a> a 40
Amthurided.. 6.4. ~« « «= » «. ©
Hlabellifera . =" s fe 2 els TB
Acknowledgements. : 2°. .«. » »« 9
IREICRCHGES Shc ok MS ke 8D
INTRODUCTION
As part of a survey of the fauna of the continental shelf of South Africa, the
Zoology Department of the University of Cape Town carried out a bottom
transect in a line off the coast from Still Bay, Cape. This line extended over the
relatively shallow platform of the Agulhas Bank in a south-easterly direction.
The samples were taken from the university vessel, the 7. B. Davie, in June 1972
and May 1973. The bottom fauna was sampled from depths of 15 to 350 metres
by means of trawls, dredges, and grabs. The isopods collected were submitted
to the South African Museum for identification, together with some samples
from False Bay, Cape, Saldanha Bay, Cape, and the southern west coast,
thought to have a bearing on the material from the Agulhas Bank.
Fifty-four species, of which fourteen are new, are described in this study;
also two new genera.
A discussion of zoogeography, ecology, and distributional patterns for the
area will be carried out by members of the Zoology Department, once all the
animal groups (most of which have proved to be extremely interesting) have
been identified.
The bulk of the collection of isopods is housed in the Zoology Department
35
Ann. S. Afr. Mus. 67 (4), 1975: 35-89, 27 figs, 3 tables.
36 ANNALS OF THE SOUTH AFRICAN MUSEUM
of the University of Cape Town, while all type material is in the collection of the
South African Museum.
In the accompanying figures, all dimensions are in millimetres.
SPECIES LIST
The material dealt with in this paper comes from the following localities:
False Bay (FAL) approximately 34°S., 18°E.
Lambert’s Bay (LBT) 32.04S., 18.20E.
South coast dredge material (SCD)
Still Bay (SST) approximately 35°S., 22°E.
Saldanha Bay (SB) 33.01S., 17.58E.
Southern west coast (WCD)
Lamberts Bay
Saldanha Bay
Cape Town
Still Bay
Cape Agulhas
Fig. 1. Coastline of the south-western Cape, showing collecting localities.
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 37
Suborder VALVIFERA
Family Arcturidae
Antarcturus kladophorus Stebbing .
Arcturella brevipes Barnard
Arcturella corniger (Stebbing) .
Arcturella lobulata Barnard
Arcturina hexagonalis Barnard.
Arcturina scutula sp. nov. .
Arcturina triangularis Barnard
Astacilla bacillus Barnard .
Austroarcturus africanus sp. Nov. .
Austroarcturus foveolatus sp. nov. .
Holidotea unicornis Barnard
Microarcturus laevis sp. nov.
Microarcturus ornatus sp. nov.
Microarcturus quadriconus sp. nov.
Cat. No.
SST.11.D
SST.47.Q
SST.109.Q
SST.11.C
SST.96.R
SST.96.W
FAL.487.D
FAL.496.0
FAL.506.1
FAL.510.U
FAL.654.T
FAL.657.E
FAL.658.T
FAL.674.X
FAL.686.X
FAL.702.R
FAL.706.S
FAL.798.E
FAL.840.H
SST.101.E
FAL.654.S
FAL.658.U
FAL.670.B
SST.61.U
SST.11.C
SST.27.U
FAL.335.D
FAL.428.P
FAL.442.E
FAL.589.E
SB.187.J
SB.270.H
SB.304.U
SCD.243.N
SCD.293.W
SCD.337.P
SCD.339.P
WCD.206.R
WCD.211.K
FAL.760.N
SCD.204.E
SCD.235.C
SCD.272.0
SCD.326.E
SST.1.Y
SST.11.J
SST.101.G
FAL.423.D
FAL.803.K
SST.10.H
SCD.217.R
SST.101.H
200
30
30
200
120
120
62
42
68
68
ie
75
87
66
61
Depth Substrate
(M)
Coarse khaki sand and rock
Coarse sand and shell
Rock
Coarse khaki sand and rock
Rock and shell
Rock and shell
Green sand and shell
Rock
Yellow sand
Yellow sand
Khaki sand and shell
Khaki sand, shell and gravel
Fine khaki sand and shell
Khaki shell and sand
Fine khaki sand
Khaki sand
Khaki sand
Green mud, shell and sand
Sand and rock
Coarse sand and shell
Khaki sand and shell
Fine khaki sand
Coarse khaki sand and shell
Sand and fine shell
Coarse khaki sand and rock
Coarse sand and shell
Coarse shell and sand
Shell
Sand
Coarse sand and shell
Green mud, shell
Coarse shell
Sand, mud, rock
Coarse sand and shell
Mud
Mud
Shell
Shell, sand, green mud
Coarse green sand
Khaki sand
Khaki sand
Khaki sand and shell
Dark mud
Coarse khaki sand
Coarse khaki sand and rock
Coarse sand and shell
Khaki sand and shell
Coarse sand and shell
Coarse khaki sand and rock
Coarse sand and shell
38 ANNALS OF THE SOUTH AFRICAN MUSEUM
Microarcturus similis (Barnard)
Neastacilla tranquilla sp. nov.
Pleuroprion chuni (zur Strassen)
Family Idoteidae
Synidotea hirtipes (Milne Edwards)
Synidotea setifer Barnard .
Suborder ANTHURIDEA
Family Anthuridae
Agulanthura serenasinus sp. nov. .
Apanthura africana Barnard
EXanthurasps = = Jo. &
Exanthura filiformis (Lucas)
Haliophasma cf coronicauda Barnard .
Haliophasma foveolata Barnard
Holoroanthura capensis sp. nov.
Katanthura laevitelson sp. nov.
Leptanthura agulhasensis sp. nov. .
Leptanthura laevigata (Stimpson) .
Cat. No.
SST.40.J
SST.46.C
SST.106.A
SST.11.F
SST.11.E
SST.46.B
SST.91.V
SST.102.N
SST.106.B
SST.101.F
FAL.487.E
FAL.496.L
Depth
(M)
FAL.654.N-R 75
FAL.685.B-D 29
FAL.666.X-Y 26
FAL.763.R-S
SCD.188.P
SCD.329.R
SST.67.S
SST.74.G
SST.5.L
SST.17.E
SST.31.V
SST.76.U
SST.92.B
SST.1.X
SST.17.D
SST.21.R
SST.101.J
SST.37.K
SST.101.C
SST.8.C
SST.19.K
LBT.72.K
WCD.64.P
WCD.109.A
WCD.111.J
WCD.114.U
SST.47.K
FAL.673.J—L
SCD.204.B
SCD.343.Q
SST.27.S
SST.31.U
WCD.77.G
SST.65.M
SST.70.Z
SST.92.A
SST.101.K
SST.114.A
Substrate
Green mud
Green mud
Coarse khaki sand and rock
Coarse khaki sand and rock
Rock
Coarse sand and shell
Green mud
Coarse sand and shell
Green sand and shell
Rock
Khaki sand, shell
White shell and sand
Sand, shell and gravel
Fine green sand
Green mud .
Fine sand and shell
Sand and coral fragments
Coarse khaki sand
Coarse khaki sand and rock -
Coarse khaki sand and shell
Coarse sand
Rock and sand
Rock and coarse khaki sand
Coarse khaki sand and rock
Sand and rock
Coarse sand and shell
Coarse sand and shell
Coarse sand and shell
Coarse khaki sand
Coarse sand and shell
Sand and clay
Green mud
Fine dark green mud
Dark green mud
Dark green mud
Coarse sand and shell
Shell, khaki sand, and gravel
Khaki sand
Sand and shell
Coarse khaki sand and shell
Coarse khaki sand and shell
Green sand and mud
Sand and fine coral fragments
Sand
Rock and sand
Coarse sand and shell
Sand
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 39
Leptanthura urospinosa sp. nov.
Paranthura punctata (Stimpson)
Suborder FLABELLIFERA
Family Cirolanidae
Cirolana borealis Lilljeborg
Cirolana cingulata Barnard :
Cirolana hirtipes Milne Edwards .
Cirolana imposita Barnard .
Cirolana obtusispina sp. nov.
Cirolana pilula Barnard
Cirolana virilis Barnard
Family Sphaeromatidae
Cymodoce alia sp. nov.
Cymodoce cf. umbonata Barnard .
Cymodoce velutina sp. nov.
Cymodocella sp. .
Dynamenella sp.
Family Aegidae
Aega antillensis Schioedte & Meinert
Aega monilis Barnard
Family Corallanidae
Lanocira gardineri Stebbing
Lanocira sp. .
Suborder GNATHIIDEA
Family Gnathiidae
Gnathia africana Barnard .
Gnathia cryptopais Barnard
Gnathia spongicola Barnard
Gnathia sp.
Cat. No.
FAL.442.K
Depth
(M)
39
FAL.654.N-R 75
FAL.666.X-Y 26
FAL.838.Z 5
SCD.310.B 50
SST.1.W 200
SST.19.L 120
SST.17.F 200
SST.21.Q 120
SST.37.M 80
SST.96.S 120
SST.57.A 30
SST.114.B 15
SST.47.R 30
SST.27.T 80
SST.87.H 350
SST.11.L 200
SST.91.W 200
SST.96.V 120
SST.19.N 120
SST.21.U 120
SST.68.Y 20
SST.76.J 15
SST.77.W 10
SST.114.C 15
SST.54.N 30
SST.101.D 80
SST.17.A 200
SST.21.8 120
Sor21T 120
SST.20.Z 120
FAL.700.R-T —
SST.21.W 120
SST.54.P 30
SST.84.A 200
SST.84.B 200
SST.21.V 120
SST.109.P 30
SST.91.Y 200
SST.11.H 200
SST.10.K 200
SST.11.H 200
SST.10.J —
Substrate
Sand
Khaki sand and shell
Sand, shell and gravel
Sand and rock
Coarse sand
Coarse khaki sand
Coarse sand and shell
Coarse khaki sand and rock
Sand and rock
Coarse sand and shell
Rock and sand
Coarse sand and shell
Sand
Coarse sand and shell
Coarse sand and shell
Rock
Coarse khaki sand and rock
Rock
Rock and sand
Coarse sand and shell
Sand and rock
Sand and fine coral fragments
Coarse sand
Coarse sand
Sand
Coarse sand and shell
Coarse sand and shell
Coarse khaki sand and rock
Sand and rock
Sand and rock
Coarse sand and shell
Sand and rock
Coarse sand and shell
Coarse sand
Coarse sand
Sand and rock
Rock
Sand and rock
Coarse khaki sand and rock
Coarse khaki sand and rock
Coarse khaki sand and rock
40 ANNALS OF THE SOUTH AFRICAN MUSEUM
Cat. No. Depth Substrate
(M) 7
Suborder ASELLOTA
Family Munnidae
Mirna spe 2s IS ees es SSE 120 Sand and rock
Family Stenetriidae
Stenetrium crassimanus Barnard . . SST.96.T 120 Rock and sand
Stenetrium dagama Barnard . . . SST.91.X 200 Rock
SST.92.C 200 Rock
SST.11.K 200 Coarse khaki sand and rock
Steneirium Sp: -..6. «2s 2s -e SSS 200 Coarse khaki sand
SST.37.N 80 Coarse sand and shell
SYSTEMATIC DISCUSSION
Suborder VALVIFERA
Family Arcturidae
Barnard (1920: 381) summarized the differences between four families of
the Valvifera. The characters used by Barnard to redefine the family Pseudido-
theidae Ohlin (1901) were partly based on his new genus Holidotea. With the
present material, and especially the two species of the new genus Austroarcturus,
as well as four species of Microarcturus available, a more critical examination of
Holidotea shows that it is not a member of the Pseudidotheidae but is rather a
member of the family Arcturidae. Reasons for the change follow the descriptions
of the species of the new genus Austroarcturus.
Holidotea unicornis Barnard
Figs 2a-j
Holidotea unicornis Barnard, 1920: 382; 1940: 493. Nordenstam, 1933: 113. Nierstrasz,
1941: 262.
Remarks
It was thought useful to include figures of some of the appendages of this
species not given in Barnard’s description, as these are relevant to the discussion
of the family position of the species.
Austroarcturus gen. nov.
Body dorso-ventrally flattened. Eyes dorsal. Lateral margins of head entire.
Peraeonal segment I fused with head, yet distinguishable. Pleon consisting of
two distinct segments plus pleo-telson. Penis single, apically bifid. Peraeopod I
shorter than following peraeopods, strongly setose. Peraeopods II to IV
sparsely setose, more slender than peraeopods V to VII. Exopod of male
pleopod 1 modified; stylet present on endopod of male pleopod 2. Inner ramus
of uropod minute.
Type species of the genus: Austroarcturus foveolatus sp. nov.
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 4]
Fig. 2. Holidotea unicornis Barnard.
a. g, dorsal view; b. 3, lateral view; c. penis; d. maxilliped; e. apex of uropod; f. pleopod 1 3;
g. pleopod 2 3; h. peraeopod I; i. peraeopod III; j. peraeopod VI.
42 ANNALS OF THE SOUTH AFRICAN MUSEUM
Austroarcturus foveolatus sp. nov.
Figs 3a—n
Description of °
Body dorso-ventrally flattened. Dorsal surface finely pitted. Peraeonal
segment I fused with head yet distinguishable. Body widest at peraeonal
segments II and III, segments IV to VII becoming gradually narrower. Pleon
consisting of two free segments plus pleo-telson. Latter with distinct indentation
proximo-laterally. Head bearing mid-dorsal sloping crest between eyes,
continued on posterior portions of segments II to IV, strongest on segments II
and IV.
Antennule about one-third length of antenna, peduncle 3-segmented,
flagellum a single segment bearing several aesthetascs.
Antennal peduncle 5-segmented, 2nd segment triquetral, with prominent
lateral flange; two proximal segments subequal; 4th and 5th segments slender.
elongate; flagellum of two segments, tipped with strong spine.
Mandible bearing tridentate incisor process; lacinia mobilis tridentate with
three penicils at its base, molar process large, bearing numerous short bristles.
Ist maxilla biramous, outer ramus tipped with about 10 spines, inner ramus
bearing three stout plumose setae.
2nd maxilla, outer ramus bilobed, outer lobe with three serrate spines, inner
with two, inner ramus bearing six plumose setae and four simple setae.
Maxilliped with 5-segmented palp, endite distally slightly convex, bearing
numerous bristles plus about seven fringed setae.
Peraeopod I shorter than rest, basal segment equal in length to merus,
ischium, carpus, and half of propodus. Merus with broad dorsal flange;
propodus broadly oval in shape; dactylus stout, somewhat hook-like; two distal
segments bearing numerous fringed setae, those on propodus arranged in
rows.
Peraeopods II to IV slender, with elongate propodi, carpi, and bases;
dactyli slender, curved.
Peraeopods V to VII stout, shorter than earlier peraeopods except peraeo-
pod I, propodus equal in length to carpus and merus together.
Uropods articulating with pleon at about midpoint of outer margin,
basally rounded, distally lanceolate, bearing two rami, outer ramus tiny, inner
ramus reduced to papilla bearing single serrate spine.
Description of 3
Head and peraeon appendages as in female. Rounded crest on head and
dorsal crests of peraeonal segments III and IV relatively stronger than in
female. Penis elongate, single, distally bifid, rounded, bearing numerous fine
bristles.
Pleopod 1, propodus shorter than rami, bearing row of short blunt spines
on outer margin, four hooks on inner margin; outer ramus equal in length to
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 43
Fig. 3. Austroarcturus foveolatus sp. nov.
a. 2, dorsal view; b. 2, lateral view; c. antennule; d. antenna; e. Ist maxilla; f. 2nd maxilla;
g. maxilliped; h. apex of uropod; i. pleopod 1 g; j. pleopod 2 6; k. penis; 1. peraeopod I;
m. peraeopod IT; n. LeAned VII.
44 ANNALS OF THE SOUTH AFRICAN MUSEUM
propodus, fringed with plumose setae; inner ramus apically modified, curved
towards median line, with convoluted tip.
Pleopod 2 bearing elongate stylet on base of inner ramus; stylet with
several rows of fine bristles.
Material
g 3
Holotype SAM-A13540 SST.1.Y 9,0 -—
Allotype SAM-A13540 SST.11.J — I
Paratypes SAM-A13541 SCD272-€ 7,4 4,5
4,9
SCD.204.D 4,9 -—
SCD.217.R a2
Single juvenile specimens from SCD.235.C, SCD.326.F, FAL.760.N.
Austroarcturus africanus sp. nov.
Figs 4a-k
Description of 2
Body dorso-ventrally flattened, integument smooth. Peraeonal segment I
fused with head yet distinguishable. Body widest at peraeonal segment II;
peraeonal segment IV not as wide as III or V. Segments V and VI equal in width,
segment VII shorter and narrower than preceding segments. Pleon consisting of
two free segments plus pleo-telson. Latter with proximo-lateral lobes, strongly
convex, distally bluntly rounded. Head evenly rounded, convex. No dorsal
crests or ridges.
Antennule about one-quarter length of antenna, peduncle 3-segmented,
flagellum of a single segment bearing several aesthetascs.
Antennal peduncle 5-segmented, segments 2 to 5 subequal in length,
2nd triquetral with prominent flattened lateral flange; flagellum of two segments.
Mouthparts as in Austroarcturus foveolatus.
Peraeopods II to IV more slender than peraeopods V to VII, with scattered
setae and numerous fine setules on ventral surface. Peraeopods V to VII stout,
also bearing numerous setae and setules.
Outer ramus of uropod tiny, with terminal fringed spine, inner ramus
reduced to a papilla bearing a single fringed spine.
Description of 3
Similar to female, but epimeres of peraeonal segments V and VI more
rounded, and more obviously extending laterally beyond segment IV. Penis
elongate, single, distally bifid for about cne-third of length, lobes distally
rounded, bearing setules.
Pleopod 1 propodus basally wider than distally, bearing about 18 short
blunt spines on outer margin, four elongate hooks on inner. Outer ramus
elongate-rectangular, fringed with plumose setae. Inner ramus distally modified,
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 45
i
Fig. 4. Austroarcturus africanus sp. nov.
a. 3, dorsal view; b. 3, lateral view; c. antennule; d. antenna; e. penis; f. pleopod 1 6;
g. pleopod 2 g; h. apex of uropod; i. peraeopod I; j. peraeopod II; k. peraeopod VII.
46 ANNALS OF THE SOUTH AFRICAN MUSEUM
bearing row of about 25 stiff plumose setae on outer margin, distally curved and
rounded.
Pleopod 2 bearing slender elongate stylet on base of inner ramus, extending
slightly beyond tips of rami.
Material
- re)
Holotype SAM-A13542 FAL.442.E — 5,4
Allotype SAM-A13542 FAL.442.E 6,0 ovig. —
Paratypes SAM-A13543 FAL.428.P 7,0 ovig. 6,5
5,9 ovig.
Numerous males (3,9-6,2 mm) and females (4,9-7,0 mm) from the following
stations: FAL.589.E., FAL.335.D., SST.102.M., SCD.337.P., SCD.339.P.,
SCD.243.N., SCD.293.W., SB.304.U., SB.187.J., SB.270.H., WCD.206.R.,
WCD.211.K.
Remarks
The present species is obviously closely related to the foregoing Austro-
arcturus foveolatus, but differs in several definite and constant features. In
A. foveolatus peraeonal segments V to VII and the pleo-telson taper gently and
evenly posteriorly; in A. africanus the epimeres of segments V and VI extend
laterally well beyond those of segments IV and VII. The proximo-lateral lobes
at the base of the pleo-telson are more strongly developed in A. africanus, and
consequently the notch formed distally at the fusion to the pleo-telson is wider.
The integument of this latter species is not so obviously granular and pitted as in
A. foveolatus, and when seen in profile, A. africanus lacks the high median crest
of A. foveolatus. In the appendages (excluding the mouthparts) there are subtle
differences but the most striking are in the antennae. In A. foveolatus the 4th
and 5th peduncle segments are very elongate and slender, each being equal in
length to the three basal segments together. A. africanus does not have elongate
segments in the antennal peduncle.
It is the author’s opinion that Holidotea should be placed in the Arcturidae
because the species is more or less intermediate between species of Austroarcturus
and Microarcturus of that family. The reasons are as follows:
A similar differentiation of the peraeopods is apparent in species of the threegenera.
The mouthparts and uropodal rami are similar.
The outer ramus of pleopod 1 in the male, although showing slight differences,
follows the same basic plan in all three genera.
The 2nd pleopods in the males are similar, as are the penes. From this list, the
three genera would seem to be quite closely related.
Barnard (1920) separated the Pseudidotheidae from the Arcturidae
(Astacillidae in his table) by the following features:
1. Body flattened in Pseudidotheidae, cylindrical in Arcturidae.
2. Peraeonal segment IV never elongate in Pseudidotheidae, often elongate in the
Arcturidae.
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 47
3. Peraeopod I prehensile in the Pseudidotheidae, slender and setiferous in the
Arcturidae.
4. Peraeopods II to IV stout (moderately) in Pseudidotheidae, slender and setiferous
in the Arcturidae.
Comparison of species of the three genera make it obvious that these
criteria can be given little strength in the separation of the two families.
Several members of the Arcturidae possess a body to some degree dorso-
ventrally depressed, especially species of Arcturella and Microarcturus. This
flattening is especially noticeable in the females.
Several species of the arcturids do not possess elongate 4th peraeonal
segments, e.g. Arcturella and Microarcturus.
From the species here discussed, and from the figures supplied, it can be
seen that the Ist peraeopods are very similar in structure, as are peraeopods
II to IV, and V to VII. Ohlin (1901), in his description of the family Pseudido-
theidae based on Pseudidothea bonnieri, notes that peraeopods II to VII are
nearly the same in structure and size. His figure of P. bonnieri shows a very
idoteid-like isopod, quite unlike Holidotea unicornis. It would seem that
Holidotea unicornis is more closely related to species of the arcturids than to
species of the Pseudidotheids. It is proposed that Holidotea thus be placed in the
Arcturidae. The criteria used to separate species of the three genera, Holidotea,
Austroarcturus, and Microarcturus, are summarized in Table 1.
Table 1
Holidotea Barnard Austroarcturus Kensley Microarcturus
Nordenstam
Lateral Entire Entire Incised
margins of
head
Eyes Dorsal Dorsal Dorso-lateral
Peraeonal Distinguishable, fused Distinguishable, fused Indistinct, completely
segment I to head to head fused to head
Pleon No distinct segments 2 distinct segments 2-3 distinct segments
anterior to pleo-telson anterior to pleo-telson anterior to pleo-telson
Inner Less than half length More than half length More than half length
ramus of of outer ramus, distally of outer ramus, distally of outer ramus, distally
pleopod 13 acute truncate-rounded truncate-rounded
Microarcturus similis (Barnard)
Figs 5a—b
Antarcturus similis Barnard, 1925: 395; 1940: 508.
Microarcturus similis: Nordenstam, 1933: 128.
Material
SST.106.A-C I1¢
SST.40.J 13
SST.46.C 19
48 ANNALS OF THE SOUTH AFRICAN MUSEUM
Remarks
For comparison with the new species described below, a male and female
of this species are figured in dorsal view.
Fig. 5. Microarcturus similis (Barnard).
a. 9, dorsal view; b. 3, dorsal view.
Microarcturus laevis sp. nov.
Figs 6a-1
Description of °
Integument smooth. Body dorso-ventrally flattened, head and peraeonal
segment I fused, but with lateral margins free. Body widest at peraeonal
segments II and III; peraeonal segments V to VII and pleo-telson narrow, all
of similar width. Pleon consisting of two free segments, with third segment
indicated on pleo-telson. Head bearing two large dorso-lateral eyes separated
by convex semicircular portion, antero-lateral corners slightly produced.
Epistome produced into blunt tapering process, tip just visible in dorsal view.
All epimeres distinct, those of peraeonal segments II and III evenly convex in
dorsal view. Peraeonal segments all dorsally convex, segment III bearing two
broadly convex submedian dorsal bulges. Rounded boss at base of pleo-telson.
Antennule slightly less than one-third length of antenna, consisting of
3-segmented peduncle and one flagellar segment. Basal segment broad, bearing
fine setae, flagellum equal in length to distal two peduncle segments.
Antennal peduncle 4-segmented, 3rd segment longest, flagellum
2-segmented, tipped with single spine.
Mandible with strongly chitinised quadridentate incisor process, lacinia
mobilis tridentate with three penicils at its base, large broad molar process
covered with fine bristles.
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 49
Fig. 6. Microarcturus laevis sp. nov.
a. 2, dorsal view;. b. 9, lateral view; c. antennule; d. antenna; e. peraeopod I; f. peraeopod IT;
g. peraeopod VII; h. maxilliped; i. 2nd maxilla; j. 1st maxilla; k. mandible; 1. apex of
uropod.
50 ANNALS OF THE SOUTH AFRICAN MUSEUM
Inner ramus of Ist maxilla bearing three long plumose setae, outer ramus
tipped with about 10 spines.
Inner ramus of 2nd maxilla bearing seven plumose setae and four slender
spines distally; inner lobe of outer ramus with two distal serrate spines, outer
lobe with three.
Maxilliped palp 5-segmented, 3rd segment broadest; endite broad, bearing
numerous simple setae plus several short fringed setae on inner distal angle.
Peraeopod I shorter than following peraeopods; basis equal in length to
ischium, merus, and carpus together; propodus bearing three rows of setae on
outer distal face; dactylus armed with serrate setae.
Peraeopods II to IV slender, dactyli, propodi, and carpi elongate, unguis of
dactylus blunt, striated.
Uropods proximally evenly rounded; outer ramus minute, inner ramus tiny,
reduced to papilla bearing a single serrate spine.
Material
Holotype SAM-A12544 1 ovigerous 2 6.4mm FAL.803.K
Paratype SAM-A13545 1 ovigerous 2 5.2mm FAL.423.D
Microarcturus ornatus sp. nov.
Figs 7a—o
Description of 2
Body with large tubercles, widest at peraeonal segment III. Head with
antero-lateral corners acute; broad transverse furrow separates rounded
posterior ridge from rest of head. Eyes large, lateral. Peraeonal segment I with
epimeres ventrally directed, tridentate. Epimeres of peraeonal segments IH to IV
expanded laterally. Peraeonal segments with two medio-lateral tubercles,
largest on segments I to IV. Pleon consisting of one indistinct and two distinct
segments plus pleo-telson. Latter terminally acute, with strong lateral teeth, and
bearing scattered tubercles. Antennule reaching to midpoint of 2nd antennal
peduncle segment. Peduncle 3-segmented, basal segment broadest, 3rd segment
one-quarter length of flagellum.
Antennal peduncle 4-segmented, basal segment about half length of
2nd segment, with antero-lateral corner acute, 2nd segment with distal spine,
3rd and 4th segments slender, subequal in length, flagellum 2-segmented with
slender terminal spine.
Mandible with tridentate incisor process; lacinia mobilis tridentate with
three penicils at base, and broad setose molar process with four slender setae at
its base.
Ist maxilla with three setae on inner ramus, several spines on outer ramus.
2nd maxilla inner ramus bearing five simple spines and five stout setae,
outer lobe of outer ramus bearing three elongate spines, inner lobe with two
fringed setae.
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 51
Fig. 7. Microarcturus ornatus sp. nov.
a. 6, dorsal view; b. 9, dorsal view; c. 9, lateral view; d. antennule; e. antenna;
f. peraeopod 1; g. peraeopod III 2; h. peraeopod VII 9; i. penis; j. pleopod 1 3; k. pleopod
2; 1. peraeonal segment IV, 3, ventral view; m. mandible; n. 2nd maxilla; o. maxilliped.
By? ANNALS OF THE SOUTH AFRICAN MUSEUM
Maxilliped palp 5-segmented, endite relatively narrow, with three setae at
inner distal angle.
Peraeopod I shorter than following peraeopods; propodus twice length of
dactylus, basis equal in length to ischium and merus together.
Peraeopods II to IV increasing in length posteriorly, five distal segments
relatively slender, bearing elongate setae; basis stout, bearing two strong
spinose processes on dorsal margin in peraeopods II and III, three processes on
peraeopod IV; terminal segment with strong serrate spine apically.
Peraeopods V to VII with all segments robust, propodus bearing small
disto-ventral spine, basis with strong spinose process at about midpoint of outer
face.
Description of 3
Body elongate, parallel-sided, hardly depressed, bearing numerous tubercles.
Peraeonal segments I to IV with more prominent submedian dorsal tubercles.
Peraeonal segment IV bearing ventrally a triangular process between bases of
peraeopods, plus two submedian oval flattened processes distally. Peraeopods II
to IV lacking processes on bases.
Pleopod 1 with outer ramus broader and longer than inner, bearing
oblique curved groove on anterior face, ending on a rounded distal prominence.
Stylet on inner ramus of pleopod 2 sabre-like. Penis basally broad, distally bifid,
rami not diverging. |
Material
Holotype SAM-A13546 SST.10.H 2 ovig. 6,4 mm
Allotype SAM-A13546 SST.10.H 4 5,0 mm
Paratypes SAM-A13547 SST.10.H 233 5,0 mm
4,1 mm
Microarcturus quadriconus sp. nov.
Figs 8a-i
Description of 2
Peraeon segments II and III broadest. Head and peraeonal segment I fused,
with shallow furrow indicating line of fusion. Anterior margin of head concave,
eyes dorso-lateral, two large submedian conical tubercles in line with eyes, each
flanked by a tiny anterior and posterior tubercle. Submedian tubercles of
peraeonal segment I tiny, two lateral rounded tubercles slightly larger. Peraeonal
segments II to IV each with four large conical tubercles. Peraeonal segments V
to VII each with two tubercles. Pleon having two segments anterior to pleo-
telson, latter with large rounded boss mid-dorsally at base. Pleo-telson
pentagonal, distally acute. Antennular peduncle 4-segmented, basal segment
equal in length to 2nd and 3rd segments together; flagellum 2-segmented.
Mouthparts typical of the genus.
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 53
Fig. 8. Microarcturus quadriconus sp. nov.
a. 2, dorsal view; b. 3, dorsal view; c. penis; d. peraeopod I; e. peraeopod II; f. peraeopod
VII; g. pleopod 1 g; h. pleopod 2 g; i. maxilliped.
Peraeopod I shorter than following peraeopods, three distal segments
bearing numerous serrate spines.
Peraeopods II to IV more slender than following ones, bearing few scattered
elongate setae.
Peraeopods V to VII somewhat shorter than preceding ones, distal segments
stouter.
54 ANNALS OF THE SOUTH AFRICAN MUSEUM
Uropod with outer ramus tiny, inner ramus reduced to papilla with single
serrate spine.
Description of 3
Body elongate, peraeonal segments II and III only slightly wider than rest
of body. Sculpture as in female.
Pleopod 1 rami subequal in length, outer ramus with oblique groove
running almost entire length, ending at slight bulge at outer distal angle.
Pleopod 2 inner ramus distally truncate, equal in length to stylet, latter
sabre-shaped. Penis single, distally bifid, distal rounded lobes not diverging.
Pleo-telson not as obviously pentagonal as in female.
Material
Holotype SAM-A13548 SST.101.C-—K 2 ovig. 5,2 mm
Allotype SAM-A13548 So 4,0 mm
Paratypes SAM-A13549 2 ovig. 5,6 mm
4,8 mm
SCD.217.R Qovig. 4,9 mm
Remarks
The genus Microarcturus was defined by Nordenstam (1933: 128). The
following characters which set it apart from species of Arcturus, are present in
the foregoing three species, as well as in M. similis:
Lateral margins of head incised; eyes dorso-lateral; peraeonal segment I fused
with head, often indistinct; pleon possessing two free segments plus one indicated on
the pleo-telson; antennae shorter than body; antennal flagellum 2-segmented
(2—4-segmented); pleo-telson never longer than last four peraeonal segments together.
M. ornatus resembles M. rugosus Nordenstam in the spination of the
peraeopods II to IV, but lacks the numerous elongate acute tubercles and
spinose epimeres of the Antarctic species. Apart from this similarity, the three
species described here are rather distinctive, and most closely resemble M. similis.
Table 2 summarizes the main differences between the four species.
Table 2
M. similis M. ornatus M. quadriconus M. laevis
Epimeres of peraeonal Angular Rounded Angular Rounded
segments IT & III 2
Dorsal integument Granular- Granular- Non-granular Smooth
tuberculate tuberculate tuberculate
Bases of peraeopods Non-spinose Spinose Non-spinose Non-spinose
II to IV 2
Pleo-telson Rounded boss No rounded Rounded boss Rounded boss
at base boss at base at base present at
present or base
absent
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 55
Genus Arcturina Koehler, 1911
Barnard (1957) mentions the error in Koehler’s description of the male of
Arcturina rhomboidalis, where the figured second pleopod is labelled and
referred to as the first pleopod. Barnard’s material of A. hexagonalis consisted
of a female and an immature male. Many males are now available, thus pleopod
2 of mature males can be recorded. Arcturina rhomboidalis and the three species
included here are compared in table 3.
Arcturina hexagonalis Barnard
Figs 9a—k
Arcturina hexagonalis Baruard, 1925: 400; 1957: 6.
Description of 3
Body slender, geniculate. Peraeonal segment I fused with head but dis-
tinguishable. Peraeonal segments I to III subequal in length, [Vth segment
subcylindrical, bearing two submedian somewhat indistinct bands of pile-like
short hairs. Peraeonal segment V slightly longer than following segments. Pleon
consisting of three fused segments plus pleo-telson. Fused segments subequal,
short, each bearing two dorsal clumps of fine hairs. Antennae, antennules, and
mouthparts as in female.
Peraeopod I within lateral wall of buccal cavity. Peraeopods II to IV
directed anteriorly; peraeopods V to VII stout, considerably longer than
anterior peraeopods.
Penial rami fused for half of length, tips hardly expanded.
Exopod of pleopod | with indentation on outer margin, bearing numerous
fine simple setae plus three stout plumose setae. Exopod slightly longer than
endopod.
Pleopod 2 bearing slender stylet on inner margin of endopod; apex of stylet
with one short and two elongate spines extending well beyond endopod apex.
Previous records
Off Cape St Blaize, 75 metres.
Material
45 3d, 33 99, False Bay.
Arcturina triangularis Barnard
Figs 10a—h
Arcturina triangularis Barnard, 1957: 4, fig. 3.
Description of 3
Body slender, peraeonal segment I distinguishable, fused with head.
Ventro-lateral margin of head slightly scalloped. Peraeonal segment IV equal in
56 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 9. Arcturina hexagonalis Barnard.
a. 2, dorsal view; b. 3, dorsal view; c. antennule; d. apex of uropod; e. penis; f. maxilla;
g. peraeopod I; h. pleopod 1 3; i. pleopod 2 3; j. peraeopod III; k. peraeopod VI.
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA a7
Fig. 10. Arcturina triangularis Barnard.
a. lateral view; b. pleopod 1 3g; c. pleopod2 3; d. penis; e. peraeopod II 3;
f. peraeopod III 3; g. peraeopod IV g; h. peraeopod VI ¢.
length to head plus anterior three peraeonal segments. Ventral margin of
peraeonal segment IV very slightly convex. Head and anterior four peraeonal
segments bearing two submedian dorsal ridges which diverge posteriorly.
Submedian ridges again visible on peraeonal segments VI and VII and on
anterior portion of pleon. Peraeonal segments V to VII having strong lateral
ridge. Mouthparts as in female. Peraeopod I hidden by lateral border of buccal
cavity. Peraeopod II with reduced dactylus; propodus and carpus inflated and
broad. Peraeopods III and IV more slender. Peraeopods V to VII stout, heavily
setose. Pleopod 1 with exopod having indentation at proximal third, with three
stout plumose setae, and numerous fine hairs. Pleopod 2 endopod bearing
slender stylet on inner margin; stylet apically bearing one short and two long
slender spines.
58 ANNALS OF THE SOUTH AFRICAN MUSEUM
Previous records
Mossel Bay, Cape, 9 metres.
Material
8 gg, 11 29. SST.61.U.
Arcturina scutula sp. nov.
Figs 1la-i, 12a—n
Description of °
Head and anterior four peraeonal segments together forming a lozenge-
shaped structure. Head with well-developed antero-lateral lobes. Peraeonal
segment I fused with head yet distinguishable. Two submedian dorsal ridges
extend from anterior margin of head to posterior margin of peraeonal seg-
ment IV. Peraeonal segments V to VII bearing strong lateral ridges, Vth segment
slightly larger than following segments. Pleon consisting of three indistinct
segments fused to pentagonal pleo-telson.
Antennule 4-segmented, basal segment broad, 2nd ae 3rd segments
together equal in length to flagellum; latter bearing single terminal aesthetasc.
Antenna having 5-segmented peduncle and 3-segmented flagellum; first
three peduncle segments together equal in length to 4th segment, latter equal to
Sth.
Mandible consisting of tridentate strongly chitinised incisor process, smaller
tridentate lacinia mobilis and two penicils at its base, and molar process bearing ©
many close-set short bristles.
Ist maxilla biramous, outer ramus tipped with about seven or eight simple
spines, inner ramus bearing three terminal plumose setae.
2nd maxilla biramous, inner ramus tipped with six simple spines, outer
ramus bilobed, each lobe bearing two elongate plumose setae.
Maxilliped with single coupling hook on inner margin of endite, few
scattered fringed setae near upper margin; palp 5-segmented, 3rd segment
longest, broadly oval, terminal segment about one-third length of penultimate
segment.
Peraeopod I within lateral border of buccal cavity, propodus bearing
numerous elongate serrate spines; dactylus bearing two setae and single strong
curved spine.
Peraeopods II and III subequal in length, dactylus reduced, bearing three
serrate spines, propodus broad.
Peraeopod IV slightly shorter than III, dactylus reduced, bearing two
serrate spines, propodus more elongate than in previous peraeopods.
Peraeopods II to IV with well-developed oostegites, that of the IVth
largest.
Peraeopods V to VII becoming successively slightly smaller.
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 59
Fig. 11. Arcturina scutula sp. nov.
a. 2, dorsal view; b. g, dorsal view; c. 9, lateral view; d. 3, lateral view;
e. pleopod 1 3; f. pleopod2 3; g. penis; h. peraeopod II g; i. peraeopod VI.
Uropods anteriorly rounded, distally tapering, bearing strong longitudinal
carina near median margin, outer ramus small, inner ramus about half length
and one-quarter breadth of outer, tipped with single simple seta.
Description of 3
Body slender. Head and anterior three peraeonal segments together
somewhat shorter than peraeonal segment IV. Head plus anterior four peraeonal
segments longer than peraeonal segments V to VII plus pleo-telson. Head with
60 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 12. Arcturina scutula sp. nov.
a. antenna; b. antennule; c. 2nd maxilla; d. 1st maxilla; e. mandible; f. maxilliped;
g. oostegite of peraeonal segment IV; h. peraeopod I 9; i. peraeopod II 9; j. peraeopod III 9;
k. peraeopod IV 92; 1. perzeopod V 2; m. uropod; n. apex of uropod.
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 61
well-developed antero-lateral lobes. Peraeonal segment I forming acute lateral
projection at level of eyes. Two indistinct submedian ridges on Ist to IlIrd
peraeonal segments. Peraeonal segment IV with two strong submedian ridges
dorsally, slightly arched in lateral view, bearing an antero-ventral rectangular
projection.
Antennae, antennules, mouthparts, and anterior four pairs of peraeopods
similar to female. Peraeopods V to VII decreasing slightly in size posteriorly,
basis with membranous ventral flange; propodus about equal in length to carpus
plus merus; all segments bearing simple setae.
Penis consisting of two slender elongate rami fused basally for about
one-quarter of their length.
Pleopod 1 endopod elongate, apically truncate, exopod with indentation on
outer margin flanked by three stout setae.
Pleopod 2 endopod bearing slender stylet on inner margin; stylet distally
consisting of one short and two elongate and more slender spines, reaching
beyond apex of endopod.
Material
Holotype SAM-A13537 False Bay 3 4,5 mm
Allotype SAM-A13537 False Bay 2 3,5 mm
Paratypes SAM-A13538 False Bay 435 4,3-5,1 mm
Paratypes SAM-A13539 False Bay 329 3,5-3,8 mm
Numerous males and females from False Bay; also 4 3g, 5 99, Algoa Bay.
Remarks
Three species of Arcturina have been described, viz. A. rhomboidalis
Koehler, from Morocco and Mauritania, A. hexagonalis Barnard from Cape
St Blaize, and A. triangularis Barnard, from Mossel Bay. Table 3 summarizes
the differences between these three species and the new species A. scutula.
Table 3
3 A. rhomboidalis A. hexagonalis A. triangularis A. scutula
Dorsal More or less Only on peraeonal More or less More or less
ridges parallel extending segment IV parallel extending parallel extending
on to anterior seg- on to head and on to head and
ments and head anterior segments anterior segments
Peraeonal Dorsally straight Dorsally straight Slightly arched Slightly arched
segment IV in lateral view in lateral view dorsally in lateral dorsally in lateral
view view
No antero-ventral No antero-ventral No antero-ventral Truncate antero-
projection projection projection ventral projection
Greatest height 4 Greatest height 4 Greatest height 4 Greatest height 4
length length length length
Total 6-7 mm Average for 10 Average for 3 Average for 10
length specimens: 4,0mm specimens:3,7mm _ specimens: 3,7 mm
62
ANNALS OF THE SOUTH AFRICAN MUSEUM
Table 3 (continued)
fe) A. rhomboidalis A. hexagonalis A. triangularis A. scutula
Total 4 mm Average for 10 Average for 7 Average for 10
length specimens: 5,5 mm specimens: 3,6mm specimens: 3,7mm
Dorsal Separate, parallel, Anteriorly diver- Extending on to Separate, more or
ridges extending on to gent absent from head converging less parallel,
head head and anterior on 3rd and extending on to .
3 segments anterior of 4th head
segment
Peraeonal Dorsally slightly Straight in lateral Smoothly curved Strongly convex in
in lateral view;
lozenge-shaped,
lateral view;
lozenge-shaped,
segmentIV curved; view;
lozenge-shaped widest anteriorly,
(with anterior 3 anterior segments margins of margins of
segments in dorsal rounded anterior segments anterior segments
view) not as rounded as_ not rounded
in hexagonalis
Antennule 2nd & 3rd seg- No serrate seg- No serrate seg- No serrate seg-
ments serrate ments ments ments
Uropods Non-carinate Non-carinate Carinate Strongly carinate
Neastacilla tranquilla sp. nov.
Figs 13a—-d
Description of 2
Head with ventral margins somewhat expanded; antero-dorsal corner also
expanded anterior to dorso-lateral eye; two low rounded submedian dorsal —
bulges present. Peraeonal segment I fused with head, line of fusion marked by a
narrow groove, and with lateral suture visible between head and Ist segment.
Peraeonal segments I to III subequal, [Vth segment about 34 times length of
three anterior segments together, tapering posteriorly in dorsal view. Peraeonal
segment V slightly longer than following segments, segments V to VII each with
three small lateral tubercles. Pleon consisting of three fused segments plus pleo-
telson, latter tapering to acute tip, sides slightly concave. Antennular flagellum
of a single segment, equal in length to two distal peduncular segments together.
Antennal peduncle 5-segmented, 4th segment longest; flagellum
3-segmented, bearing about eight ventral spines. Three pairs of oostegites
present.
Material
Holotype SAM-A13615 SST.11.F. 2 ovig. 6,3 mm
Paratype SAM-A13616 SST.11.F. 2 ovig. 6,3 mm
Remarks
Nordenstam (1933) defined the genus Neastacilla and mentioned some of
the differences from Astacilla. The present material agrees with Neastacilla in
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 63
Fig. 13. Neastacilla tranquilla sp. nov.
a. 2, dorsal view; b. 9, lateral view; c. apex of uropod; d. maxilliped.
having peraeonal segment I fused with the head (the fusion marked by a shallow
groove), the pleon consisting of three fused segments plus pleo-telson, and the
inner ramus of the uropod bearing a long apical seta. The material does,
however, show characteristics of species of Astacilla. These include the presence
of a short lateral suture between the head and peraeonal segment I, and the
lateral portions of the head and peraeonal segment I somewhat expanded.
Peraeopod I however, does possess an unexpanded unguis on the dactylus. It
would seem that Nordenstam’s remark (1933: 119) concerning the superfluity of
Neastacilla in view of species intermediate between Astacilla and Neastacilla
may well be accurate.
N. tranquilla differs from the often recorded N. bacillus in several features,
especially in the shape of the eyes, in being a relatively less slender species, and in
possessing a ‘shoulder’ on the antero-lateral corners of peraeonal segment IV
These shoulders are lacking in N. bacillus.
N. mediterranea, the other species recorded from South Africa, has a
granulate integument and a spinose head, while N. tranquilla is quite smooth
and non-spinose.
64 ANNALS OF THE SOUTH AFRICAN MUSEUM
Suborder ANTHURIDEA
Family Anthuridae
Leptanthura agulhasensis sp. nov.
Figs 14a—k, 15a—d
Description of &
Head about half length of peraeonal segment I. Peraeonal segments I to VI
subequal, segment VII two-thirds length of VIth. Pleonal segments free and
distinct, pleon longer than peraeonal segment VII. Dorso-lateral keels only
obvious on anterior peraeonal segments when seen in lateral view. Eyes absent.
Antennular peduncle 3-segmented, distal segment bearing several setae;
flagellum very short, 4(5)-segmented.
Antennal peduncle 5-segmented, flagellum 4-segmented.
Mouthparts modified for piercing and sucking.
Mandible typical of the genus, elongate-acute; palp 3-segmented, broad
middle segment three times length of basal segment, terminal segment short and
narrow, bearing two terminal serrate spines.
Maxilla slender, elongate, distally serrate on inner margin, bearing three
barbs on outer membranous margin.
Maxilliped elongate, basal segment at least four times longer than broad;
followed by two (?three) distal segments bearing setae.
Peraeopod I with palm of propodus straight, with no thumb at base, with a
row of seven short stout fringed setae flanked by simple elongate setae; carpus ~
triangular, with three distal fringed setae, 2nd and 3rd segments subequal in
length, 2nd segment wider.
Peraeopods II and III similar to I but becoming progressively more
elongate.
Peraeopods IV to VII with dactyli slightly shorter than propodi, bearing
10-12 short setae on ventral margin; ventral margin of propodus with three
spines; carpus triangular, small, underriding propodus, with two spines.
Exopod of pleopod 1 broadly oval, endopod half width of exopod, both
rami fringed with elongate plumose setae.
Uropod exopod lanceolate, shorter than basis; latter with strong dorsal
ridge, triangular in cross-section; endopod tapering to narrowly rounded apex
extending beyond telsonic apex.
Telson with proximal two-thirds parallel-sided, distal third tapering to
acute apex bearing a few fine setae. Single Siatoeyst pore opening dorsally near
base. Four pairs of oostegites present.
Description of 3
Peraeopod I differs from female in having numerous setae on the palm,
and in lacking a row of short fringed setae. Propodus with short blunt proximal
projection.
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 65
Fig. 14. Leptanthura agulhasensis sp. nov. 9.
a. 9, dorsal view; b. maxilliped; c. maxilla; d. mandible; e. antennule; f. antenna;
g. peraeopod I; h. peraeopod VII; i. uropod; j. pleopod 1; k. telson.
66 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 15. Leptanthura agulhasensis sp. nov. é.
a. antenna; b. pleopod 2; c. peraeopod II; d. peraeopod I.
Peraeopod II more slender than peraeopod I, palm of propodus concave; —
blunt proximal projection of propodus outflanked by more elongate projection
of carpus.
Antenna with brush-like flagellum of about 12 segments.
Pleopod 2 endopod with stylus not quite reaching apex of rami, apically
rounded.
Material
Holotype SAM-A13550 FAL.673.J-—L 6 9,0 mm
Allotype SAM-A13551 SST.27.S 2 8,0 mm
Paratypes SAM-A13617 FAL.673.J—L 6d 9,1 mm
9,0 mm
Paratypes SAM-A13618 SCD.343.Q ele) 7,9 mm
6,5 mm
SCD.204.B Q 6,8 mm
WCD.77.G Q 7,9 mm
SST.31.U Q 8,0 mm
Remarks
The present species would seem to be most closely related to L. tenuis (Sars)
recorded from the North Atlantic. The two species agree in the structure of the
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 67
antennae, antennules, mouthparts, telson, and pleopods. In the structure of the
peraeopods, however, the two species do differ. The palm of peraeopod I (as
figured by Sars 1897, pl. 20; and Schultz 1969, fig. 129) of L. tenuis is concave,
with a marked thumb, whereas the present species has a straight palm and no
thumb. Peraeopods IV to VII differ in spination and setation. The uropods are
also different. The present species has a marked dorsal ridge on the basis which
is triangular in cross-section. This feature is not present in L. tenuis. The uropod
exopod of the latter species is broader and proportionally longer than in
L. agulhasensis.
Leptanthura urospinosa sp. nov.
Figs 16a—k
Description of °
Head about half length of peraeonal segment I. Peraeonal segments
I to VI subequal in length. Peraeonal segment VII two-thirds length of VIth.
Pleon equal in length to peraeonal segment VII. Pleonal segments free and
distinct, 5th segment longer than preceding segments, 6th semicircular. Eyes
absent.
Antennular peduncle 3-segmented, distal segment bearing several setae,
flagellum 6-segmented.
Antennal peduncle 5-segmented, second segment longest, flagellum
3-segmented. Mouthparts modified for piercing and sucking.
Mandibular palp 3-segmented, median segment about twice longer than
broad, distal segment bearing two serrate spines.
Maxilla slender, inner margin serrate, outer membranous margin bearing
three separate barbs.
Maxilliped slender, elongate, basal segment extended distally on inner
margin followed by two setae-bearing segments.
Peraeopod I with palm of propodus straight, bearing six stout serrate setae
plus several simple setae; carpus triangular, bearing three stout sensory
setae.
Peraeopod VII with propodus carrying four sensory setae on ventral
margin; carpus underriding propodus, triangular, carrying two sensory setae;
unguis of dactylus very short and blunt.
Pleopod 1 exopod broadly oval, operculate.
Uropod exopod broadly leaf-shaped, inner margin proximally with a right-
angled bend marked by short spine, inner margin dentate, slightly longer than
basis, not reaching telsonic apex; basis with strong dorsal ridge, triangular in
cross-section; endopod triangular, extending beyond telsonic apex.
Telson tapering gently, but distal quarter tapering more acutely; single
Statocyst situated near base.
68 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 16. Leptanthura urospinosa sp. nov.
a. 2, dorsal view; b. antennule; c. antenna; d. mandible; e. maxilla; f. telson; g. uropodal
exopod; h. maxilliped; i. peraeopod VII; j. peraeopod I; k. uropodal basis and endopod.
SS
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 69
Material
Holotype SAM-A13619 FAL.666.X-Y 10,5 mm
2
Paratype SAM-A13620 FAL.838.Z 2 13,3 mm
FAL.654.N-R 2 ovig. 8,8 mm
2 9,6 mm
FAL.442.K 3) Qe 8,5 mm
6,4 mm
6,0 mm
SST.1.W 2 5,4 mm
SST.19.L Q 8,0 mm
SCD.310.B 2 5,2 mm
Remarks
In several respects, L. urospinosa resembles the foregoing species,
L. agulhasensis. These similarities include the mandible, maxilla, antennae,
peraeopods I to VII, and the telson. The major difference is to be seen in the
uropod exopods. In L. agulhasensis the exopod is a short lanceolate structure,
while in L. urospinosa, this is a broadly oval structure, dentate on the inner
margin, with the exopod of each side almost touching basally. The maxilliped of
L. agulhasensis appears to have at least one segment more than in L. urospinosa,
while the antennular flagellum of the latter species consists of six segments, of
four to five segments in the former. The shape of the uropod exopod (except for
its dentate inner margin) is similar to that of L. tenuis.
Katanthura laevitelson sp. nov.
Figs 17a—k
Description of °
Head and first six peraeonal segments of equal length, peraeonal segment VII
very short; segments IV to VI with shallow transverse furrow anteriorly. Pleon
slightly shorter than peraeonal segment VI. Pleonal segments distinct. Dorsal
surface of pleon and peraeon bearing irregular brown reticulate pattern. Frontal
margin of head with tiny rostral point; large well-developed oval eyes on
antero-lateral corner.
Antennule shorter than antenna, basal segment twice length of 2nd segment,
flagellum of 11 segments.
Antennal peduncle 5-segmented, 2nd segment widest and longest, flagellum
of 20 segments.
Mouthparts adapted for piercing and sucking.
Mandibular palp 3-segmented, basal segment short, 2nd segment equal in
length but narrower than 3rd, bearing single simple setae; 3rd segment bearing
a single elongate fringed seta distally, plus row of 17 or 18 spines on outer
margin. Mandible with piercing portion tapering, with single strong spine-like
process, ensheathed by an apparently membranous structure.
70 ANNALS OF THE SOUTH AFRICAN MUSEUM
A ee
Fig. 17. Katanthura laevitelson sp. nov.
a. 9, dorsal view; b. antenna; c. antennule; d. pleopod 1; e. mandible; f. uropod;
g. maxilla; h. maxilliped; i. peraeopod I; j. peraeopod II; k. peraeopod VI.
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 71
Maxilla slender, harpoon-like, distally serrated on one margin, and with
fine groove running length of the appendage, widening proximally.
Maxilliped of three segments; proximal segment slightly longer than two
distal segments, with slender lobe-like extension on medio-distal angle; terminal
segment tapering to narrowly-rounded apex with cluster of about 13 slender
setae.
Peraeopod I strong, subchelate, dactylus strongly curved, with no obvious
unguis; propodus proximally broader than distally, palm only slightly curved,
with row of more or less alternating long and short fringed spines, and scattered
long setae; palm flanked by single row of fringed spines; base of palm with
rounded lobe. Carpus short, triangular, merus with broadly rounded dorsal
area; ischium and basis subequal, more elongate.
Peraeopod II with propodus not as strong or as wide as in peraeopod I;
dactylus curved, with short unguis; propodus with row of seven spines on inner
margin, two distal spines tripartite, proximal five spines having sensory tip.
Peraeopod III similar to II.
Peraeopods IV to VI with dactylus having distinct small unguis, propodus
with five or six stout spines on inner margin, carpus with three spines on ventral
margin. Peraeopod VII absent.
Pleopod 1 not operculiform, outer ramus elongate, distally broadly
rounded, about three times width of inner ramus, with several distal plumose
setae; inner ramus narrow, not curved.
Pleopod 2 with outer ramus only slightly longer than inner.
Uropod with exopod longer than basis, slender lanceolate, apically
narrowly rounded, with numerous setae; endopod twice longer than wide,
apically rounded with numerous setae, extending beyond telsonic apex.
Telson elongate, tapering gently to point, bearing distal setae, large median
proximal statocyst, but no sculpture.
Material
Holotype SAM-A13552 SST.47.K g 6,4 mm
Remarks
The genus Katanthura was defined by Nierstrasz (1941) for K. barnardi
from the Solar Straits. It has the following characteristics, recorded by Barnard
(1925):
Mouthparts modified for piercing and sucking.
No statocyst in telson.
Carpus of peraeopods IV to VI not underriding propodus.
Maxilliped 4-segmented.
Peraeona! segment VII very short, lacking peraeopods.
The male is not known. Nierstrasz based his description on a single female,
as is done in the present case. The specimen described here is so well preserved
(P2 ANNALS OF THE SOUTH AFRICAN MUSEUM
and so markedly different from the female of K. barnardi that it is described as
a new species.
The differences between the two species of Katanthura are given in the
following table.
K. barnardi K. laevitelson
Mandibular palp Terminal segment with 2 rows of One row of spines, one elongate
spines, no elongate spine spine
Maxilla Possessing free lancets No free lancets
Antennule Flagellum of 14 segments Flagellum of 11 segments
Antenna Flagellum of 22 segments Flagellum of 20 segments
Peraeopod VI Propodus with 3 (?) spines Propodus with 6 ventral spines
Pleopod 1 Endopod distally curved Endopod not curved
Telson Bearing 6 weak ridges Unsculptured
Statocyst Absent Present
Total length 14 mm 6,4 mm
Uropod Relatively slender Not as slender as K. barnardi
Agulanthura gen. nov.
Diagnosis
Mouthparts normal, not modified for piercing and sucking. Maxilliped
5-segmented. Third segment cof mandibular palp shorter than Ist or 2nd. Eyes
present. Unguis of peraeopod I long. Peraeopod II smaller than I. Carpus of
peraeopods IV to VII not underriding propodus, but distally expanded, distal
margin straight. Pleopod 1 operculiform. Pleonal sutures indistinct in male and .
female. Stylet on pleopod 2 of male slender, acute. Exopod of uropod folding
over telson. Latter lanceolate, dorsally convex, with two statocysts at base.
Type species of the genus: Agulanthura serenasinus.
Agulanthura serenasinus sp. nov.
Figs 18a—o
Description of 3
Body elongate, head half length of peraeonal segment I. Peraeonal segments
subequal in length, each segment about twice as long as wide. Pleonal segments
indistinct, apparently fused, except segment 6 which is free. Segmental part of
pleon equal in length to telson.
Antennule shorter than antenna, three basal peduncular segments short,
flagellum consisting of 10 segments each bearing numerous short setae.
Antennal peduncle with basal segment longest, slightly longer than 2nd and
3rd segments together, 4th segment slightly longer than 3rd; flagellum with six
segments each with disto-ventral tuft of about 10 setae.
Mandibular palp 3-segmented, basal and 2nd segments equal in length,
each with single distal fringed seta, terminal segment shortest, with five distal
spines; incisor process bearing bluntly rounded irregular teeth.
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 73
Fig. 18. Agulanthura serenasinus sp. nov.
a. g, dorsal view; b. antennule; c. antenna 3; d. antenna 9; e. maxilliped; f. maxilla;
g. mandible; h. peraeopod I; i. peraeopod II; j. peraeopod VII; k. pleopod 1 3; 1. pleopod
2 3; m. telson, with cross-section; n. uropod.
74 ANNALS OF THE SOUTH AFRICAN MUSEUM
Maxilla slender, distally curved, tapering to point, with four or five
subterminal spines. |
Maxilliped 5-segmented, terminal segment set obliquely on penultimate
segment, bearing five distal setae; 2nd segment three times length of 3rd, latter
half length of 4th segment.
Peracopod I longer and stouter than I, unguis almost same length as rest
of dactylus, gently curved; propodus three times longer than wide, palm
sinuous, flanked by irregular row of 15 spine-like setae; carpus triangular, with
seven or eight setae; ischium and basis subequal in length.
Peraeopod II dactylus with very short unguis, slightly curved, with ventral
groove flanked by membranous flange; propodus twice as long as wide, disto-
ventral corner with two short spines, plus short spine bearing accessory spinules;
carpus small, triangular.
Peraeopods IV to VII with dactylus slightly shorter than propodus, gently
curved, ventrally serrate, unguis very short; propodus twice width of dactylus,
bearing several short spines disto-ventrally, plus strong blunt spine bearing
accessory spinules; carpus almost square, distal margin not underriding
propodus but straight, disto-ventral corner serrate, bearing strong blunt spine;
ventral margin bipartite, proximal portion defined by few serrations and very
short spines; merus equal in length to carpus, ventral margin with three equally-
spaced groups of serrations.
Pleopod | outer ramus operculiform, median margin straight, outer margin
evenly convex, bearing elongate distal plumose setae; inner ramus lying ventral
to outer ramus, half width of outer, and slightly shorter.
Pleopod 2 rami subequal in length, tipped with plumose setae, stylet
slender, elongate, apically acute.
Telson lanceolate, widest at midpoint, tapering to narrowly-rounded apex
with proximo-lateral ridge covered by closely adpressed exopod of uropod.
Uropod endopod slightly shorter than telson, tapering evenly to narrowly
rounded apex; basis medially hollowed to accommodate telson; exopod narrowly
leaf-shaped, curved dorsally over, and adpressed to telson; margin fringed with
plumose setae.
Description of 2
Antennule longer than antenna, basal segment equal in length to following
three segments, median face hollowed to accommodate antenna, flagellum of
Six segments, each with distal cluster of setae.
Antenna with three basal peduncle segments subequal, stout, 4th segment
more elongate, flagellum reduced, consisting of only two tiny segments.
All appendages with exception of pleopod 2 as in male.
Material
Holotype SAM-A13553. FAL.487.F 3S 11,0 mm
Allotype SAM-A13554 FAL.685.B-D 9 14,5 mm
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 45
Paratype SAM-A13621 SCD.329.R 2 13,0 mm
Paratype SAM-A13622 SB.310.S 2 14,0 mm
Paratypes SAM-A13623 SST.74.G 392 12,9mm 7,1 mm 6,9 mm
3 11,0 mm
Single female specimens from SST.67.8, SCD.188.P, FAL.763.R-S,
FAL.496.L, FAL.685.B—-D, FAL.666.X-Y, FAL.654.N-R, 6,0 mm-12,3 mm.
Remarks
Agulanthura serenasinus has several features in common with species of
Haliophasma. These include the unsegmented pleon, the 5-segmented maxilliped,
operculiform pleopod 1; also, the carpi of the posterior peraeopods do not
underride the propodi. Several other features in combination, however, seem to
indicate the separation of the present species into a new genus. These features
include the feebly developed eyes, the uropod exopod which is closely adpressed
to the telson, the proximo-laterally ridged telson, the long unguis of the dactylus
of peraeopod I, the square-ended carpus of peraeopod VII, and the uninterrupted
body profile of the animal.
Holoroanthura capensis sp. nov.
Figs 19a—l, 20a—c
Description of °
Head shorter than peraeonal segment I, having slight rostral point. Eyes
absent. Peraeonal segments I to IV gradually increasing in length, segments V
and VI subequal, segment VII slightly shorter. Pleonal segments distinct,
together equal to VIth peraeonal segment in length. Dorso-lateral ridges
distinct only on anterior two peraeonal segments.
Antennule slightly shorter than antenna, consisting of three peduncle
segments, and 4-segmented flagellum tipped with two aesthetascs.
Antennal peduncle 4-segmented, flagellum 7-segmented.
Mandible with incisor portion consisting of upper chitinised portion of
three teeth, and five weakly chitinised teeth, separated from a blunt tooth by a
row of tiny denticles; palp 3-segmented, terminal segment short, bearing three
distal spines, middle segment twice length of basal segment. Maxilla moderately
stout, distally curved with one strong and four or five smaller spines.
Maxilliped slender, Ist free segment three to three and a half times longer
than wide, with digitiform extension at medio-distal angle; 2nd and 3rd segments
subequal, 4th segment half length of 3rd, terminal segment tiny.
Peraeopod I no different in size from peraeopod II, unguis one-third length
of dactylus; palm of propodus bearing three setae; carpus triangular, ventro-
distal extension forming thumb, but not as marked as in following two pairs of
peraeopods.
Peraeopods II and III with thumb-like projection of carpus well developed,
76 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 19. Holoroanthura capensis sp. nov.
a. 9, dorsal view; b. antenna; c. antennule; d. maxilla; e. maxilliped; f. mandible;
g. peraeopod I; h. peraeopod II; i. peraeopod VII; j. uropod; k. telson, with cross-section
at level of arrow; 1. pleopod 1.
bearing two setae and short terminal blunt spine; palm of propodus armed with
two spines.
Peraeopods IV to VII with carpus triangular, underriding propodus; three
distal segments bearing ventral fringe of very fine setules.
Pleopod 1 not operculiform, similar to following pleopods, rami subequal
in length, fringed with long plumose setae.
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 41
Fig. 20. Holoroanthura capensis sp. nov. 3.
a. antenna; b. pleopod 2; c. peraeopod I.
Uropods and telson indurated, slightly splayed. Endopod of uropod almost
twice length of basis, apically truncate, bearing few denticles proximally; outer
margin of exopod sinuous, ending in narrowly rounded lobe, inner margin
straight, bearing four to six denticles, ending in strong triangular tooth well
separated from distal narrow lobe.
Telson margins serrate, distally lanceolate, with strong medio-ventral
rounded ridge.
Description of 3
Antenna consisting of three peduncular segments, basal one longest, plus
flagellum of 13 to 14 segments bearing numerous fine setae.
Peraeopod I unguis one-third length of dactylus, latter curved; propodus
with concave palm, bearing about 10 setae; carpus triangular, distally forming
tiny thumb.
Pleopod 2 with slender slightly curved stylet on inner margin of endopod,
extending well beyond endopod apex, apically blunt, rounded.
78 ANNALS OF THE SOUTH AFRICAN MUSEUM
Material |
Holotype SAM-A13555 WCD.64.P 3 3,8 mm
Allotype SAM-A13624 LBY.72.K. 2 6,1 mm
Paratypes SAM-A13625 WCD.109.A 3 3,1 mm
5 29 4,2-5,0 mm
WCD.64.P 2) 4,0 mm
WCD.111.J 2 4,0 mm
WCD.114.U 49° 3,5-4,9 mm
Remarks
The present material agrees well with the generic definition of Holoroanthura
Menzies & Frankenberg (1966: 41) which is characterized by the lack of eyes,
the possession of normal biting mouthparts, a short unguis on peraeopod I of
the female, the carpus of peraeopods IV to VII underriding the propodus, the
maxilliped 5-segmented, and all pleonal segments distinct.
The type species of the genus, H. irpex, is represented by a single female
recorded off Georgia, U.S.A. H. capensis differs in several respects from the
American species. In the latter the telson is sharply pointed and not indurated,
the spines on the exopods of the uropods elongate, and the inner margin of the
endopod of the uropod as well as the telson margins are entire. In H. capensis
the telson is broadly lanceolate (not sharply tapering) and indurated, the spines
on the inner margin of the exopod of the uropod tiny, the inner margin of the
uropod endopod and the telson margins finely denticulate. The basal maxilliped
segment in H. capensis is more obviously lobed than in the American species.
Suborder FLABELLIFERA
Family Cirolanidae
Cirolana borealis Lilljeborg
Figs 21a—g
Cirolana borealis: Sars, 1897: 70. Hansen, 1905: 342. Richardson, 1905: 101. Schultz, 1969: 182.
Riedl, 1970: 345.
Material
SAM-A13556 SST.57.A—B 2 6,3 mm
SAM-A13557 SST.114.A-—G 2 damaged
Previous records
Atlantic coast of North America, eastern North Atlantic, Mediterranean.
Remarks
The present material agrees well with the above-mentioned descriptions.
A very few differences with the northern species do exist. Unfortunately, no male
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 719
He
SN
Nj
Wl
=at\
'/
Fig. 21. Cirolana borealis Lilljeborg.
a. 2, dorsal view; b. peraeonal epimeres; c. epistome; d. maxilliped;
e. peraeopod I; f. peraeopod VI; g. uropod.
80 ANNALS OF THE SOUTH AFRICAN MUSEUM
is available for comparison of pleopod structure. In the present material the
epimeres are postero-ventrally acute and also have a tiny notch subterminally
especially on epimeres III to VI. The northern species seems to lack this tiny
notch. This feature may, however, be due to the immaturity of the specimens,
as the species apparently matures sexually at about 12 mm length.
Cirolana obtusispina sp. nov.
Figs 22a-i, 23a-f
Description of 3
Body two and a half times longer than wide, smooth, widest at peraeonal
segment III. Head with impressed line joining posterior margins of eyes. Each
peraeonal segment with transverse impressed line in posterior third. Pleo-telson
triangular, tapering to subacute apex, with two submedian dorsal rounded
longitudinal ridges, strongest proximally, becoming obscure distally. Epimeres II
to IV rounded to quadrate. Epimeres V to VII becoming progressively more
acute and elongate. Epimere of pleon segment 4 overlapping 5th. Epistome
longer than wide, distally rounded, with slight lateral ‘shoulders’. Antennule
shorter than antenna, basal segment with ventral groove to accommodate basal
segment of antenna; 3rd peduncular segment twice length of 2nd, flagellum
9-10-segmented. Antennal peduncle 5-segmented, two distal segments largest,
flagellum of about 12 segments.
Mandibles with 3-segmented palp, terminal segment bearing 12-14 curved
simple spines, middle segment with 12-14 serrate spines, molar process produced,
with row of teeth on upper margin; incisor process with three strong chitinised
teeth, and secondary cluster of six smaller spine-like teeth.
Ist maxilla with outer ramus bearing 10 slightly curved spines, some of
which are denticulate; inner ramus with three stout plumose setae.
2nd maxilla outer ramus bilobed, each lobe digitiform, bearing several
plumose setae; inner ramus broadly rounded, bearing several plumose setae.
Maxilliped with 5-segmented palp, 3rd segment broad, 4th segment some-
what lobed on inner margin; endite about half width of basal lobe, tapering
slightly distally, bearing four plumose setae, single very short blunt spine, and
single coupling hook.
Peraeopod I dactylus with distinct unguis; propodus armed with three
spines on ventral margin; carpus triangular, merus bearing five blunt knob-like
modified spines, and three acute spines on ventral margin.
Peraeopod VII with propodus, carpus, merus, and ischium each bearing
several simple and serrate spines on distal margin, plus pair of spines at midpoint
of ventral margin.
Penial processes moderately elongate, digitiform.
Pleopod 1 with roughly rectangular basis bearing five coupling hooks;
endopod one-third width of exopod, latter oval-round.
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 81
Fig. 22. Cirolana obtusispina sp. nov.
a. 2, dorsal view; b. peraeonal epimeres; c. antenna; d. antennule;
e. epistome; f. Ist maxilla; g. maxilliped; g. mandible; i. 2nd maxilla.
Pleopod 2 endopod bearing sabre-shaped stylet on inner margin, extending
well beyond end of rami.
Uropod base produced on inner margin of inner ramus, latter with numerous
setae and few spines; outer ramus lanceolate, shorter than inner, bearing several
setae and two short spines on inner margin.
Material
Holotype SAM-A13558 SST.19.N 3 8,0 mm
Paratypes SAM-A13559 SST.21.U es 5,2 mm
5,0 mm
82 ANNALS OF THE SOUTH AFRICAN MUSEUM
Remarks
The present species is distinct as regards several features. These include the
lack of dorsal sculpture, the 2-ridged pleo-telson, the form of the epistome, and
the knob-like spines of the meri of peraeopods I to HI. Cirolana theleceps
Barnard, 1940 possesses a pleo-telson bearing two longitudinal ridges, but its
denticulate distal margin, and also the shape and character of the uropods
immediately distinguishes this species from C. obtusispina.
Fig. 23. Cirolana obtusispina sp. nov.
a. peraeopod VII; b. peraeopod I; c. penial processes; d. uropod;
e. pleopod 1 3; f. pleopod 2 6.
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 83
Family Sphaeromatidae
Cymodoce alia sp. nov.
Figs 24a-h, 25a—h
Description of 3
Body about two and a half times longer than broad, dorsally strongly
convex. Peraeonal segment I broader than II, ventrally expanded with
anterior lobe running ventral to eye, dorsally with broad transverse groove just
behind head, plus row of small granules near posterior margin, and two larger
submedian granules. Peraeonal segments II to IV with narrow epimeres, each
with tiny granules near posterior margin, and two larger submedian granules.
Epimeres of segments V to VII broader than preceding segments; peraeonal
segment VII overlapping anterior pleon dorsally, finely granular, hind margin
with two prominent tubercles on each side. First three pleon segments granular,
overlapped by VIIth peraeonal segment. Pleon segment 4 with two large conical
submedian tubercles and smaller lateral tubercle. Pleo-telson with two large
conical submedian tubercles, apex notched, trilobed, median lobe smaller and
lower than lateral lobes.
Antennule with basal segment strongly chitinised, large, external face
granular; 2nd segment one-quarter length of first, also strongly chitinised;
3rd segment slender, slightly longer than 2nd; flagellum of 11-12 segments.
Antennal peduncle of five segments, two distal segments longest; flagellum
of 11-12 segments; base hidden by basal segment of antennule. Basal segments
of antennule flanking and closely adpressed to epistome.
Mandible with incisor process strongly chitinised, cutting edge evenly
rounded; molar process also chitinised; six spines between incisor process and
lacinia mobilis; palp 4-segmented.
Ist maxilla outer ramus with 10 curved spines, inner ramus with four
fringed setae.
2nd maxilla with both lobes of exopod tipped with about eight curved
serrated spines; endopod bearing 11-12 fringed setae.
Maxilliped palp with 2nd, 3rd, and 4th segments lobed, lobes tipped with
setae.
Peraeopod I dactylus with short spine at base of strong unguis, propodus
two and a half times longer than wide, with four spines on ventral margin, spines
all apically trifid; carpus triangular, also bearing four trifid spines; merus with
one short and five long spines.
Peraeopod VII more slender and somewhat longer than first three pairs of
peraeopods; propodus twice length of dactylus, with four simple spines on
ventral margin; ventral margins of propodus, carpus, and merus with thick pile
of short setules.
Pleopod 1 with triangular endopod, shorter than exopod.
Pleopod 2 with endopod bearing elongate slender stylet.
Pleopod 3 exopod 2-segmented, endopod broad, median margin straight.
84 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 24. Cymodoce alia sp. nov.
a. 6, dorsal view; b. posterior peraeon and pleon 9; c. 3, lateral
view; d. epistome; e. Ist maxilla; f. 2nd maxilla; g. antenna;
h. antennule.
Pleopod 4 endopod with transverse pleats, apex with tiny lobe.
Pleopod 5 exopod bearing five spinule-bearing cushions, endopod bilobed,
bearing transverse pleats.
Uropod with inner ramus fused to base, leaf-shaped, oval in cross-section,
outer ramus longer than inner, curved towards midline, basally oval in cross-
section, distally circular in cross-section, both rami bearing short setules.
Description of &
Not as granular as male, conical submedian process on 4th pleon segment
and pleo-telson not as large as in male. Telsonic apex notched, trilobed, but
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 85
Fig. 25. Cymodoce alia sp. nov.
a. pleopod 1 3; b. pleopod 2 3; c. pleopod 3; d. pleopod 4; e. pleopod 5; f. peraeopod VI;
g. maxilliped; h. peraeopod I.
curved ventrally. In dorsal view, median lobe longer than lateral lobes, latter
just visible. Inner uropodal ramus distally quadrate, flattened, as long as outer
ramus, latter leaf-shaped. Peraeonal segment VII not extending over pleon.
Material
Holotype . SAM-A13560 SST.21.S 3 9,0 mm
Allotype SAM-A13560 SST.21.S 2 ovig. 8,5 mm
Paratypes SAM-A13561 SST.21.S Q 8,5 mm
SST.17.A-B @ juv.
86 ANNALS OF THE SOUTH AFRICAN MUSEUM
Remarks
The following features are used to place the present material in the genus
Cymodoce; hemibranchiate pleopods, pleo-telsonic apex notched in both sexes.
maxillipedal segments lobed, both uropodal rami well-developed, no medial
process on pleo-telson, mouthparts sexually dimorphic.
The present species most closely resembles Cymodoce amplifrons Stebbing
of the numerous South African representatives of this genus, in possessing a
strong pair of conical processes on both the last pleon segment and on the pleo-
telson. The nature of the uropods and the trilobed medial lobe of the pleo-telson
in the male of Stebbing’s species, distinguish it from the present species.
Cymodoce velutina sp. nov.
Figs 26a—k, 27a-e
Description of 3
Body about twice longer than wide, dorsally strongly convex. Most of
dorsal surface of head, peraeon, pleon, and pleo-telson covered with tiny
delicate membranous scale-like structures, with stout apically bifid seta arising
from base of each. Pleo-telson smoothly convex, apex notched, trilobed, median
lobe slightly longer than lateral lobes.
Antennular peduncle 3-segmented, basal segment strongly chitinised,
flagellum | 1—12-segmented.
Antennal peduncle 5-segmented, distal segment longest; flagellum
10-segmented. Mandible with strongly-chitinised incisor process having straight
edge, tridentate lacinia mobilis; molar process broad, bristle-covered; palp
3-segmented.
Ist maxilla outer ramus with about 10 curved spines, inner ramus with four
apical fringed setae.
2nd maxilla with both lobes of exopod tipped with curved serrate spines,
endopod bearing about six fringed setae.
Maxilliped palp with 2nd, 3rd, and 4th segments lobed, lobes tipped with
setae. Peraeopod I shorter than following peraeopods, dactylus with short spine
at base of strong unguis, propodus, carpus, and merus each carrying four fringed
spines, ischium with numerous fine setae on ventral margin. Following
peraeopods similar to I, but with meri and bases somewhat longer.
Penial processes slender, elongate, apically slightly broadened.
Pleopod 1 endopod about half width of exopod.
Pleopod 2 with stylet on endopod extending beyond apex of ramus, slender,
apically slightly hooked.
Uropod with inner ramus apically truncate, outer ramus about half length
of inner, leaf-shaped.
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 87
Fig. 26. Cymodoce velutina sp. nov.
a. g, dorsal view; b. 92, pleo-telson dorsal view; c. antenna; d. antennule; e. 1st maxilla;
f. maxilliped; g. peraeopod I; h. 2nd maxilla; i. epistome; j. penial processes; k. one scale
and seta.
88 ANNALS OF THE SOUTH AFRICAN MUSEUM
Description of °
Very similar to male, pleo-telson slightly broader, apex trilobed, lobes
subequal.
Material
Holotype SAM-A13629 FAL.700.R-T 3 5,9 mm
Allotype SAM-A13562 SST.20.Z 2 ovig. 8,0 mm
Paratypes SAM-A13630 FAL.700.R-T 3 go 6,0-6,9 mm
Remarks
The unmistakable body covering of this species is not found in any of the
other southern African species of Cymodoce, nor in any of the exotic species.
The unsculptured pleo-telson with trilobed apex is also distinctive.
Fig. 27. Cymodoce velutina sp. nov. 3.
a. pleopod 1; b. pleopod 2; c. pleopod 3; d. pleopod 4; e. pleopod 5.
MARINE ISOPODA FROM THE CONTINENTAL SHELF OF SOUTH AFRICA 89
ACKNOWLEDGEMENTS
I am grateful to Professor J. H. Day of the Department of Zoology of the
University of Cape Town, for making the present collection available to the
South African Museum for description.
My thanks are due to Dr J. Field of the Department of Zoology, for
preliminary identifications of some of the isopods, and to Dr N. Christie of the
Department of Zoology for the specimen of Holoroanthura from Lambert’s Bay.
My sincere thanks are due to Dr George A. Schultz of Hampton, New
Jersey, for reading the manuscript, and for his many helpful comments and
criticisms.
REFERENCES
BARNARD, K. H. 1920. Contributions to the crustacean fauna of South Africa.— Ann. S. Afr.
Mus. 17: 319-438.
BARNARD, K. H. 1925. A revision of the family Anthuridae (Crustacea Isopoda) with remarks
on certain morphological peculiarities.—J. Linn. Soc. (Zool.) 36: 109-160.
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
identifications of the hitherto recorded marine and fresh-water species.—Ann. S. Afr.
Mus. 32: 381-543.
BARNARD, K. H. 1957. Additions to the fauna-list of South African Crustacea.— Ann. Mag.
nat. Hist. (12) 10: 1-12.
HAnsEN, H. J. 1905. Revision of the European marine forms of the Cirolaninae a subfamily of
the Crustacea Isopoda.—J. Linn. Soc. (Zool.) 29: 337-372.
KOEHLER, R. 1911. Arcturidés nouveaux provenant des campagnes de la ‘‘Princesse-Alice”’
ou appartenant au Museé Océanographique de Monaco.— Bull. Inst. océanogr. Monaco
214: 1-65.
MeNzzeS, R. J. & FRANKENBERG, D. 1966. Handbook of the common marine isopod Crustacea
of Georgia. Athens, Georgia: University of Georgia Press.
NIERSTRASZ, H. F. 1941. Die Isopoden der Siboga-Expedition. IV. Isopoda Genuina. III.
Gnathiidae, Anthuridae, Valvifera, Asellota, Phreatocoidea.—Siboga Exped. monogr.
32d: 1-72.
NORDENSTAM, 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): 1-284.
OHLIN, A. 1901. Isopoda from Tierra del Fuego and Patagonia.— Wiss. Ergebn. schwed.
Exped. Magellansland 2: 261-306.
RICHARDSON, H. 1905. A monograph on the Isopods of North America. — Bull. U.S. natn. Mus.
54: 1-727.
RIEDL, R. 1970. Fauna und Flora der Adria. Hamburg & Berlin: Paul Parey.
Sars, G. O. 1897. An account of the Crustacea of Norway with short descriptions and figures of
all the species. 2: Isopoda. Bergen: Bergen Museum.
SCHULTZ, G. A. 1969. How to know the marine isopod crustaceans. Dubuque, Iowa: William C.
Brown.
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.
r 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 Haniey, 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 .. .’
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(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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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.
BRIAN KENSLEY
MARINE ISOPODA
FROM THE CONTINENTAL SHELF
OF SOUTH AFRICA
OLUME 67 PART 5 AUGUST 1975 : ISSN 0303-2515
_ ANNALS
OF THE SOUTH AFRICAN
CAPE TOWN
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(a) Author’s name and year of publication given in text, e.g.:
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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.
Koun, 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
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
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Part 5 Deel
THE AMPHIPODA OF SOUTHERN AFRICA
PART 5
THE GAMMARIDEA AND CAPRELLIDEA OF
fae CAPE PROVINCE WEST OF CAPE AGULHAS
By
Cc. L. GRIFFITHS
Cape Town Kaapstad
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THE AMPHIPODA OF SOUTHERN AFRICA
PART 5
THE GAMMARIDEA AND CAPRELLIDEA OF THE
CAPE PROVINCE WEST OF CAPE AGULHAS
By
C. L. GRIFFITHS
C.S.I.R. Oceanographic Research Unit, University of Cape Town
(With 21 figures)
[MS accepted 18 December 1974]
ABSTRACT
A systematic account of the marine gammaridean and caprellid Amphipoda of the Cape
Province of South Africa west of Cape Agulhas (20°E) is provided. The analysis is based on
collections totalling over 70 000 specimens in the possession of the University of Cape Town
and of the South African Museum, as well as records from the existing literature.
Two hundred and thirty-two species are recognized from the area. These include fourteen
species which are recorded for the first time from southern Africa, as well as the following
eleven species which are described as new to science: Panoploea stegosaura, Maera emarginata,
Maera komma, Maera thrixa, Melita mucronata, Listriella saldanha, Socarnes septimus,
Perioculodes pallidus, Heterophoxus cephalodens, Heterophoxus opus and Podocerus pyurae.
In addition the following changes in taxonomy are proposed:
A new family—the Temnophliidae—is erected for Temnophlias K. H. Barnard, 1940.
Ceradocus aviceps K. H. Barnard, 1940 is removed to Quadrivisio. Tryphosella africana
K. H. Barnard, 1955 is synonymized with Hippomedon longimanus. Tryphosella normalis
K. H. Barnard, 1955 is removed to Hippomedon and H. rotundipleura Ledoyer, 1973 synony-
mized with it. Microlysias indica K. H. Barnard, 1937 is synonymized with M. xenoceras.
Uristes induratus K. H. Barnard, 1925, is removed to Procyphocaris where P. primata
J. L. Barnard, 1961 falls into synonymy with it. Podocerus cristatus of K. H. Barnard (1916)
and Griffiths (1973, 1974a, c) is referred to P. inconspicuus. Allorchestes inquirendus is
synonymized with Hyale grandicornis, Talorchestia inaequalipes with Orchestia gammarella
and Caprella falsa with C. penantis.
CONTENTS
PAGE
Introduction : . , 2 : : 91
The collecting stations . : : : ; 93
Collections from estuaries . ' : : 93
Collections from the marine environment . 95
Systematics . : ; : : : : 99
Gammaridea i ; , : : ; 100
Caprellidea . ; : : : : Sa an fe
Acknowledgments 2 é : : oS lies
References . ‘ : : é ; ee he)
INTRODUCTION
The following account forms the fifth and last part of a series surveying
the known gammaridean and caprellid Amphipoda of various geographical
zones within southern Africa (defined as Africa south of 20°S). Previous sections
a1
Ann. S. Afr. Mus. 67 (5), 1975: 91-181, 21 figs.
92 ANNALS OF .THE SOUTH AFRICAN MUSEUM
have dealt with Mocambique (Griffiths 1973), South West Africa (Griffiths
1974a), Natal (Griffiths 1974b) and the Cape Province east of Cape Agulhas
(Griffiths 1974c), while the present portion deals with the remaining area between
Cape Agulhas and the South West African border.
Oceanographic conditions along the Atlantic coast from Cape Point to
South West Africa are primarily influenced by the Benguela Current. The cold
waters of this current originate in the sub-antarctic, but before reaching the
African coast they have been covered by a thick layer of warmer, less dense
South Atlantic water, 200-500 m thick. However, along the coastal margin the
dominant south or south-easterly winds of summer frequently blow the warm
oceanic water offshore, allowing cold Benguela water to upwell to the surface.
Thus although surface temperatures a few kilometres offshore average 15—17°C
the narrow band of inshore upwelled water may be as cold as 8°C (normally
10-14°C). As wind stress varies so the width of the upwelled zone waxes and
wanes, indeed during periods of north-westerly winds the warmer Atlantic
water extends to the shoreline.
Conditions in the area between Cape Agulhas and Cape Point are even
more variable than those experienced along the west coast, since warm Agulhas
Current and cold Benguela Current waters mix in this region. The-limits of the
warm and cold waters are controlled largely by wind conditions and are
consequently highly variable. During periods of south-easterly winds, par-
ticularly in summer, warm Agulhas water may extend into False Bay, and as an
offshore tongue of water, right around Cape Point. Where this water reaches —
the shore the temperature may rise above 20°C, but this rise is not universal
since local upwelling cells are produced along the west coasts of projecting shore
features, creating much colder conditions. Conversely north-westerly winds of
winter force warmer surface waters offshore and allow cold Benguela water to
round Cape Point and upwell along the east coasts of peninsulas. As a result of
this upwelling system the fauna of peninsulas along this stretch of coast tends to
contain more cold-water forms while in sheltered bays more warm-water forms
are to be found.
These unusual physical conditions and the proximity to the marine research
centres of Cape Town have combined to attract a great deal of research effort to
the south-western Cape coast. The marine benthos of the area has been
extensively sampled, with the exception of the region north of Lambert’s Bay,
which remains virtually untouched. The University of Cape Town alone has
collected more than 2 000 benthic samples between Lambert’s Bay and Cape
Agulhas, particular attention having been given to the Saldanha Bay complex
and to False Bay, each of which accounts for over 500 samples. In addition,
other South African institutions and international expeditions have made
smaller benthic collections in the area. Extensive intertidal collections also
exist, notably those of K. H. Barnard (presently housed at the South African
Museum) and of the University of Cape Town. These collections largely
originate from the Cape Peninsula but include samples from numerous localities
THE AMPHIPODA OF SOUTHERN AFRICA 93
between Cape Agulhas and Port Nolloth. Compared with the Indian Ocean
coast, the Atlantic coast of the Cape Province is poor in estuaries but those
which do exist have been comparatively well studied, allowing fruitful
comparison with the more numerous estuaries of other regions.
THE COLLECTING STATIONS
The scope of the collections from the area considered here is so great that
it has become impracticable to list individual stations, as has been the format in
previous parts of this series. Instead a generalized discussion of the various
collecting areas is given and the distributions of individual species are provided
in the more compact form outlined in the systematic section.
Collections from estuaries
(a) Klein River Estuary, Hermanus
A full description of the Klein River Estuary and of its fauna is provided
by Scott, Harrison & Macnae (1952). The estuary extends from Walker Bay for
a distance of about 12 km to the village of Stanford. Initially the river is canal-
like before opening up into a shallow lagoon about 0,75 km wide. The lagoon
consists mostly of muddy shallows through which deeper channels meander, but
near the mouth the bottom becomes sandy with rocky outcrops along the shore.
The lagoon is closed for most of the year but during the winter rains it fills
steadily until a passage is artificially cut through the sandbar at the mouth to
prevent flooding. The mouth remains open during the spring, when the lagoon
is tidal, and then gradually closes, remaining so until the next year’s rains.
On sandy shores around the lagoon numerous Talorchestia australis are to
be found, whereas this species is replaced by Orchestia ancheidos under stones
and in gravel areas. Amongst the weeds of the lagoon Melita zeylanica is
abundant, as is Corophium triaenonyx. The tubicolous Grandidierella lutosa, a
species found only in this vicinity, is common on the mudflats, although
Corophium triaenonyx is present here too. Orchestia rectipalma occurs under
stones on gravel bottoms as well as amongst weeds. In the upper reaches of the
estuary Quadrivisio aviceps has its only known habitat.
(6) Milnerton River system
This system, which consists of a river, the Diep River, which flows into a
shallow lake, Riet Vlei, and from there through the Milnerton Estuary to the
sea, has been fully described by Millard & Scott (1954). At the time of this study
the system was relatively undisturbed, but Riet Vlei is at present being developed
into a marina. In its original state the system experienced a consistent fresh-water
flow during winter, but in summer a sandbar closed the mouth while Diep River
and Riet Vlei dried out, causing the estuary to become hypersaline.
Under these rigorous conditions of fluctuating salinity the fauna of the
94 ANNALS OF THE SOUTH AFRICAN MUSEUM
Honieklip
Bay
Fig. 1. The Atlantic coast of the Cape Province showing main collecting areas.
THE AMPHIPODA OF SOUTHERN AFRICA 95
system was relatively impoverished. The amphipod fauna consisted of only two
species, Afrochiltonia capensis and Melita zeylanica. Both these well-known
euryhaline forms were found amongst weeds and polychaete tubes in the lower
reaches of the estuary, where they were most abundant in early summer.
(c) Berg River Estuary
The Berg River Estuary was the subject of a brief collecting expedition by
the University of Cape Town in September 1949. The estuary is about 60 m wide
and 6 km long with a maximum depth of about 10 m, and enters the sea at the
head of St Helena Bay. Talorchestia quadrispinosa is common on the sandbanks
around the mouth, while the waters of the estuary harbour the typical estuarine
species Afrochiltonia capensis, Melita zeylanica, Orchestia rectipalma and
Orchestia ancheidos.
(d) Olifants River Estuary
The Olifants River Estuary consists of a fairly straight stretch of water
about 200 m wide and 2-5 m deep flowing through a deep valley surrounded by
arid scrubland. The flood plain consists of a muddy saltmarsh partially cut off
by a shallow blind arm which represents a previous river mouth. The present
mouth is fringed by sandbanks to the south, while the north bank is rocky. The
river bed is mostly soft mud which becomes progressively more sandy towards
the mouth.
Eight amphipod species have been collected from the estuary. Five of these
are typical marine forms and were found only around the mouth, while the
remaining three species were common further upstream. These euryhaline
species are Afrochiltonia capensis, Melita zeylanica and Orchestia rectipalma.
Collections from the marine environment
(a) Shore collections
During early studies by the University of Cape Town on the constitution
of the rocky intertidal fauna around the South African coast, collections were
made at a number of west coast localities. These stations were more or less
evenly spaced between Cape Agulhas and Buffels River, there being eleven
stations between these two points. More recently numerous other shore
collections have been made, particularly around the Cape Peninsula.
Sandy beaches in these areas may harbour enormous populations of either
Talorchestia quadrispinosa or Talorchestia capensis along the drift line and above,
while at lower levels Bathyporeia sp. and Urothoe elegans are to be found
burrowing in the sand. The fauna of rocky areas is considerably more complex
but is dominated by Hyale spp., principally H. grandicornis and H. saldanha.
At lower levels Ceradocus rubromaculatus and Lysianassa ceratina are common
under rocks, while Aora kergueleni, Amaryllis macrophthalma, Paramoera
96 ANNALS OF THE SOUTH AFRICAN MUSEUM
capensis and Caprella spp. occur in great numbers amongst algae and bryozoa.
The unusual Temnophlias capensis is to be found crawling over rock surfaces
while many other species are locally abundant.
(b) Langebaan Lagoon
A full description of Langebaan Lagoon and of its fauna is given by
Day (1959). The lagoon is about 3 km wide and extends for about 15 km south
of its origin on the southern shore of Saldanha Bay. The whole area is completely
protected from the sea by a narrow finger of land projecting parallel to the shore
and by two small islands at the mouth of the lagoon. Tidal flow is considerably
restricted and retarded, particularly at the head of the lagoon. The water is
generally clear and the bottom mainly sand. The consistency of the sand varies
greatly, being fine and clean at the mouth of the lagoon, whereas in the body of
the lagoon the beach slopes fairly steeply to mean sea-level and then flattens out
into a wide, waterlogged bed of fine, often muddy sand. These sandflats are
riddled with Callianassa holes and abound in patches of the loose alga Gracilaria
and of Zostera. At the high-water mark saltmarsh vegetation occurs, this zone
becoming extremely wide at the head of the lagoon. Salinity in the lagoon is
generally as high or slightly higher than experienced in the open sea, while solar
radiation also tends to raise the temperature somewhat.
The amphipod fauna of Langebaan Lagoon is rich and varied, some
65 species having been recorded there. The rocky islands at the mouth of the.
lagoon and isolated rocky patches elsewhere support large populations of
Elasmopus affinis, Paramoera capensis, Cymadusa filosa and Hyale and Caprella
spp. living amongst the rich algal cover. Ceradocus rubromaculatus is common
under rocks, while Polycheria atolli is frequently recovered from sponges and
compound ascidians. Along the beaches Talorchestia spp. are fairly common at
the driftline, while the rich fauna of the Zostera and Gracilaria beds is dominated
by Cymadusa filosa and Lysianassa ceratina. At lower tidal levels the mudflats
are inhabited by the tubicolous Ampelisca palmata and burrowing Urothoe spp.
as well as Lysianassa ceratina. Bathyporeia sp. becomes common at lower water
springs. The sand-bottomed channels which cut through the sandflats yield a
variety of species of which Lysianassa ceratina and Paramoera capensis are the
most abundant. Both these species, as well as Ampelisca palmata and Perioculodes
longimanus, also occur in the plankton at night.
(c) Saldanha Bay
Lying about 120 km north of Cape Town, Saldanha Bay consists of an
almost semicircular bay about 8 km across, from the southern end of which
stretches Langebaan Lagoon (above). The main bay is moderately protected
from wave action by rocky headlands, between which lie three small islands.
The eastern shore of the bay is formed of a long sandy beach interrupted by a
few isolated rocky points. The bottom of the bay consists almost entirely of
sands of various textures, these tending to be coarse and shelly in areas of
THE AMPHIPODA OF SOUTHERN AFRICA 97
turbulence, particularly around the islands, and finer elsewhere. Extensive
harbour development is at present taking place in the area and this can be
expected to result in major changes in the physical conditions and fauna in the bay.
The fauna of Saldanha Bay has been extensively sampled and 72 amphipod
species have been recorded there. Along the driftline of sandy beaches Talor-
chestia australis is common, while rocky points along the shore are populated by
numerous Ceradocus rubromaculatus, Aora kergueleni, and Hyale spp. Caprella
spp. are also common on algae and hydroids and Polycheria atolli is frequently
found burrowing into compound ascidians and sponges. Amongst the benthos
of the bay Lysianassa ceratina and Paramoera capensis are the most frequently
encountered species, while Ampelisca anomala dominates areas of shelly sand.
Where the sand is fine and clean Ampelisca brevicornis is abundant with a
number of other species also common, particularly Urothoe grimaldi, Orcho-
mene plicata, Bathyporeia sp., Perioculodes longimanus and Megaluropus
namaquaeensis. Local concentrations of Photis spp. and of Siphonoecetes
dellavallei are associated with solid objects on the bottom.
(d) False Bay
A description of False Bay and its biology is given by Day (1970). The bay
is roughly square with a side of some 35 km. Its eastern and western shores are
generally precipitous, although sandy beaches do occur along the Cape
Peninsula to the west, notably Simonstown and Fish Hoek. The northern shore
of the bay is flat and low-lying, forming wide sandy beaches from which the
bottom shelves gently towards the mouth, where it reaches a maximum depth of
about 90 m. The substrate of the bay consists largely of sand; this may be fine
and clean, or coarse and mixed with shell in the shallower areas, but over most
of the bay it is fine and khaki-coloured, merging into green mud in deeper water
near the mouth. Numerous rocky patches are to be found throughout but are
particularly numerous to the east.
Hydrological conditions in the area are of particular interest since the bay
is subject to both Atlantic and Indian Ocean regimes. Under the influence of
north-westerly winds in winter, surface water is blown south and cold Benguela
water enters the bay around Cape Point, causing the temperature to drop to
13-14°C at the surface and as low as 10°C in the deeper parts. With the spring
south-easterly winds Agulhas water drifts into the bay, striking the western shore
and drifting clockwise from there. At this stage surface temperatures average
17-18°C but may rise to 20°C by the end of summer when blue Agulhas water
dominates the bay. At this stage a distinct thermocline is usually found at
about 20 m.
As a result of its unusual hydrological regime and its proximity to Cape
Town, False Bay has been the subject of a good deal of research. The Zoology
Department of the University of Cape Town has collected some 500 benthic and
numerous intertidal samples from the bay and recorded over 140 amphipod
species from these.
98 ANNALS OF THE SOUTH AFRICAN MUSEUM
The amphipod fauna of False Bay is both rich and complex, the species
composition varying widely with depth and substrate composition. The clean
sands of the surf zone are dominated by burrowing forms, particularly
Perioculodes longimanus and Urothoe grimaldi in the shallower areas, and
Urothoe pulchella and Mandibulophoxus stimpsoni slightly deeper. The fine
hard-packed sands beyond the surf zone are predominantly occupied by
tubicolous filter-feeders, notably Ampelisca anomala, A. palmata and A. brevi-
cornis, although both Photis uncinata and Aora gibbula are locally common. The
deepest areas of the bay are composed of green muds and here Hippomedon
normalis, a scavenging species, is the most common amphipod, although
Ampelisca brevicornis also extends into this zone.
The fauna of rocky areas is very diverse with numerous Amaryllis macro-
phthalma. Other lysianassids such as Lysianassa variegata and Orchomene plicata
are also common. Ampithoe ramondi is well represented, as are Caprella spp.
Paramoera capensis is the most common species in the bay and is found both in
rocky and sandy areas as well as at night in the plankton.
(e) Lambert’s Bay
The University of Cape Town has recently undertaken a study of benthic
distribution in the Lambert’s Bay area. Sampling has taken the form of a
transect running from high-water springs to a depth of 800 m. The substrate
from 0 to 60 m consists of fine or medium well-sorted sands, while between
80 and 100 m there is an area of muddy sand characterized by the presence
of the large tubicolous polychaete, Diopatra monroi. Below this zone lies a
further area of fine silty sand which extends to the limit of the transect line
at 800 m.
The amphipod fauna can be considered as falling into a number of faunistic
zones. The first of these is the driftline fauna, the only amphipod represented
here being Talorchestia quadrispinosa. The lower intertidal levels and surf zone
(to 5 m depth) are characterized by a dominance of burrowing forms such as
Perioculodes longimanus, Bathyporeia sp., Urothoe grimaldi and especially the
deep-burrowing Cunicus profundus. The area between 5 and 40 m has a rich
amphipod fauna with Paramoera capensis the most common species. Burrowing
forms, particularly Bathyporeia sp. and Perioculodes longimanus, are still
common, while tubicolous forms, notably Ampelisca brachyceras and Photis
longidactylus, become important. Cunicus profundus is replaced by other
haustoriids, notably Urothoe grimaldi and Urothoe pulchella. The muddy sands
of the Diopatra zone between 80 and 100 m have a similar fauna, particularly
abundant in Paramoera capensis, Megaluropus namaquaeensis and Bathyporeia
sp., however Ampelisca brachyceras is replaced by Ampelisca anomala. Ampelisca
anomala continues as the dominant ampeliscid to 120 m, where it is replaced by
the larger A. brevicornis. Stations deeper than 120 m are also marked by the
appearance of such deep-water forms as Paraphoxus oculatus, Eriopisella
capensis and Hippomedon onconotus.
THE AMPHIPODA OF SOUTHERN AFRICA 99
(f) Other benthic samples
In addition to the discrete collections discussed above numerous other
collections exist, particularly in the possession of the University of Cape Town.
These include material originating from commercial trawlers; donated by the
Division of Sea Fisheries and collected by the university’s research vessels
R.V. Gilchrist and Thomas B. Davie. The area covered is primarily that between
Cape Point and Lambert’s Bay. Very few of the samples are from further north
or from depths of more than 200 m and many species undoubtedly await
discovery in these regions.
Since the collections were not planned with any predefined concepts of
faunistic analysis in mind, their interpretation is complicated by the interaction
of variables of location, season, depth, substrate and collecting gear. Although
over 100 species are represented, some are notable in their scarcity. For example,
Mandibulophoxus stimpsoni, so abundant in False Bay and eastward, is rare
along the west coast, as is Photis uncinata, which is replaced by the similar
P. longidactylus. Sandy and muddy areas are normally dominated by burrowers
(Perioculodes, Urothoe, Bathyporeia) and tubicolous forms (Byblis, Photis and
especially Ampelisca), although Paramoera, Lysianassa and Hippomedon are also
common. Relatively few samples originate from rocky areas, but these include
numerous Lysianassa, Paramoera, Chevalia, Maera and Leucothoe as well as
various caprellids.
SYSTEMATICS
The taxonomy of Gammaridea follows the system outlined by J. L. Barnard
(1969) and subsequently revised by J. L. Barnard (1970a, 1972b, 1973). The
revision of the family Gammaridae proposed by Bousfield (1973) is still
contentious (Holsinger 1974) and has thus been disregarded here. Familial taxa
within the Caprellidea are those proposed by McCain (1970).
The system of listing individual records for each species, as employed in
earlier portions of this series, has proved too cumbersome to encompass the
2 000 stations and 70 000 specimens reported on here. A more concise system
has thus been devised in which the ranges of species are indicated by the
latitude/longitude squares in which they have been recorded. These are followed
by letters denoting the depth of the various records (E = estuarine, I = inter-
tidal, T = 0-29 m, O = 30-99 m, D = 100-499 m, VD = 500-1 000 m). The
latitude/longitude square 34°S/18°E includes both the cold Atlantic and warmer
False Bay coasts of the Cape Peninsula, and in order to distinguish between
these the letters FB are appended to records from False Bay. Thus a range
indicated as 32/18/I, S to 34/18/FB/I, T indicates that the species in question is
found intertidally and at depths of less than 30 m between 32°S/18°E and
34°S/18°E (False Bay).
In the account which follows the arrangement of families, genera and
species is alphabetic. Limbs of the pereon are referred to as gnathopods | and 2,
100 ANNALS OF THE SOUTH AFRICAN MUSEUM
followed by pereiopods 1-5 and the segments of these limbs are referred to as
articles 1-7, article 1 being the coxal plate (whether this is present or absent).
The analysis is restricted to species occurring between the driftline and 1 000 m
depth, with estuarine species being included. Holotypes of all new species and
representative material of all other species have been placed in the South
African Museum, Cape Town.
Suborder GAMMARIDEA
Family Acanthonotozomatidae
Iphimedia capicola K. H. Barnard, 1932
Iphimedia capicola K. H. Barnard, 1932: 118, fig. 66.
Records: 34/18/FB/T, O to 32/18/T, O, moderately common.
Distribution: Endemic, Port Elizabeth to Lambert’s Bay.
Panoploea excisa K. H. Barnard, 1932
Panoploea excisa K. H. Barnard, 1932: 129, fig. 73.
Records: 33/17/T, a single record.
Diagnosis: Rostrum acute, not strongly deflexed; pereon segments 1-6 smooth,
7 with a small pair of dorsal denticles; pleon segments 1-3 each with a pair of
dorsolateral procumbent teeth; gnathopod 1 chelate, gnathopod 2 subchelate;
article 2 of pereiopod 5 posteriorly serrate, postero-distally excised to leave a
large semicircular concavity; telson oblong, apically truncated, slightly
emarginate.
Distribution: Endemic, the above record is the only one to date.
Panoploea stegosaura sp. nov.
Fig. 2
Description of male (4 mm): Head slightly shorter than enlarged pereon
segment 1, rostrum downturned, extending to tip of article 1 of antenna 1, eyes
round, rust brown; articles 1 and 2 of antenna 1 medio-distally produced into
an acute tooth, article 3 slender (flagellum broken), accessory flagellum absent;
flagellum of antenna 2 broken; upper lip slightly incised apically; mandible
without molar, consisting simply of a subacute process, palp 3-articulate
(Fig. 2B); lower lip (Fig. 2C) with inner lobes, their apices obscurely incised ;
palp of maxilla 1 (Fig. 2D) bi-articulate, not extending to apex of outer plate,
outer plate bearing 14 strong serrate spines, inner plate with two short apical
setae; plates of maxilla 2 subequal; palp of maxilliped (Fig. 2E) 3-articulate,
article 2 medially produced to apex of article 3, both articles terminally setose,
THE AMPHIPODA OF SOUTHERN AFRICA 101
Fig. 2. Panoploea stegosaura sp. nov.
Male, 4mm: A —lateral aspect; B—mandible; C—lower lip; D—maxilla 1; E—maxilliped;
F—gnathopod 1; G—tip of gnathopod 1 enlarged; H—gnathopod 2; I—articles 6 and 7 of
gnathopod 2 enlarged; J—telson.
102 ANNALS OF THE SOUTH AFRICAN MUSEUM
outer plate bearing short marginal setae, inner plate with 18 short marginal
plumose setae. :
Pereon segments all medio-dorsally keeled, 1 produced anteriorly over head,
7 with a posterior pair of sub-dorsal teeth; anterior coxae acuminate,
4 posteriorly excavate, 5—7 posterodistally produced into acute teeth; gnathopod
1 minutely chelate (Fig. 2F—G), articles 5 and 6 subequal, 6 and 7 distally
bearing a few long plumose setae; gnathopod 2 weakly chelate (Fig. 2H-l),
palm defined by a small spine, posterior margin with a row of pectinate setae; —
pereiopods | and 2 slender, article 4 antero-distally lobed (articles 5—7 missing);
posterior margin of article 2 of pereiopods 3 and 4 with a strong medial tooth,
pereiopod 5 with four teeth on posterior margin of article 2; article 4 of
pereiopods 3-5 slightly produced antero-distally, strongly lobed postero-distally
(articles 5-7 missing).
Pleon segments 1 and 2 each with an anterior medio-dorsal tooth and a
posterior pair of upright sub-dorsal processes; pleon segment 3 with two
triangular medio-dorsal teeth and a pair of reverted sub-dorsal processes
posteriorly; first pleonal epimeron postero-distally rounded but with an acute
process at centre of posterior margin, second pleonal epimeron similar but
postero-distally acute; third pleonal epimeron with stronger posterior and
postero-distal teeth; uropods | and 3 extending equally, slightly exceeding apex
of uropod 2, rami lanceolate, outer ramus of uropods 2 and 3 slightly the
longer; telson apically truncated, broadly excavate (Fig. 2J).
Holotype: SAM—A13228, male, 4 mm, unique.
Type-locality: 34°14’S/18°29’E, 15 February 1965, depth 40 m, substrate coarse
khaki sand.
Relationships: The highly developed dorsal armature of this species and the
unusual manner in which pereon segment | is produced anteriorly over the head
distinguish it from other members of the genus, none of which show carinae on
all pereon segments.
Family Ampeliscidae
Ampelisca acris Griffiths, 1974
Ampelisca excavata (non K. H. Barnard, 1925): K. H. Barnard, 1955: 82, fig. 40A.
Ampelisca acris Griffiths, 1974c: 268, fig. 3.
Records: 34/18/FB/T, O, 34/18/O, a few records.
Distribution: Endemic, Port Elizabeth to Cape Peninsula.
Ampelisca anisuropa (Stebbing, 1908)
Byblis anisuropus Stebbing, 19085: 12, pl. 10.
Ampelisca anisuropa: Griffiths, 19746: 220.
Records; 32/16/D to 34/18/D and 34/18/FB/T, O, quite common.
Distribution: Endemic, Natal to Lambert’s Bay.
THE AMPHIPODA OF SOUTHERN AFRICA 103
Ampelisca anomala Sars, 1882
Ampelisca anomala: Sars, 1895: 178, pl. 62 (fig. 2).
Records: 31/18/O,D and 32/16/VD to 34/18/T,O,D and 34/18/FB/T, O,
locally abundant.
Distribution: Scandinavia, southern Africa.
Ampelisca brachyceras Walker, 1904
Ampelisca brachyceras Walker, 1904: 252, pl. 2 (fig. 13).
Records: 32/18/T to 34/18/FB/T, O, many records.
Distribution: Ceylon, southern Africa.
Ampelisca brevicornis (Costa, 1853)
Ampelisca brevicornis: Reid, 1951: 204-210, figs 9-15. Kaim Malka, 1969: 928-932, pls 1-6.
Records: 32/18/T, O, 32/17/D, 32/16/D to 34/18/FB/T, O, numerous records.
Distribution: Cosmopolitan.
Ampelisca chiltoni Stebbing, 1888
Ampelisca chiltoni: J. L. Barnard, 1961: 61, fig. 31.
Records: 32/16/D, 34/18/I and 34/18/FB/T, O, a few records.
Distribution: Australia, New Zealand, southern Africa.
Ampelisca diadema (Costa, 1853)
Ampelisca diadema: Chevreux & Fage, 1925: 82, fig. 74.
Records: 32/18/T, O, 32/17/D, 32/16/D to 34/18/FB/T, O, locally common.
Distribution: Cosmopolitan.
Ampelisca excavata K. H. Barnard, 1925
Ampelisca excavata K. H. Barnard, 1925: 336, pl. 34 (figs 5-7). Gray & J. L. Barnard, 1970:
67-83, figs 1-5, pl. 1.
(non) Ampelisca excavata: K. H. Barnard, 1955: 82 (= A. acris, above).
Records: 34/18/FB/I, T, uncommon.
Diagnosis: Antennae subequal, about 40°%% body length; three pairs of eyes
with obscure corneal lenses; article 5 of pereiopods 3 and 4 not greatly lobed
distally; article 2 of pereiopod 5 posteriorly produced to tip of article 5, posterior
margin distally excavate, apex bifurcate, article 3 half length of 4, neither article
lobed; third pleonal epimeron postero-distally rounded; inner ramus of
uropod 3 apically bifid; pereon segment 7 bearing two hard dorsal ridges
enclosing an elongate furrow.
Distribution: Endemic to False Bay and vicinity; apparently limited to cirripede
burrows in encrusting algae, usually on the shells of large gastropods.
104 ANNALS OF THE SOUTH AFRICAN MUSEUM
Ampelisca fusca Stebbing, 1888
Ampelisca fusca Stebbing, 1888: 1052, pl. 105.
Records: 31/16/D to 34/18/0,D and 34/18/FB/T,O, more common at the
deeper stations.
Distribution: Endemic, Mocgambique to South West Africa.
Ampelisca palmata K. H. Barnard, 1916
Ampelisca palmata K. H. Barnard, 1916: 136, pl. 28 (figs 30-31).
Records: 33/17/T, O to 34/18/FB/T, O, abundant in sandy and muddy sub-
strates.
Distribution: Senegal to Mocambique.
Ampelisca spinimana Chevreux, 1887
Ampelisca spinimana: Chevreux & Fage, 1925: 81, fig. 73.
Records: 32/18/T, O to 34/18/FB/T, O, fairly common in sandy areas.
Distribution: Europe, West and southern Africa.
Byblis gaimardi (Kroyer, 1846)
Byblis gaimardi: Mills, 1971: 367-370, figs 6A, 7.
Records: 34/18/D, a single record.
Distribution: Probably cosmopolitan.
Triodos insignis K. H. Barnard, 1916
Triodos insignis K. H. Barnard, 1916: 140, pl. 26 (figs 8-10).
Records: 34/18/D, several records.
Distribution: Endemic, Natal to Cape Peninsula.
Remarks: These records mark the rediscovery of this genus and species,
previously known solely from Barnard’s original two specimens collected off
Natal in 1900.
Family Amphilochidae
Amphilochus neapolitanus Della Valle, 1893
Amphilochus neapolitanus: J. L. Barnard, 1962b: 126, fig. 3.
Records: 34/18/FB/O, a single record.
Distribution: Cosmopolitan in tropical and temperate seas.
Cyproidea ornata Haswell, 1880
Cyproidea ornata: J. L. Barnard, 1972a: 21, figs 4-5.
Records: 33/18/T to 34/18/FB/T, O, a few records.
Distribution: Indo-Pacific, extending to South West Africa.
THE AMPHIPODA OF SOUTHERN AFRICA 105
Gitanopsis pusilla K. H. Barnard, 1916
Gitanopsis pusilla K. H. Barnard, 1916: 144.
Records: 33/18/1, T and 33/17/O to 34/18/FB/I, T, O, fairly common.
Distribution: South Atlantic, southern Indian Ocean.
Hoplopleon australis (K. H. Barnard, 1916)
Peltocoxa australis K. H. Barnard, 1916: 146, pl. 26 (fig. 13).
Records: 33/18/I, a single record.
Diagnosis: Coxae 1 and 2 concealed by greatly enlarged coxae 3 and 4;
gnathopods | and 2 subchelate, article 6 ovate, palm oblique, convex, defined by
a small spine and studded with numerous small denticles, dactyl subequal to
palm, inner margin bearing a comb-like row of strong upstanding teeth;
article 2 of pereiopod 3 linear, that of pereiopods 4 and 5 expanded; pleon
segment 4 elongate with a high medio-dorsal crest along its whole length;
outer ramus of uropods | and 2 the shorter.
Distribution: Endemic, known only from the above record.
Hoplopleon medusarum K. H. Barnard, 1932
Hoplopleon medusarum K. H. Barnard, 1932: 105, fig. 54.
(non) Hoplopleon medusarum: Penrith & Kensley, 1970: 230 (= Cyproidea ornata).
Records: 32/18/T to 34/18/FB/O, a few records.
Distribution: Endemic; Mossel Bay to Lambert’s Bay.
Remarks: The distribution given by Griffiths (1974a) was based on the erroneous
records of Penrith & Kensley; this species has not in fact been found farther
north than Lambert’s Bay.
Family Ampithoidae
Ampithoe africana K. H. Barnard, 1925
Ampithoe africana K. H. Barnard, 1925: 361.
Records: 34/18/FB/T, a single record.
Distribution: Endemic, Natal to False Bay.
Ampithoe falsa K. H. Barnard, 1932
Ampithoe brevipes: K. H. Barnard, 1916: 255, pl. 28 (fig. 34).
Ampithoe falsa: Ruffo, 1969: 57, figs 18-20.
Records: 33/18/T, 34/18/FB/I, a few records.
Distribution: Indian Ocean, South Africa.
106 ANNALS OF THE SOUTH AFRICAN MUSEUM
Ampithoe ramondi (Audouin, 1826)
Ampithoe vaillanti K. H. Barnard, 1916: 253.
Ampithoe ramondi: J. L. Barnard, 19705: 50, figs 18-19.
Records: 29/16/I to 34/18/FB/I, T, fairly common.
Distribution: Circumtropical.
Cymadusa filosa Savigny, 1818
Grubia australis K. H. Barnard, 1916: 258.
Cymadusa australis: K. H. Barnard, 1940: 480.
Cymadusa filosa: J. L. Barnard, 1955: 29, fig. 15.
Records: 29/17/I to 34/18/FB/I, abundant in Langebaan Lagoon, otherwise
uncommon.
Distribution: Cosmopolitan in tropical and temperate seas.
Macropisthopus stebbingi K. H. Barnard, 1916
Macropisthopus stebbingi K. H. Barnard, 1916: 260, pl. 28 (figs 15-17).
Records: 34/18/I, 34/18/FB/I, T, a few records.
Distribution: Endemic, Port Elizabeth to Table Bay.
Family Argissidae
Argissa hamatipes Norman, 1869
Argissa Stebbingi: Chevreux & Fage, 1925: 90, figs 81-82.
Argissa hamatipes: Nagata, 1965: 154, fig. 7. Bousfield, 1973: 121, pl. 20 (fig. 2).
Records: 34/18/FB/O, a few records.
Diagnosis: Accessory flagellum bi-articulate; coxae 1-3 successively smaller,
coxa 4 larger than 1; gnathopods 1 and 2 simple; pereiopods 1 and 2 not
glandular; urosome segments 2 and 3 slightly carinate; telson deeply cleft.
Distribution: Cosmopolitan.
Remarks: It is now generally accepted that A. stebbingi is synonymous with
A. hamatipes (Nagata 1965; J. L. Barnard 1967). The eyes of this species are
highly variable, ranging from absent to well developed with cuticular lenses.
The specimens recorded here, which are the first from this family to be found in
southern Africa, show well-developed eyes each bearing four cuticular lenses.
Family Cheluridae
Chelura terebrans Philippi, 1839
Chelura terebrans: Bousfield, 1973: 207, pl. 69 (fig. 1).
Records: 33/\8/T, a single record.
Distribution: Cosmopolitan, wood-boring.
THE AMPHIPODA OF SOUTHERN AFRICA 107
Family Colomastigidae
Colomastix pusilla Grube, 1864
Colomastix pusilla: J. L. Barnard, 1971: 55, fig. 24.
Records: 34/18/T, O, D, 34/18/FB/T, a few records.
Distribution: Cosmopolitan in tropical and temperate seas.
Family Corophiidae
Aora anomala Schellenberg, 1926
Aora typica forma anomala Schellenberg, 1926a: 372, fig. 59.
Records: 34/18/FB/T, O, a few records.
Diagnosis: Coxa 1 3 not enlarged; article 2 of gnathopod 1 ¢ anteriorly smooth,
articles 2 and 3 not antero-distally lobed, article 4 distally produced into a long
process which extends beyond tip of article 5, article 5 postero-distally produced
into an acute tooth, 6 half as large as 5, palm very short, defined by a small
spine, dactyl serrate; article 6 of gnathopod 2 half length of article 5, palm
convex; uropod 1 with strong interramal spine.
Distribution: Southern Atlantic.
Remarks: Both this species and the two following species have been raised to
specific rank by J. L. Barnard (19725).
Aora gibbula K. H. Barnard, 1932
Aora typica forma gibbula K. H. Barnard, 1932: 220, fig. 135.
Records: 34/18/FB/T, O, numerous records.
Diagnosis: Coxa 1 ¢ greatly enlarged, concealing most of head; gnathopod 1 3
greatly elongate, article 2 produced near its origin into a marginally crenulate
anterior lobe, articles 2 and 3 not antero-distally lobed, 4 distally produced into
a long process extending almost to apex of article 5, article 5 postero-distally
rounded, 6 half as large as 5, palm obscure, dactyl weakly serrate; article 6 of
gnathopod 2 half as long as 5, palm excavate; uropod 1 with strong interramal
spine.
Distribution: Endemic, known only from False Bay.
Aora kergueleni Stebbing, 1888
Aora kergueleni Stebbing, 1888: 1073, pl. 109A, D.
Aora typica (non Kroyer, 1845): K. H. Barnard, 1916, 236; 1940: 478 (partim). Griffiths, 1973:
278; 1974a: 179; 1974b: 255; 1974c: 277.
Records: 29/16/I to 34/18/FB/I, T, O, abundant.
Diagnosis: Coxa 1 g not enlarged; article 2 of gnathopod 1 ¢ anteriorly smooth,
articles 2 and 3 not antero-distally lobed, article 4 distally produced into an
108 ANNALS OF THE SOUTH AFRICAN MUSEUM
elongate process extending to apex of article 5, 5 postero-distally rounded,
6 almost as large as 5, palm short, defined by a spine, dactyl elongate, serrate;
articles 5 and 6 of gnathopod 2 subequal, palm linear; uropod 1 with strong
interramal spine.
Distribution: Kerguelen Island, southern Africa.
Aorcho delgadus J. L. Barnard, 1961
Aorcho delgadus J. L. Barnard, 1961: 114, fig. 80.
Records: 34/18/D, a few records.
Diagnosis: Article 3 of peduncle of antenna | as long as article 1, accessory
flagellum 2-4 articulate; gnathopods subequal; coxae short, hardly serially
touching, pereiopods 1 and 2 glandular; uropods 1 and 2 with large terminal
peduncular spines; rami of uropod 3 subequal, longer than peduncle, outer
ramus with terminal fascicle of setae, inner with small terminal spine; telson
entire, subcircular.
Distribution: Tasman Sea, South Africa.
Remarks: The above records are the first of this species from southern Africa.
The material differs from that of J. L. Barnard (1961) in that the accessory
flagellum of the only specimen possessing a first antenna is 2-articulate, whereas
Barnard described it as 4-articulate.
Cheiriphotis megacheles (Giles, 1885)
Cheiriphotis megacheles: Walker, 1904: 284, pl. 6 (fig. 42). K. H. Barnard, 1937: 167, fig. 14A.
?Cheiriphotis durbanensis K. H. Barnard, 1916: 247. Ruffo, 1969: 55.
Cheiriphotis walkeri Stebbing, 1918: 68, pl. 13.
?Cheiriphotis megacheles forme durbanensis: Ledoyer, 1973: 65, pl. 14A.
Records: 33/17/T, O, D to 34/18/FB/T, O, fairly common.
Distribution: Indo-Pacific.
Remarks: The taxonomic status of this species has historically been somewhat
confused and I do not wish to compound that confusion here by making definite
decisions on synonymy. However, material from southern Africa can be
regarded as falling into two distinct forms. The first, ‘megacheles’ form, common
in the Cape Province, has the palm of gnathopod 2 ¢ transverse and multi-
dentate. This form corresponds to Walker’s C. megacheles and Stebbing’s
C. walkeri as well as K. H. Barnard’s (1937) figure 14B. The second form, found
in Natal and Mogambique, corresponds with that figured by K. H. Barnard
(1937) figure 14A and described by Ledoyer (1973) and can be regarded as the
‘durbanensis’ form. Here the palm of gnathopod 2 3 is oblique and bears three
large teeth, that nearest the finger-hinge being apically notched.
J. L. Barnard (1962a) figures a third form (subsequently assigned to
C. delloyei by Ruffo) and suggests that the various forms of gnathopod 2
represent growth stages. However, mixed populations of the two southern
THE AMPHIPODA OF SOUTHERN AFRICA 109
African forms have not yet been found. Moreover, the ‘durbanensis’ form, the
presumed juvenile, is often larger than the “‘megacheles’ form and juvenile
‘megacheles’ specimens show no tendency to resemble the ‘durbanensis’ form.
This would suggest that the two morphs represent distinct population groups
and should be given at least sub-specific status.
Chevalia aviculae Walker, 1904
Chevalia aviculae: J. L. Barnard, 1971: 88, fig. 42.
Records: 32/18/O to 34/18/T, O, D and 34/18/FB/T, O, numerous records.
Distribution: Cosmopolitan in tropical and temperate seas.
Corophium acherusicum Costa, 1857
Corophium acherusicum: J. L. Barnard, 1971: 59, figs 17, 26. Bousfield, 1973: 201, pl. 62 (fig. 2)-
Records: 33/18/T, 34/18/FB/T, a few records.
Distribution: Cosmopolitan in tropical and temperate seas.
Corophium triaenonyx Stebbing, 1904
Corophium triaenonyx Stebbing, 1904: 25, pl. 6A.
Records: 34/18/FB/T, 34/19/E, a few records.
Distribution: Widespread tropical and subtropical, particularly in brack waters.
Gammaropsis afra Stebbing, 1888
Eurystheus afer: K. H. Barnard, 1916: 249, pl. 28 (fig. 11).
Gammaropsis afra: J. L. Barnard, 1961: 113, fig. 79; 19705: 170, fig. 108.
Records: 31/18/O, 32/16/D, 34/18/FB/O, 35/18/D, a few records.
Distribution: Circumtropical.
Gammaropsis atlantica Stebbing, 1888
Eurystheus atlanticus: Stebbing, 1910a: 461.
Gammaropsis atlantica: J. L. Barnard, 19706: 174, figs 111-113.
Records: 33/18/T to 34/18/FB/T, O and hues D, abundant.
Distribution: Circumtropical.
Gammaropsis holmesi (Stebbing, 1908)
Eurystheus holmesi Stebbing, 19085: 85, pl. 14A. K. H. Barnard, 1955: 95, fig. 48A—D.
Eurystheus semidentatus K. H. Barnard, 1916: 250, pl. 28 (figs 13, 14).
Gammaropsis holmesi: Griffiths, 19745: 244.
Records: 34/18/T, O, 34/18/FB/I, T, O, fairly common.
Distribution: Endemic, Natal to Cape Peninsula.
Gammaropsis longicarpus (Reid, 1951)
Fig. 3
Eurystheus longicarpus Reid, 1951: 259, fig. 50.
Records: 34/18/FB/T, O, a few records.
110 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 3. Gammaropsis longicarpus (Reid, 1951)
Male, 7 mm: A-—lateral aspect; B—gnathopod 1; C—gnathopod 2 (outer aspect);
D—gnathopod 2 (inner aspect); E—third pleonal epimeron ; F-—uropod 3; G—telson.
Female, 6 mm: H—gnathopod 2.
THE AMPHIPODA OF SOUTHERN AFRICA 111
Diagnosis: Eyes quadrate; coxa 1 not strongly produced forwards; articles 5
and 6 of gnathopod 2 3 subequal, article 6 of terminal male with two peg-like
teeth on medial margin and three such teeth along palmar margin (subadult
with palmar teeth but lacking medial teeth); third pleonal epimeron with an
oblique ridge running to a small postero-distal tooth, hind margin strongly
convex; urosome segments | and 2 each with a pair of small subdorsal teeth;
uropod 3 slightly exceeding 1 and 2, rami subequal to peduncle.
Distribution: West and southern Africa.
Remarks: Adult males from the present material are larger and show more
highly developed second gnathopods than those described by Reid (1951).
However, subadults from False Bay agree closely with Reid’s brief description
and on this basis I have associated the two.
Gammaropsis palmoides (K. H. Barnard, 1932)
Eurystheus palmoides K. H. Barnard, 1932: 231, fig. 144; 1955: 96, fig. 48e-g.
Records: 34/18/FB/T, O, numerous records.
Diagnosis: Eyes round, optic lobes short, rounded; article 6 of gnathopod 2 3
much longer than 5, palm oblique, occupying almost entire posterior margin of
hand, defined by an acute tooth and bearing a large medial tooth and two small
distal teeth, dactyl equal to palm, closing within it; pleon segments dorsally
smooth; third pleonal epimeron minutely produced postero-distally; uropod 3
very short, not extending to apex of uropod 2.
Distribution: Endemic to False Bay.
Gammaropsis scissimanus (K. H. Barnard, 1925)
Fig. 4
Eurystheus scissimanus K. H. Barnard, 1925: 361, pl. 34 (fig. 15).
Records: 33/17/O, 34/18/D, 34/18/FB/O, a few records.
Diagnosis: Eyes oval; ocular lobes of head acute; article 6 of gnathopod 2 3
considerably larger than 5, palm transverse, shorter than hind margin, defined
by a small tooth and with a deep semicircular concavity distally, remainder of
palm minutely crenulate; pleon segments dorsally smooth; third pleonal
epimeron postero-distally rounded-quadrate; uropod 3 moderately elongate,
reaching apex of uropod 2.
Distribution: Endemic, False Bay to Saldanha Bay.
Remarks: This species has not been adequately figured before so I have provided
full figures here. Antenna 1 was previously unknown and has an 11-articulate
flagellum with a 5-articulate accessory flagellum. This confirms the correct
placement of G. scissimanus in Gammaropsis rather than Podoceropsis (in which
the accessory flagellum is 0-2 articulate).
112 ANNALS OF THE SOUTH AFRICAN MUSEUM
YI
bah \\
sy)
= a
nM
Fig. 4. Gammaropsis scissimanus (K. H. Barnard, 1925)
Male, 4 mm: A-—lateral aspect; B—gnathopod 1; C—gnathopod 2; D-—uropod 3;
E—telson.
Grandidierella lignorum K. H. Barnard, 1935
Grandidierella lignorum K. H. Barnard, 1935: 300, fig. 14.
Records: 34/19/E, a single record.
Distribution: Endemic, Cape Agulhas to Natal, estuarine.
Grandidierella lutosa K. H. Barnard, 1952
Grandidierella lutosa K. H. Barnard, 1952: 280, fig. 3.
Records: 34/19/E, two records.
Diagnosis: Pereon segments without medio-ventral processes; coxae 1 and 2
i a ee
THE AMPHIPODA OF SOUTHERN AFRICA 3
quadrate, not acutely pointed; article 2 of gnathopod 1 3 widening distally,
article 5 enlarged, two strong teeth on distal margin, article 6 curved, palm
transverse, dactyl hardly exceeding palm; gnathopod 2 ¢ subchelate, much
smaller than gnathopod 1.
Distribution: Endemic to estuaries near Hermanus.
Lemboides acanthiger K. H. Barnard, 1916
Lemboides acanthiger K. H. Barnard, 1916: 239, pl. 28 (figs 7-8).
Records: 34/18/FB/O, a single record.
Distribution: Endemic, Natal to False Bay.
Lemboides afer Stebbing, 1895
Lemboides afer: K. H. Barnard, 1932: 222, fig. 137.
Records: 33/18/T, 34/18/FB/T, O, fairly common.
Distribution: Endemic, False Bay to South West Africa.
Lemboides crenatipalma K. H. Barnard, 1916
Lemboides crenatipalma K. H. Barnard, 1916: 240, pl. 28 (figs 9-10).
Records: 32/18/T, 33/18/T, 33/17/O, a few records.
Distribution: Endemic, Saldanha Bay to South West Africa.
Lembos hirsutipes Stebbing, 1895
Lembos hirsutipes Stebbing, 1895: 207, pl. 8, 9B. Karaman, 1972: 101, figs 1-2.
Records: 32/18/T to 34/18/FB/T, O, a few records.
Diagnosis: Pereon segments ventrally smooth; article 2 of gnathopod 1 ¢
strongly setose posteriorly, palm slightly oblique, a small tooth near finger
hinge separated from long defining tooth by a deep cleft; article 2 of gnathopod
2 § not antero-distally produced, article 5 longer than 6; article 4 of pereiopods 1
and 2 ¢ strongly setose.
Distribution: South Africa to tropical West Africa.
Lembos hypacanthus K. H. Barnard, 1916
Lembos hypacanthus K. H. Barnard, 1916: 237, pl. 28 (figs 5-6).
Records: 33/18/T, 34/18/I, 34/18/FB/I, a few records.
Diagnosis: Pereon segments 3-7 in 3 each bearing a strong medio-ventral
114 ANNALS OF THE SOUTH AFRICAN MUSEUM
spinose process; article 2 of gnathopod 1 3 weakly setose, palm slightly oblique,
defined by a single spine and bearing a minute tooth near finger hinge and a
larger spiniform one near defining angle; article 2 of gnathopod 2 3 with
antero-distal apex produced as a recurved hook, article 6 as long as, but
narrower than, 5; article 4 of pereiopods 1 and 2 ¢ not strongly setose.
Distribution: Endemic, Natal to South West Africa.
Lembos leptocheirus Walker, 1909
Lembos leptocheirus: Walker, 1909: 338, pl. 43 (fig. 7). Schellenberg, 1926a: 373.
Records: 33/17/T, 33/18/T, 34/18/FB/T, a few records.
Diagnosis: Pereon segments ventrally smooth; article 2 of gnathopod 2 3 naked,
palm slightly oblique, convex distally and with a semicircular concavity next to
defining tooth; article 2 of gnathopod 2 3 not produced antero-distally, article 6
much longer than 5; article 4 of pereiopods | and 2 3 sparsely setose.
Photis dolichommata Stebbing, 1910
Photis dolichommata Stebbing, 19105: 609, pl. 55B.
Records: 34/18/FB/T, O, a few records.
Distribution: Australia, South Africa.
Photis longidactylus Griffiths, 1974
Photis longidactylus Griffiths, 1974a: 193, fig. 6.
Records: 32/18/T, O to 34/18/FB/T, O, numerous records.
Distribution: Endemic, False Bay to South West Africa.
Photis longimanus Walker, 1904
Photis longimanus: Rabindranath, 19715: 71, figs 3-4.
(non) Photis longimanus: K. H. Barnard, 1916: 224 (= P. kapapa).
Records: 32/18/T to 34/18/FB/T, O, common, especially in Saldanha Bay and
False Bay.
Distribution: India, Ceylon, southern Africa.
Remarks: The material from Durban Bay attributed to this species by K. H.
Barnard (1916) and subsequently reported by Griffiths (19745) in fact represents
P. kapapa J. L. Barnard. This species is distinguished from P. Jongimanus by the
shape of articles 2 and 6 of the ¢ second gnathopod and the presence of stridu-
lation ridges on the anterior coxae.
THE AMPHIPODA OF SOUTHERN AFRICA is
Photis uncinata K. H. Barnard, 1932
Photis longicaudata: K. H. Barnard, 1916: 243, pl. 28 (fig. 26).
Photis uncinata K. H. Barnard, 1932: 223, fig. 138.
Records: 33/17/T, O to 34/18/FB/T, O, abundant, particularly in False Bay.
Distribution: Endemic to South Africa.
Podoceropsis sophiae Boeck, 1861
Podoceropsis sophiae: Chevreux & Fage, 1925: 316, fig. 324. Reid, 1951: 264, fig. 53. Karaman,
1972: 121, figs 9-11.
Records: 34/18/FB/O, two records.
Diagnosis: Accessory flagellum absent; palm of gnathopod 1 ¢ undefined,
dactyl almost as long as hand, its inner margin serrate; article 5 of gnathopod 2 3
very short, bearing a narrow posterior lobe, article 6 with medial surface
strongly setose, palm oblique, subequal to hind margin, strongly concave
proximally, a large lobe near finger hinge distally divided into two teeth, dactyl
sinuous near base, closing within palm; article 2 of pereiopod 3 posteriorly
produced into a distinctive rectangular projection.
Distribution: Atlantic.
Remarks: This is the first record of this species from southern Africa.
Pseudomegamphopus jassopsis (K. H. Barnard, 1951)
Lembos jassopsis K. H. Barnard 1951, 706, fig. 6.
Pseudomegamphopus jassopsis: Myers 1974: 195, figs 1-4
Records; 32/18/T, 33/17/T, 33/18/T, a few records.
Diagnosis: Article 5 of gnathopod | narrow, triangular, article 6 enormous,
palm oblique, defined by a huge lobe which becomes fleshy and triangular in
terminal male, dactyl bearing long setae on posterior margin; outer ramus of
uropod 3 shorter and broader than inner, bearing a small article 2 terminating
in two long setae.
Distribution: Endemic, the above records (most of which are from Saldanha
Bay) are the only ones to date.
Remarks: This species has been removed from Lembos to Pseudomegamphopus
by Myers 1974, who also supplies a detailed redescription of the species.
Pseudomegamophopus may be distinguished from Lembos by the deeply recessed
head and elongate article 3 of antenna 1.
Siphonoecetes dellavallei Stebbing, 1893
Siphonoecetes dellavallei: Chevreux & Fage, 1925: 361, fig. 369.
Records: 33/18/T, 33/17/T,O to 34/18/FB/T,O and 34/19/E, common,
particularly in False Bay and Saldanha Bay.
Distribution: Mediterranean, southern Africa.
116 ANNALS OF THE SOUTH AFRICAN MUSEUM
Unciolella foveolata K. H. Barnard, 1955
Uniolella foveolata K. H. Barnard, 1955: 97, fig. 49.
Records: 34/18/FB/O, 32/16/D, a few records.
Diagnosis: Integument coarsely pitted; accessory flagellum 4-5 articulate;
pereon segments without ventral spines; gnathopods subequal, subchelate,
medial surfaces of articles 5 and 6 of gnathopod 2 strongly setose; uropods 1
and 2 hardly extending beyond tip of uropod 3, uropod 3 uniramous, ramus with
apical tuft of plumose setae.
Distribution: Endemic, False Bay to Lambert’s Bay.
Unciolella spinosa Griffiths, 1974
Unciolella spinosa Griffiths, 19746: 229, fig. 3.
Records: 33/17/T, O, D to 34/18/FB/T, O, a few records.
Distribution: Endemic, Natal to Saldanha Bay.
Remarks: The discovery of further specimens of this form has shown that, as is
the case in other Corophiidae (e.g. Grandidierella bonnieroides), the medio-
ventral spines of the J pereon may be present or absent within the same species.
This feature should thus not be relied upon during identification. In the present
instance U. spinosa is best distinguished from U. foveolata on the basis of its
elongate first and second uropods, which exceed uropod 3 by about half their
length.
Family Dexaminidae
Atylus granulosus (Walker, 1904)
Atylus granulosus: Ledoyer, 1967: 127, fig. 8.
Records: 32/18/T, O to 34/18/FB/T, O, numerous records.
Distribution: Indian Ocean.
Atylus guttatus (Costa, 1851)
Nototropis guttatus: Chevreux & Fage, 1925: 194, figs 201-203.
Records: 32/18/T, O to 34/18/FB/T, fairly common, especially in Saldanha
Bay.
Distribution: Eastern Atlantic, extending around south coast of South Africa.
Atylus homochir Haswell, 1885
Atylus homochir: Stebbing, 1888: 908-913, pl. 74. J. L. Barnard, 1974: 12, figs 7-9.
Records: 33/17/O, a single record.
Distribution: Australia, South Africa.
THE AMPHIPODA OF SOUTHERN AFRICA 117
Atylus swammerdami (Milne-Edwards, 1830)
Atylus swammerdami: Bousfield, 1973: 131.
Records: 32/18/T, a few records.
Distribution: Atlantic, extending along south coast of South Africa.
Dexamine spiniventris (Costa, 1853)
Dexamine spiniventris: Chevreux & Fage, 1925: 262, figs 271-273.
Records: 33/18/T, a single record.
Diagnosis: Accessory flagellum absent; mandible without palp; maxillipedal
palp 3-articulate; article 2 of pereiopods 3-5 successively wider, article 4
shorter than 5 plus 6; pleon segments 2-3 (sometimes 1-3) each with a medio-
dorsal tooth and a pair of lateral teeth; third pleonal epimeron acutely produced
postero-distally; urosomite 1 with an acute medio-dorsal carina; urosomites 2
and 3 coalesced; telson 60% cleft, reaching almost to apex of uropod 3.
Distribution: Mediterranean, Atlantic.
Guernea rhomba Griffiths, 1974
Guernea laevis (non Chevreux, 1887): K. H. Barnard, 1916: 213.
Guernea rhomba Griffiths, 1974a: 183, fig. 3.
Records: 34/18/I, a single record.
Distribution: Endemic, Cape Town to South West Africa.
Paradexamine pacifica (Thomson, 1879)
Paradexamine pacifica: J. L. Barnard, 19725: 60.
Records: 34/18/FB/T, a single record.
Diagnosis: Article 5 of gnathopods 1-2 about 1,3 times length of article 6,
palms pectinate; pleon segment | dorsally smooth, pleon segments 2 and 3 each
with a medio-dorsal tooth and a pair of lateral teeth; third pleonal epimeron
postero-distally acutely produced; urosomite 1 with a sharp medio-dorsal tooth
and a pair of subdorsal spines; urosomites 2 and 3 fused, bearing two subdorsal
spines on each side; apices of telson cut into 8—10 serrations.
Distribution: Indo-Pacific.
Polycheria atolli Walker, 1905
Polycheria antarctica: Chilton, 1912: 502. K. H. Barnard, 1916: 211.
Polycheria atolli: Ledoyer, 1972a: 205, pl. 27.
Records: 32/18/T to 34/18/FB/T, O, numerous records.
Distribution: Southern oceans, extending to tropical Indian Ocean.
118 ANNALS OF THE SOUTH AFRICAN MUSEUM
Family Eusiridae
Calliopiella michaelseni Schellenberg, 1925
Calliopiella michaelseni: K. H. Barnard, 1940: 451, fig. 24. Griffiths, 1974a: 180.
Records: 33/17/1, T to 34/19/I, common under the shells of Patella spp.
Distribution: Endemic, Cape Agulhas to South West Africa.
Cleonardopsis carinata K. H. Barnard, 1916
Cleonardopsis carinata K. H. Barnard, 1916: 176, pl. 27 (figs 7-9).
Records: 35/18/VD, a single record.
Diagnosis: Accessory flagellum uni-articulate; gnathopods subchelate, article 5
almost as long as 6, lobed posteriorly, palm very oblique, undefined; pereon
segments 6 and 7 and pleon segments | and 2 with medio-dorsal carinae ending
in acute teeth; pleon segment 3 with medio-dorsal carina posteriorly extended
with a hooked process; telson unarmed, 30% cleft.
Distribution: Endemic to deep waters around Cape Peninsula.
Eusiroides monoculoides (Haswell, 1880)
Eusiroides monoculoides: J. L. Barnard, 1964: 221, fig. 1.
Records: 32/18/T to 34/18/FB/T, O, fairly common.
Distribution: Circumtropical.
Paramoera bidentata K. H. Barnard, 1932
Paramoera bidentata K. H. Barnard, 1932: 211, figs 118m, 129.
Records; 32/18/I to 34/18/FB/I and 34/18/T, a few records.
Distribution: Endemic, Still Bay to Liideritz.
Paramoera capensis (Dana, 1853)
Paramoera capensis: K. H. Barnard, 1916: 183-186.
Paramoera schizurus Stebbing, 1918: 66, pl. 10.
Records: 29/16/I to 34/18/FB/T, O and 34/19/I, E, the most abundant species
found in this area at depths of less than 100 m.
Distribution: Southern oceans.
Rhachotropis grimaldi (Chevreux, 1887)
Rhachotropis grimaldii: Stebbing, 1888: 1641. K. H. Barnard, 1916: 179.
Records: 32/17/D, 34/18/D, VD, a few records.
Distribution: Atlantic, extending to Natal.
THE AMPHIPODA OF SOUTHERN AFRICA 119
Rhachotropis kergueleni Stebbing, 1888
Rhachotropis kergueleni Stebbing, 1888: 955, pl. 85.
Records: 34/18/VD, a single record.
Diagnosis: Pereon dorsally smooth; pleon segments | and 2 each with an acute-
tipped medio-dorsal carina and a pair of acutely tipped subdorsal ridges; pleon
segment 3 with a single carina, 4 with a single carina terminating in a large
tooth; pleonal epimera 1 and 2 postero-distally rounded, 3 postero-distally
serrate; hind margin of article 2 of pereiopods 3-5 serrate, postero-distal corner
acute, slightly produced.
Distribution: Kerguelen Island, South Africa.
Rhachotropis paeneglaber K. H. Barnard, 1916
Rhachotropis paeneglaber K. H. Barnard, 1916: 181, pl. 27 (fig. 10).
Records: 34/18/D, VD, two records.
Diagnosis: Pereon dorsally smooth; pleon segment | with an obscure median
keel, 2 with three keels each terminating in an acute tooth, 3 with three distally
untoothed keels; pleon segment 4 with median keel ending acutely, subdorsal
keels not extending as far as posterior margin of segment; pleonal epimera 1
and 2 postero-distally rounded, 3 postero-distally serrate; hind margin of
article 2 of pereiopods 3 and 4 smooth, that of pereiopod 5 faintly serrate,
postero-distal corner rounded.
Distribution: Endemic, the above records are the only ones to date.
Rhachotropis palporum Stebbing, 1908
Rhachotropis palporum Stebbing, 1908a: 194, pl. 28.
Records: 34/18/D—VD, a single record.
Diagnosis: Pereon dorsally smooth; pleon segments 1-3 each with three keels,
all ending in elongate acute teeth; pleon segment 4 with a minute medio-dorsal
tooth; pleonal epimera 1 and 2 with a single acute tooth at centre of posterior
margin, postero-distally rounded, third pleonal epimeron similar but postero-
distally acutely produced; article 2 of pereiopods 3-5 with a large blunt process
arising from centre of posterior margin, otherwise smooth.
Distribution: North and South Atlantic.
Family Gammaridae
Ceradocus rubromaculatus (Stimpson, 1855)
Ceradocus rubromaculatus: J. L. Barnard, 1972a: 220, fig. 129.
Records: 29/16/I to 34/18/FB/I, T,O, and 35/18/D, abundant, particularly
intertidally and in shallow rocky areas.
Distribution: Indo-Pacific, extending to South West Africa.
120 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 5. Cheirocratus inermis Ledoyer, 1968
Male, 4 mm: A-—lateral aspect; B—accessory flagellum; C—articles 5-7 of gnathopod 2.
Female, 3,5 mm: D-—articles 6 and 7 of gnathopod 1; E—articles 6 and 7 of gnathopod 2;
F—uropod 3; G—telson.
THE AMPHIPODA OF SOUTHERN AFRICA 12)
Cheirocratus inermis Ledoyer, 1968
Fig. 5
Cheirocratus inermis Ledoyer, 1968: 36, pl. 12.
Records: 34/18/FB/O, a few records.
Diagnosis: Antenna 1 shorter than peduncle of antenna 2, accessory flagellum
2-articulate; body dorsally smooth; gnathopod 1 simple; gnathopod 2 9 simple;
gnathopod 2 ¢ strongly subchelate, article 6 elongate, palm occupying almost
whole hind margin of hand, distally cut into three teeth, dactyl sinuous,
exceeding length of hand; uropod 3 greatly exceeding 1 and 2, rami equal, the
inner uni-articulate; telson cleft nearly to base.
Distribution: Madagascar, South Africa.
Remarks: This is the first record of a male of this species and only the second
time the species has been found. The female agrees closely with Ledoyer’s
original description, while the male shows powerful second gnathopods, a
feature unusual for this genus. C. inermis is the only member of the genus
Cheirocratus in which the pereon is dorsally smooth.
Elasmopus affinis Della Valle, 1893
Elasmopus affinis: Sars, 1895: 521, pl. 183.
Records: 33/17/¥, 33/18/I, T, fairly common.
Distribution: Mediterranean, Atlantic, southern Indian Ocean.
Elasmopus japonicus Stephensen, 1932
Elasmopus japonicus: Sivaprakasam, 1968: 278, figs 3-S.
Records: 34/18/FB/I, a few records.
Distribution: Indo-Pacific, extending to South West Africa.
Elasmopus pectenicrus Bate, 1862
Elasmopus pectenicrus: J. L. Barnard, 19705: 125, figs 73-74.
Records: 34/18/FB/I, a single record.
Distribution: Cosmopolitan in tropical and temperate seas.
Eriopisa epistomata Griffiths, 1974
Eriopisa epistomata Griffiths, 1974a: 186, fig. 4.
Records: 34/18/FB/O, a single record.
Distribution: Endemic, Port Elizabeth to South West Africa.
122 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 6. Jerbarnia mecochira Croker, 1971
Male, 6 mm: A—antenna 2; B—mandibular palp; C—maxilla 1; D—maxilliped;
E—gnathopod 1; F—gnathopod 2; G—uropod 3; H—telson.
THE AMPHIPODA OF SOUTHERN AFRICA 123
Eriopisella capensis (K. H. Barnard, 1916)
Eriopisa capensis K. H. Barnard, 1916: 187, pl. 27 (figs 16-19).
Records: 32/17/D, 32/16/D to 34/18/D, a few records.
Distribution: Endemic to south and west coasts of South Africa.
Jerbarnia mecochira Croker, 1971
Fig. 6
Jerbarnia mecochira Croker, 1971: 382-386, figs 1-2.
Records: 34/18/FB/T, O, a few records.
Diagnosis: Accessory flagellum multi-articulate; article 3 of antenna 2 with a
posterior setose lobe; maxillipedal palp 3-articulate; gnathopod 2 extremely
long, article 3 elongate; coxae 5—7 and pleonal epimera 1-3 each with a spine at
postero-distal corner; pleon segments 1-3 dorsally cut into five teeth, a seta in
each concavity; pleon segments 4 and 5 with two posterior teeth, 6 with two
dorsal spines; uropod 3 greatly exceeding 1 and 2, outer ramus minutely
bi-articulate or not; telson quadrate, 50% cleft.
Distribution: Eniwetok Atoll, South Africa.
Remarks: As shown by figure 6 the present material conforms closely to that
described by Croker from the Pacific. The specimen figured is considerably
larger than Croker’s (6 mm as against 4,5 mm) but is apparently less mature,
having a relatively shorter article 3 of gnathopod 2 and less highly developed
coxae.
Maera boecki (Haswell, 1879)
Elasmopus boeckii: K. H. Barnard, 1916: 199, pl. 27 (figs 13-14).
Maera boeckii: K. H. Barnard, 1940: 460.
Records: 34/18/FB/T, O, a few records.
Distribution: Australia, South Africa.
Maera bruzeli Stebbing, 1888
Fig. 7
Meera bruzelii Stebbing, 1888: 1014, pl. 97.
Maera mastersi (non Haswell, 1880): Griffiths, 1974c: 290.
Records: 33/17/O, 34/18/D, 34/18/FB/T, O, 34/19/?, a few records.
Distribution: Endemic, Port Elizabeth to Saldanha Bay.
Remarks: This species was originally described only from the female and when
samples containing only males were recovered by Griffiths (1974c) these were
erroneously identified as M. mastersi. However, the present samples contain both
males and females and clearly show the two to represent the same species. The
124 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 7. Maera bruzeli Stebbing, 1888
Male, 14mm: A—mandible; B—gnathopod 1; C—articles 5-7 of gnathopod 2 (inner aspect);
D—pereiopod 5; E—pleonal epimera; F—telson; G—urosome.
THE AMPHIPODA OF SOUTHERN AFRICA 125
male M. bruzeli is distinguished by the unusual second gnathopod, in which the
inner surface of the palm is covered by a dense pad of short tightly-packed setae
(Fig. 7C). 3
(The record of M. mastersi from Mocambique by K. H. Barnard (1916), is
of the true mastersi.)
Maera emarginata sp. nov.
Fig. 8
Description of male (5 mm): Head as long as first two pereon segments, lacking
lateral cephalic notch, eyes absent; antenna | as long as pereon, articles 1 and 2
subequal, much longer than 3, flagellum subequal to peduncle, 16-articulate,
accessory flagellum (Fig. 8B) of two long articles and one short one; antenna 2
much shorter than 1, gland cone conspicuous, flagellum 7-articulate; mandible
(Fig. 8C) with tridentate incisor, lacinia mobilis of four strong teeth, spine row
of four spines, molar powerful, palp 3-articulate, article 2 the longest; lower lip
without inner lobes; lobes of maxillae 1 and 2 setose only terminally; palp of
maxilliped 4-articulate.
Gnathopod 1 considerably smaller than 2, articles 5 and 6 subequal,
densely setose posteriorly, palm oblique, subequal to hind margin, but not
clearly defined from it, palmar margin finely pectinate with submarginal rows
of small spines; gnathopod 2 strongly subchelate, article 6 considerably larger
than 5, palm oblique, subequal to hind margin, defined by two spines and
bearing three equally spaced teeth (Fig. 8D), dactyl subequal to palm; pereiopods
1 and 2 slender, 2 somewhat the shorter; (pereiopods 3 and 4 missing); pereiopod
5 elongate, extending to apex of uropods 1 and 2, article 2 not greatly expanded
posteriorly, posterior margin cut into seven moderate serrations, articles 4 and 5
with strong posterior spines.
Pleonal epimera 1 and 2 smoothly rounded, 3 with a slight postero-distal
tooth; none of pleon segments dorsally dentate or carinate; uropod | (Fig. 8E)
slightly exceeding 2, peduncle with a strong proximal spine and a large distal
spine on medial surface, outer ramus 70% length of inner, both strongly
spinose apically; peduncle of uropod 2 (Fig. 8F) with five dorsal spines, outer
ramus slightly the shorter, bearing four dorsal and three apical spines, inner
ramus with a row of ten upright dorsal spines and five apical spines; uropod 3
(Fig. 8G) greatly exceeding 1 and 2, rami subequal, outer ramus with one large
and two small dorsal spines, also three apical spines, inner ramus with four
ventral fascicles of spines and four spines apically; telson (Fig. 8H) not more
than 25% cleft, a single blunt spine within a shallow notch at apex of each lobe.
Holotype: SAM-A13473, male, 5 mm.
Type-locality: 34°17'S/18°29’E, 15 February 1965, depth 27 m, substrate shelly
sand.
Relationships: The telson of M. emarginata sp. nov. 1s highly unusual in that it
126 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 8. Maera emarginata sp. nov.
Male, 5mm: A-—lateral aspect; B—accessory flagellum; C—mandible; D-—articles 5-7 of
gnathopod 2; E, F, G—uropods 1, 2, 3; H—telson.
THE AMPHIPODA OF SOUTHERN AFRICA 7
is hardly cleft, most other species having telsons at least 50% cleft. M. eugeniae
Schellenberg also has an incompletely cleft telson but here the apices are
strongly divergent. Moreover, in M. euginiae the palm of gnathopod 2 ¢ is
undefined.
Material: Three specimens from the type locality.
Maera grossimana (Montagu, 1808)
Maera grossimana: Chevreux & Fage, 1925: 239, figs 248, 250. Karaman & Ruffo, 1971:
114, figs 1-3.
Records: 33/17/T, two records.
Distribution: Mediterranean, Atlantic.
Maera hamigera (Haswell, 1880)
Meera hamigera: J. L. Barnard, 1965: 507, fig. 16. Karaman & Ruffo, 1971: 152, figs 21-23.
Records: 32/17/D, 33/18/I, T to 34/18/FB/I, T and 34/19/I, a few records.
Distribution: Indo-Pacific, extending around west coast of South Africa.
Remarks: The form of gnathopod 2 ¢ in this species is highly variable. Males in
the present collection show a series of regular teeth along the distal portion of
the palm, as figured by K. H. Barnard (1916), while females agree closely with
the form figured by Karaman & Ruffo (1971).
Maera hirondellei Chevreux, 1910
Meera hirondellei: K. H. Barnard, 1916: 194. Chevreux & Fage, 1925: 241, fig. 252. Karaman
& Ruffo, 1971: 122, figs 4-7.
Records: 33/18/I, 33/17/D, two records.
Distribution: Mediterranean, Atlantic.
Remarks: This species can be distinguished from M. grossimana by virtue of its
elongate third uropod, which exceeds the apex of uropod 2 by about half the
length of its rami.
Maera inaequipes (Costa, 1851)
Meera inaequipes: Karaman & Ruffo, 1971: 143, figs 17-20.
Records: 29/16/I to 34/18/FB/T, O and 34/18/D, abundant.
Distribution: Cosmopolitan in tropical and temperate seas.
128 ANNALS OF THE SOUTH AFRICAN MUSEUM
Maera komma sp. nov.
Fig. 9
Description of male (7 mm): Head without cheek notch, eyes large (half height
of head), composed of 20-30 separate ocelli; antenna | as long as pereon plus
pleon, article 2 longer than 1, 3 short, flagellum 26-articulate, accessory
flagellum of three elongate articles; antenna 2 extending 25 % along flagellum of
antenna 1, flagellum 12-articulate; mandible (Fig. 9A) with large strongly
triturative molar, article 2 of palp slightly longer than 3.
Coxa 1 (Fig. 9B) strongly produced forwards, lower margin with one
anterior tooth and two posterior teeth, remaining coxae quadrate; articles 5
and 6 of gnathopod | subequal, palm (Fig. 9C) oblique, defined by a single
small spine, palmar margin finely pectinate throughout, dactyl subequal to
palm; gnathopod 2 (Fig. 9D) strongly subchelate, article 2 anteriorly smooth,
3 not lobed, 5 strongly setose posteriorly, 6 elongate, widest at its base, posterior
margin strongly setose proximally, concave distally, palm not defined, dactyl
half length of hand, strongly curved so that when closed an oval gap remains
between finger and palm; pereiopods 1 and 2 slender; article 2 of pereiopods 3—5
elongate-oval, posteriorly serrate, distal articles linear. .
First pleonal epimeron very small, postero-distally quadrate; second
pleonal epimeron much larger than first, postero-distally slightly produced;
third pleonal epimeron postero-distally slightly produced, posterior margin.
with four small serrations, lower margin with five anterior spines; uropods 1
and 2 extending equally, slightly exceeding apex of peduncle of uropod 3;
uropod | (Fig. 9G) with four pairs of dorsal spines and a strong spine on medial
distal margin, inner ramus dorsally smooth, outer with three dorsal spines;
outer ramus of uropod 2 marginally shorter than inner; uropod 3 (Fig. 9H)
large, rami broad, laminar, margins finely pectinate, weakly spinose; telson
(Fig. 91) 80% cleft, each lobe dorsally bearing two proximal setae and a
subterminal seta in a small notch plus a minute terminal seta.
Female: Similar to the male except for shorter third uropods and smaller
gnathopod 2 (Fig. 9J) which has an oblique palm defined by four strong spines
and minutely pectinate distally.
Holotype: SAM-A13477, male, 7 mm.
Type-locality: 34°05’S/17°45’E, 8 February 1963, depth 142 m, substrate dark
green mud.
Relationships: M. komma sp. nov. lies close to M. thrixa sp. nov. (below) and
M. knudseni Reid, 1951. However, it may be distinguished from these species by
the shape of the palm and dactyl of gnathopod 2 3 and, in the case of M. thrixa,
by the absence of an anterior keel on article 2 of gnathopod 2.
Material: 32/17/D, 34/18/D, two records.
THE AMPHIPODA OF SOUTHERN AFRICA 129
Fig. 9. Maera komma sp. nov.
Male, 7mm: A—mandible; B—coxa 1; C—articles 6 and 7 of gnathopod 1; D—gnathopod 2;
E—dactyl of pereiopod 5; F—third pleonal epimeron; G, H—uropods 1, 3; I— telson.
Female, 7mm: J—articles 6 and 7 of gnathopod 2.
130 ANNALS OF THE SOUTH AFRICAN MUSEUM
Meera thrixa sp. nov.
Fig. 10
Description of male (9 mm): Head slightly exceeding length of first pereon
segment, with a small cheek notch, eyes small, round, black; antenna 1 as long
as pereon, article | slightly shorter than 2, 3 very short, flagellum 30-articulate,
accessory flagellum 4-articulate; antenna 2 reaching 30% along flagellum of
antenna 1, article 4 considerably longer than 5, flagellum 11-articulate; mandible
with strong triturative molar, article 1 of palp not produced distally (Fig. 10C),
articles 2 and 3 subequal.
Coxa 1 not strongly produced forwards, lower margin faintly serrate,
remaining coxae sub-quadrate; gnathopod 1 (Fig. 10A) with articles 5 and 6
subequal, palm oblique, defined by two small spines, minutely pectinate
throughout, dactyl smooth, subequal to palm; gnathopod 2 (Fig. 10B) power-
fully subchelate, anterior margin of article 2 distally produced into a triangular
keel, article 3 with an anterior pellucid lobe, article 5 strongly setose posteriorly,
article 6 elongate, anterior margin setose, palm not distinct from hind margin,
setose, dactyl closely appressed to palm, half length of hand; pereiopods | and 2
slender; pereiopod 3 short, extending only to tip of article 4 of pereiopod 4;
article 2 of pereiopods 4 and 5 elongate-oval, posteriorly serrate, none of distal
articles strongly expanded.
First and second pleonal epimera with oblique ridge running to minutely
produced postero-distal corner; third pleonal epimeron (Fig. 10E) acutely
produced postero-distally, posterior margin with two faint serrations, lower
Margin not serrate, anteriorly bearing five small spines; urosomites dorsally
smooth; uropods 1 and 2 extending to tip of peduncle of uropod 3, rami
subequal, weakly spinose dorsally and apically; uropod 3 large, rami elongate-
oval (Fig. 10F), weakly spinose; telson (Fig. 10G) longer than peduncle of
uropod 3, 80% cleft, each lobe with two dorsal setae, a sub-apical spine in a
small notch and two minute apical setae.
Female: Similar to male except for smaller second gnathopod (Fig. 10H) which
has smooth article 2 and oblique palm lined with strong spines.
Holotype: SAM-A13230, male, 9 mm.
Type-locality: 34°18'S/18°48’E, 18 February 1965, depth 51 m, substrate coarse
shelly sand and rock.
Relationships: This species falls into a group typified by M. knudseni Reid and
M. othonis (Milne-Edwards). The anterior keel on article 2 of gnathopod 2 is
not, however, found in other members of the group. This keel appears in
specimens of about 7 mm and in fully adult males (9-12 mm) becomes extremely
prominent, reaching a maximum width about 1,5 times that of the body of
article 2 of gnathopod 2.
Material: 34/18/FB/T, O, several records.
THE AMPHIPODA OF SOUTHERN AFRICA ott
Ze
amainlyy
/
Fig. 10. Maera thrixa n. sp.
Male, 9 mm: A—gnathopod 1; B—gnathopod 2; C—mandibular palp; D-—dactyl of
pereiopod 5; E-—third pleonal epimeron; F—uropod 3; G-—telson. Female, 10 mm:
H-—articles 6 and 7 of gnathopod 2.
132 ANNALS OF THE SOUTH AFRICAN MUSEUM
Maera vagans K. H. Barnard, 1940
Elasmopus laevis K. H. Barnard, 1916: 200, pl. 27 (fig. 15).
Maera vagans K. H. Barnard, 1940: 459.
Records: 31/17/I to 34/18/FB/T, O, fairly common.
Distribution: Endemic, False Bay to Liideritz.
Megaluropus agilis Hoek, 1889
Phylloropus capensis K. H. Barnard, 1932: 146, figs 84-85.
Megaluropus agilis: Pillai, 1957: 50, fig. 10.
Records: 33/18/1, T, 34/18/FB/T, O, a few records.
Distribution: Europe, India. South Africa.
Megaluropus namaquaeensis Schellenberg, 1953
Megaluropus namaquaeensis Schellenberg, 1953: 117, fig. 5.
Records: 32/18/T, O to 34/18/FB/T, O, abundant.
Distribution: Endemic, Natal to South West Africa.
Melita machaera K. H. Barnard, 1955
Melita machaera K, H. Barnard, 1955: 90-92, fig. 45.
Records: 33/18/I, T, 34/18/FB/T, O, a few records.
Distribution: Endemic, Plettenberg Bay to Saldanha Bay.
Melita mucronata sp. nov.
Fig. 11
Description of female (4,5 mm): Head equal to first two pereon segments, eyes
round, brown (as preserved in 70% alconol); antenna 1 almost as long as body,
article 1 slightly shorter than 2, 3 short, flagellum of 29 elongate articles,
accessory flagellum of three long articles and one short article; antenna 2
shorter than 1, peduncle longer than that of antenna 1, flagellum 11-articulate;
mandible (Fig. 11B) with incisor of three strong teeth, lacinia mobilis bifurcate,
spine row of nine spines, molar quadrate, articles 2 and 3 of palp subequal, much
longer than 1; inner plate of maxilla 2 strongly setose medially.
Coxae 1-4 quadrate, subequal, postero-distal corners minutely toothed;
gnathopod 1 with articles 5 and 6 subequal, palm oblique, dactyl equal to palm;
gnathopod 2 (Fig. 11C) larger than 1, article 4 postero-distally acutely produced,
5 slightly shorter than 6, palm oblique, defined by a single spine, dactyl slightly
longer than palm, inner margin faintly crenulate; pereiopods 1 and 2 slender,
dactyl with strong anterior accessory cusp; (pereiopods 3-5 missing).
THE AMPHIPODA OF SOUTHERN AFRICA 133
LA) A
Fig. 11. Melita mucronata sp. nov.
Female, 4,5 mm: A—lateral aspect; B—mandible; C—gnathopod 2; D—dorsal margins of
pleon segments 1-5; E—uropod 3; F—telson.
134 ANNALS OF THE SOUTH AFRICAN MUSEUM
Pleonal tooth formula 4: 6:5: 2:2 (Fig. 11D) with a suggestion of a
third pair of teeth on pleon segment 1; pleonal epimera 1-3 acutely produced
postero-distally, distal margins with 1, 2 and 3 spines respectively; uropods 1
and 2 extending equally, peduncles and rami dorsally spinose, apices of rami
truncated, surrounded by rings of slender spines; uropod 3 (Fig. 11E) greatly
exceeding 1 and 2, inner ramus about 10% length of outer, bearing a single
apical spine, outer ramus with small spiniform article 2, article 1 with three
lateral fascicles of spines on each side and a terminal fascicle of spines sur-
rounding base of article 2; telson (Fig. 11F) cleft to base, each lobe with a medio
dorsal and two lateral strong spines, apices of lobes acute.
Holotype: SAM-—A13233, female, 4,5 mm, unique.
Type-locality: 34°21'S/18°41’E, 22 February 1965, depth 85 m, substrate fine
green mud.
Relationships: The pleonal tooth formula of this species is diagnostic. Similar
formulae are those of M. dentata (Kroyer) and M. gladiosa Bate, which have a
central tooth on pleon segments 4 and 5, and M. pallida Sars which lacks a
central tooth on pleon segment 3.
Melita orgasmos K. H. Barnard, 1940
Melita orgasmos K. H. Barnard, 1940: 454. Sivaprakasam, 1966: 114, fig. 12k—m.
Records: 29/16/I to 34/19/I, a few records, mostly intertidal.
Distribution: India, southern Africa.
Melita subchelata (Schellenberg, 1925)
Melita fresnelii var. subchelata Schellenberg, 1925: 153. K. H. Barnard, 1932: 211, fig. 130.
Records; 32/18/T, 33/17/T, O, a few records.
Distribution: Endemic, Saldanha Bay to South West Africa.
Melita zeylanica Stebbing, 1904
Melita zeylanica: J. L. Barnard, 1972a: 235, figs 139-141.
Records: 33/18/I and 31/18/E to 34/19/E, common in estuaries and areas of
lowered salinity.
Distribution: Indo-Pacific, a brack-water species.
Parelasmopus suluensis (Dana, 1853)
Parelasmopus suluensis: Stebbing, 1888: 1029, pl. 100. Ledoyer, 1972a: 233, pls 48-49. Griffiths,
1974¢: 292.
Records: 34/18/FB/T, a single record.
Distribution: Indo-Pacific.
THE AMPHIPODA OF SOUTHERN AFRICA 135
Quadrivisio aviceps (K. H. Barnard, 1940)
Ceradocus aviceps K. H. Barnard, 1940: 456, fig. 25.
Records: 34/19/E, a few records.
Diagnosis: None of pereon or pleon segments dorsally dentate; gnathopod 2 ¢
with article 6 elongate-oval, distally scabrous, dactyl very short and strongly
hooked, closing inwards against inner surface of hand; pleonal epimera 1-3
weakly crenulate posteriorly, minutely produced postero-distally; rami of
uropod 3 elongate-oval, apically rounded, setose ventrally and distally; telson
cleft to base, apices notched with a single spine within notch.
Distribution: Endemic to estuaries in the Hermanus district.
Remarks: A re-examination of the existing material of this species has shown
article 1 of the mandibular palp to be distally rounded and article 3 to be almost
as long as 2. These characters are inconsistent with the definition of Ceradocus
and C. aviceps should be removed from that genus to Quadrivisio Stebbing,
1907. Members of this genus resemble ‘C’. aviceps closely in the foliaceous
condition of the rami of uropod 3 and in their estuarine habits. As originally
defined Quadrivisio showed two pairs of eyes, but this has been shown to be a
variable character (Schoemaker 1933), the eyes in fact developing from oval to
dumbbell-shaped and then separating into two portions. In the present species
the eyes are distinctly bilobed although the two halves are still connected even
in the largest known specimens.
Q. aviceps differs from other species of Quadrivisio in the unusual condition
of gnathopod 2 J, in which the stunted dactyl closes medially against the inner
surface of the hand, rather than longitudinally against the palmar margin.
Family Haustoriidae
Bathyporeia sp.
Bathyporeia gracilis (non Sars, 1891): K. H. Barnard, 1951: 704.
Bathyporeia sp.: Vader, 1970: 161.
Records: 32/18/T, O to 34/18/FB/I, T, O, numerous records.
Distribution: Endemic, eastern Cape Province to South West Africa.
Remarks: These specimens were sent to Dr Wim Vader, who is to describe a
new species from them.
Cunicus profundus Griffiths, 1974
Cunicus profundus Griffiths, 1974c: 293-297, figs 8-9.
Records: 32/18/T, 34/18/D, 34/18/FB/O, abundant locally.
Distribution: Endemic, Port Elizabeth to Lambert’s Bay.
Remarks: This species appears to be an extremely efficient burrower. Although
136 ANNALS OF THE SOUTH AFRICAN MUSEUM
rarely collected by conventional methods it has been recovered in considerable
numbers by a diver-operated suction-sampler which extracts sand samples of
60 cm depth and more.
Urothoe coxalis Griffiths, 1974
Urothoe coxalis Griffiths, 19745: 239, fig. 5.
Records: 32/17/O, a single record.
Distribution: Endemic, Natal to Saldanha Bay.
Remarks: U. coxalis is closely related to U. cuspis Imbach (1967), a description
of which I have only recently been able to obtain. The two species may be
distinguished by the presence of an antero-distal cusp on coxa 2 of U. cuspis
and the elongate posteriorly-directed process of coxa 4 of U. coxalis.
Urothoe elegans Bate, 1857
Urothoe elegans: Chevreux & Fage, 1925: 101, fig. 95.
Records: 32/17/D, 33/18/I, T to 34/18/FB/I, T, O, numerous records.
Distribution: Atlantic, Indian Ocean.
Urothoe grimaldi Chevreux, 1895
Urothoe grimaldii: Chevreux & Fage, 1925: 99, fig. 93. K. H. Barnard, 1955: 84, fig. 41B.
Records: 32/18/T, O, 32/17/D to 34/18/FB/T, O, one of the most abundant
species found in sandy sediments.
Distribution: Mediterranean, Atlantic, Indian Ocean.
Urothoe pinnata K. H. Barnard, 1955
Urothoe pinnata K. H. Barnard, 1955: 86, fig. 42.
Records: 34/18/FB/T, O, fairly common.
Distribution: Endemic, Natal to False Bay.
Urothoe pulchella (Costa, 1853)
Urothoe pulchella: Chevreux & Fage, 1925: 99, fig. 92. K. H. Barnard, 1955: 83, fig. 41A.
Records: 32/18/T, 32/17/D to 34/18/FB/T, O, numerous records.
Distribution: Mediterranean, Atlantic, South Africa.
THE AMPHIPODA OF SOUTHERN AFRICA 137
Urothoe tumorosa Griffiths, 1974
Urothoe tumorosa Griffiths, 19746: 241, fig. 6.
Records: 34/18/FB/T, O, a few records.
Distribution: Endemic, Natal to False Bay.
Family Ischyroceridae
Cerapus tubularis Say, 1818
Cerapus tubularis: J. L. Barnard, 1962a: 61, figs 27-28. Bousfield, 1973: 197, pl. 60 (fig. 1).
Records: 33/18/O, 34/18/FB/T, O, a few records.
Distribution: Cosmopolitan in tropical and temperate seas.
Ericthonius brasiliensis (Dana, 1853)
Ericthonius brasiliensis: J. L. Barnard, 1971: 61, fig. 17E. Bousfield, 1973: 195, pl. 59 (fig. 2).
Records: 31/18/E, 32/18/T to 34/18/FB/T, O, numerous records.
Distribution: Cosmopolitan in tropical and temperate seas.
Isaeopsis tenax K. H. Barnard, 1916
Tsaeopsis tenax K. H. Barnard, 1916: 267, pl. 28 (figs 19-21).
Records: 33/18/1, 34/18/I, two records.
Diagnosis: Accessory flagellum of a single elongate article; coxae serially
touching, 1 about 75% length of 2, 6 half as long as 5; articles 5 and 6 of
gnathopod 1 subequal; gnathopod 2 3, palm lacking defining tooth (resembling
that of Ischyrocerus anguipes); pereiopods prehensile, article 2 widened.
Distribution: Endemic, known only from the above records.
Ischyrocerus anguipes Kroyer, 1835
Ischyrocerus anguipes: Bousfield, 1973: 192, pl. 58 (fig. 1).
Records: 33/18/T, O to 34/18/FB/I, T, O, fairly common.
Distribution: Cosmopolitan in tropical and temperate seas.
Ischyrocerus carinatus K. H. Barnard, 1916
Ischyrocerus carinatus K. H. Barnard, 1916: 266, pl. 28 (fig. 18).
Records: 33/18/I, 34/18/FB/I, T, a few records.
Distribution: Endemic, False Bay to South West Africa.
138 ANNALS OF THE SOUTH AFRICAN MUSEUM
Ischyrocerus ctenophorus Schellenberg, 1953
Fig. 12
Ischyrocerus ctenophorus Schellenberg, 1953: 121, fig. 7.
Records: 34/18/FB/I, a single record.
Distribution: Endemic, False Bay to Lideritz.
Remarks: The above record is the first of a male of this species and only the
second of a female. The female is identical with that figured by Schellenberg.
Gnathopod 1 of the male is similar to that of the female while gnathopod 2
(Fig. 12D) is much larger and of typical Ischyrocerus type. I. ctenophorus is
distinguished from other members of the genus by the large dorsal teeth on the
outer ramus of uropod 3 (Fig. 12H) and by the comb-like rows of setae along
the lateral margins of the dactyls of gnathopods | and 2 in both sexes.
Ischyrocerus gorgoniae K. H. Barnard, 1940
Ischyrocerus gorgoniae K. H. Barnard, 1940: 481, fig. 35.
Records: 34/18/FB/T, a few records.
Diagnosis: Pereon segments 1-6 each medio-dorsally carinate; article 2 of
gnathopod 2 ¢ long and slender, margins entire, article 6 elongate-oval, palm
straight with a narrow elongate tooth near hinge, dactyl laterally compressed;
inner ramus of uropod 3 slightly the shorter, bearing a small apical spine, outer
ramus with a large apical spine and two large dorsal teeth. 7
Distribution: Endemic, known only from the above records.
Jassa falcata Montagu, 1808
Jassa falcata: Sexton & Reid, 1951: 30-47, pls 4-30. Bousfield, 1973: 190, pl. 58 (fig. 2).
Records: 32/18/T, O, D to 34/18/FB/I, T, numerous records.
Distribution: Cosmopolitan.
Parajassa chikoa Griffiths, 1974
Parajassa chikoa Griffiths, 1974c: 300, fig. 11.
Records: 34/18/FB/O, a few records.
Distribution: Endemic, Cape St. Francis to False Bay.
Ventojassa frequens (Chilton, 1883)
Jassa frequens: Schellenberg, 1953: 119, fig. 6. Griffiths, 1974a: 196.
Ventojassa frequens: J. L. Barnard, 19726: 135, figs 74-75.
Records: 34/18/FB/T, O, two records.
Distribution: New Zealand, southern Africa.
Remarks: The genus Ventojassa, as created by J. L. Barnard (19725), differs
THE AMPHIPODA OF SOUTHERN AFRICA 139
Fig. 12. Ischyrocerus ctenophorus Schellenberg, 1953
Male, 4 mm: A-—accessory flagellum; B—coxae 1-7; C—gnathopod 1 with tip of dactyl
enlarged; D-—gnathopod 2. Female, 3,5 mm: E, F—gnathopods 1, 2; G—uropod 3;
H—rami of uropod 3 enlarged.
140 ANNALS OF THE SOUTH AFRICAN MUSEUM
from Jassa in the form of uropod 3, the dorsal ornamentation of the outer ramus
consisting of one or more wire-like setae, rather than the one to three sharp
reverted cusps typical of Jassa.
Family Leucothoidae
Leucothoe ctenochir K. H. Barnard, 1925
Leucothoe ctenochir K. H. Barnard, 1925: 342, pl. 34 (fig. 8).
Records: 34/18/T, 34/18/FB/T, O, a few records.
Distribution: Endemic, Durban to Cape Peninsula.
Leucothoe dentitelson Chevreux, 1925
Leucothoe dentitelson Chevreux, 1925: 297, figs 7-8. Reid, 1951: 227, fig. 25.
Records; 33/18/T, two records.
Diagnosis: Antennae short, extending only to pereon segment 2; article 7 of
gnathopod 1 about 20% length of article 6; gnathopod 2 quite different in the
two sexes, that of 3 large, article 5 apically serrate, palm oblique, nodulose
proximally, distally bearing four fairly large flat-topped teeth; gnathopod 2 2
smaller, article 5 apically crenulate, 6 antero-distally produced over base of
dactyl, process acute, setose, palm slightly oblique, finely nodulose, defining
angle almost rectangular, dactyl very stout; second pleonal epimeron acutely
produced; third pleonal epimeron with a notch above a postero-distal tooth,
posterior margin sinuous.
Distribution: Senegal, west coast of South Africa.
Remarks: This species has not previously been recorded from southern Africa.
Leucothoe dolichoceras K. H. Barnard, 1916
Leucothoe dolichoceras K. H. Barnard, 1916: 157, pl. 26 (fig. 14); 1925: 343.
Records: 33/17/T, O, 33/18/FB/T, O, a few records.
Distribution: Endemic, Natal to Saldanha Bay.
Leucothoe richiardi Lessona, 1865
Leucothoe richiardi: Sivaprakasam, 1967: 385, fig. 2.
Records: 32/18/T, 32/17/D to 34/18/FB/I, T, O, numerous records.
Distribution: Mediterranean, India, South Africa.
Leucothoe spinicarpa (Abildgaard, 1789)
Leucothoe spinicarpa: Sivaprakasam, 1967: 384, fig. 1.
Records: 33/18/T to 34/18/FB/T, O and 35/18/D, a few records.
Distribution: Cosmopolitan.
THE AMPHIPODA OF SOUTHERN AFRICA 141
Family Liljeborgiidae
Liljeborgia dubia (Haswell, 1880)
Eusirus dubius Haswell, 1880: 331, pl. 30 (fig. 3).
Records: 33/17/O, 34/18/FB/T, O, a few records.
Distribution: Indo-Pacific.
Liljeborgia epistomata K. H. Barnard, 1932
Liljeborgia epistomata K. H. Barnard, 1932: 114, fig. 83; 1955: 89, fig. 44.
Records: 31/18/O, 32/17/D to 34/18/FB/T, O, numerous records.
Distribution: Endemic, Natal to Lambert’s Bay.
Liljeborgia kinahani (Bate, 1862)
Liljeborgia kinahani: Chevreux & Fage, 1925: 157, fig. 157.
Records: 34/18/D, 34/18/FB/T, O, a few records.
Distribution: North Atlantic, South Africa.
Liljeborgia palmata Griffiths, 1974
Liljeborgia palmata Griffiths 1974c: 304, fig. 12.
Records: 32/17/D, 32/16/D to 34/18/D, a few records.
Distribution: Endemic, Still Bay to Lambert’s Bay.
Liljeborgia proxima Chevreux, 1908
Liljeborgia proxima Chevreux, 1908: 475, figs 4-5.
Records: 34/18/FB/T, a single record.
Diagnosis: Palm of gnathopod 2 ¢ smoothly convex; pleon segments 1, 2 and 4
each bearing a single medio-dorsal tooth, segments 3 and 5 dorsally smooth.
Distribution: Atlantic coast of Africa.
Listriella lindae Griffiths, 1974
Listriella lindae Griffiths, 1974a: 197, fig. 7.
Records: 32/18/O, a single record.
Distribution: Endemic, Lambert’s Bay to South West Africa.
Listriella saldanha sp. nov.
Fig. 13
Description of male (8 mm): Head as long as 1,5 pereon segments, eyes round,
black; antenna 1 half as long as 2, flagellum 9-articulate, accessory flagellum
(Fig. 13B) 3-articulate; antenna 2 extending to pereon segment 5, flagellum
8-articulate; mandible (Fig. 13C) with incisor cut into five strong teeth, lacinia
mobilis of five teeth, spine row of nine spines, molar represented by five setae,
142 ANNALS OF THE SOUTH AFRICAN MUSEUM
- g 4 é VA
Na ae
Fig. 13. Listriella saldanha sp. nov.
Male, 8 mm: A-—lIateral aspect; B—accessory flagellum; C—mandible; D—articles 4-7 of
gnathopod 2; E, F, G—uropods 1, 2, 3; H—telson.
THE AMPHIPODA OF SOUTHERN AFRICA 143
palp 3-articulate, article 3 lined with plumose setae; inner plate of maxilla |
terminating in a single seta, outer plate bearing about eight pectinate spines,
palp bi-articulate, with two terminal spines and a double row of setae along
distal margin.
Coxae 1-4 without distal teeth, | slightly produced forwards; gnathopod 1
subchelate, considerably smaller than 2, palm slightly oblique, evenly convex,
lined by alternating setae and spines; gnathopod 2 (Fig. 13D) powerful, palm
sinuous, almost transverse, defined by a large spine and with a row of short spines
on either side of its setose margin, dactyl indistinctly serrate throughout, excavate
near its base; pereiopods | and 2 slender, pereiopods 3-5 successively longer.
Pleonal epimera I-—3 each postero-distally produced into a small tooth,
posterior margins entire; urosome segments dorsally smooth; uropods | and 2
(Fig. 13E—F) extending equally; peduncle of uropod | with a medio-dorsal row
of about 20 close-packed spines, outer margin with about six more widely spaced
spines, outer ramus slightly the shorter, both rami dorsally and apically spinose,
longest terminal spine of each ramus striated; uropod 3 (Fig. 13G) reaching well
beyond apices of | and 2, outer ramus with a minute second article; telson
(Fig. 13H) cleft to base, each lobe truncated distally with a medio-distal tooth
and three strong terminal setae.
Female: The accessory flagellum of the only 2 recorded to date is 4-articulate, as
opposed to 3-articulate in the 3. The palm of gnathopod 2 9 is transverse but
not sinuous as in the J, also the telsonic apices each bear four rather than
three setae.
Holotype: SAM-—A13227, male, 8 mm.
Type-locality: 33°01'S/17°58’E, 2 May 1972, depth 12 m, substrate sand.
Relationships: All representatives of this genus with the exception of the three
southern African species, L. /indae Griffiths, L. sinuosa Griffiths and L. saldanha
sp. nov. have bi-articulate accessory flagellae. L. saldanha can be distinguished
from the other two local species by virtue of the transverse palm of the second
gnathopod and the shape of the third pleonal epimeron.
Material: 33/18/T, 34/18/FB/O, a few records.
Family Lysianassidae
Acidostoma obesum (Bate, 1862)
Acidostoma obesum: K. H. Barnard, 1925: 322. Chevreux & Fage, 1925: 32, fig. 9.
Records: 33/17/D, a single record.
Diagnosis: Antenna 1 3 very stout; mouthparts forming a conical bundle;
mandibular palp attached proximal to the weak molar; gnathopod 1 simple;
gnathopod 2 lacking dactyl; pereiopods 3-5 very stout, articles 4 and 5 as wide
as long; uropod 3 very short, extending only to middle of rami of uropod 2;
telson 50° cleft.
Distribution: North and South Atlantic.
144 ANNALS OF THE SOUTH AFRICAN MUSEUM
Amaryllis macrophthalma Haswell, 1880
Amaryllis macrophthalma: J. L. Barnard, 1972a: 262-269, figs 156-158.
Records: 29/16/I, 32/18/T, 32/17/O, D to 34/18/FB/I, T,O and 34/19/I, one
of the most abundant species in the area.
Distribution: Southern hemisphere.
Aristias symbiotica K. H. Barnard, 1916
Aristias symbiotica K. H. Barnard, 1916: 122.
Records: 32/18/I, T, 32/17/D to 34/18/I, D, 34/18/FB/T, O, fairly common.
Distribution: Endemic, Mocambique to South West Africa.
Cyphocaris anonyx Boeck, 1871
Cyphocaris anonyx: Schellenberg 1926b: 210, figs 2b, 5a—b, pl. 5 (fig. 2).
Records: 34/16/VD, a single record.
Diagnosis: Pereon segment 1 overhanging head, not produced into a ‘horn’;
article 2 of pereiopod 3 postero-distally produced to tip of article 5, both margins
of process strongly serrate.
Distribution: Cosmopolitan, bathypelagic.
Cyphocaris challengeri Stebbing, 1888
Cyphocaris challengeri Stebbing, 1888: 661, pl. 17.
Records: 34/16/VD, a single record.
Distribution: Cosmopolitan, bathypelagic.
Cyphocaris richardi Chevreux, 1905
Cyphocaris richardi: J. L. Barnard, 1954: 53, pls 2-3.
Records: 33/15/VD, 34/16/VD, two records.
Diagnosis: Pereon segment | of adult overhanging head and anteriorly produced
into an elongate, forward-projecting ‘horn’; article 2 of pereiopod 3 not greatly
produced postero-distally, posterior margin with about 10 strong serrations.
Distribution: Cosmopolitan, bathypelagic.
Euonyx biscayensis Chevreux, 1908
Euonyx biscayensis: K. H. Barnard, 1916: 110. J. L. Barnard, 1961: 34, fig. 4.
Records: 35/18/VD, a single record.
Diagnosis: Article 1 of antenna 1 slender, linear; eyes weak or absent; gnatho-
pod | distinctly chelate, chela about 40 % length of hand; article 6 of gnathopod 2
half as long as 5; third pleonal epimeron quadrate; dorsal surface of pleon
segment 4 with basal depression and rounded distal hump.
Distribution: Mediterranean, eastern Atlantic, East Africa.
THE AMPHIPODA OF SOUTHERN AFRICA 145
Euonyx conicurus K. H. Barnard, 1955
Euonyx conicurus K. H. Barnard, 1955: 80, fig. 38.
Records: 34/18/FB/T, a few records.
Distribution: Endemic, Port Elizabeth to False Bay.
Eurythenes obesus (Chevreux, 1905)
Katius obesus: K. H. Barnard, 1932: 56-58, fig. 21, pl. 1 (fig. 1).
Eurythenes gryllus: K. H. Barnard, 1940: 440.
Eurythenes obesus: J. L. Barnard, 1961: 38, fig. 8.
Records: 33—34/15-16/VD, a single record.
Diagnosis: Coxa 1 much smaller than 2 and partially concealed by it;
gnathopod 1 subchelate, article 6 twice as long as 5; gnathopod 2 subchelate,
article 6 half as long as 5; article 2 of pereiopod 3 very small, 20% or less of
length of limb; dactyl of pereiopods 3—5 large, over 50% length of article 6.
Distribution: Cosmopolitan in tropical and temperate seas.
Hippomedon longimanus (Stebbing, 1888) new synonymy
Platamon longimanus Stebbing, 1888: 643, pl. 13.
Hippomedon longimanus: K. H. Barnard, 1916: 125.
Tryphosa africana K. H. Barnard, 1955: 81.
Tryphosella africana: Griffiths, 1974c: 315.
Records: 32/18/T, O, 32/17/D to 34/18/FB/T, O, one of the most abundant
species in this area.
Distribution: Atlantic, South African east coast.
Remarks: In the past both K. H. Barnard and I have designated specimens to
either Hippomedon longimanus or Tryphosella ( =Tryphosa) africana on the
basis of the length of the postero-distal tooth of the third pleonal epimeron.
However, as more material has become available, this situation has become
untenable, since a continuous range of variation in the size of this process has
been found to occur. Other than this feature, the description of Tryphosa africana
in Barnard (1955) applies equally well to Hippomedon longimanus. Unfortunately
Barnard failed to allocate a holotype of Tryphosa africana, but he based his
brief description on specimens in my possession and which are apparently
identical with those of Hippomedon longimanus from the same area.
The composite species formed by the amalgamation of the above forms is
definitely a Hippomedon rather than a Tryphosella since the mandibular molar
is ridged and unsetulose, the dactyl of gnathopod 1 simple, the inner ramus of
uropod 2 unconstricted and the branchiae complete with accessory lobes.
Hippomedon normalis (K. H. Barnard, 1955) new synonymy
Tryphosa normalis K. H. Barnard, 1955: 80, fig. 39.
Tryphosella normalis: Griffiths, 1974a: 201; 19746: 249; 1974c: 315.
Hippomedon rotundipleura Ledoyer, 1973: 75, pl. 19.
Records: 33/18/1, T, O, to 34/18/FB/T, O, numerous records.
146 ANNALS OF THE SOUTH AFRICAN MUSEUM
Distribution: Madagascar, South Africa.
Remarks: Barnard’s incorrect placement of ‘Tryphosa africana’ (above) has
prompted a re-examination of the taxonomic status of his other ‘Tryphosa’—
T. normalis. This species also appears to be a Hippomedon for the same reasons
as given for ‘Tryphosa africana’.
A survey of the literature of Hippomedon has shown that the same species
described by Barnard as Tryphosa normalis has subsequently been redescribed as
a Hippomedon—H. rotundipleura—by Ledoyer (1973). His figures (which are of
a 2, not a 3) agree closely with females in my possession (gnathopods | and 2 are
transposed in Ledoyer’s figures), the only notable difference being that my
specimens have an accessory flagellum of five articles and a few fine setae along
the dorsal margins of the rami uropod 3. Male specimens differ from females in
the length of antenna |, which is usually as long as the body, and in the more
strongly setose margins of the rami of uropod 3.
Hippomedon onconotus (Stebbing, 1908)
Tryphosa onconotus Stebbing, 1908b: 65, pl. 35.
Hippomedon onconotus: J. L. Barnard, 1962c: 29.
Records: 32/17/D, 32/16/D to 34/18/FB/T, O, 35/18/VD, numerous records.
Distribution: Endemic, Natal to Lambert’s Bay.
Lepidepecreum clypeatum Chevreux, 1888
Fig. 14
Lepidepecreum clypeatum: Chevreux, 1900: 28, pl. 4 (fig. 2). Chevreux & Fage, 1925: 63, fig. 52.
Records; 34/18/FB/O, a single record.
Diagnosis: Articles 1 and 2 of antenna 1 dorsally carinate and produced
(Fig. 14A), accessory flagellum absent; eyes lacking; article 2 of pereiopod 5
greatly elongate (Fig. 14G), postero-distally produced to tip of article 5;
pereon segments dorsally smooth; pleon segment 3 postero-dorsally raised into
an acute tooth, segment 4 with a triangular dorsal carina; third pleonal epimeron
with an oblique ridge running to acutely produced postero-distal corner.
Distribution: North Atlantic, South Africa.
Remarks: The above record is the first of a male of this species and the first of
L. clypeatum from the Southern hemisphere. As can be seen from the figure the
material agrees closely with the females described and figured by Chevreux
(1900) and Chevreux & Fage (1925) except for such obviously sexually dimorphic
characters as antenna 2 and uropod 3.
Lepidepecreum clypodentatum J. L. Barnard, 1962
Lepidepecreum clypodentatum J. L. Barnard, 1962c: 27, figs 9-10.
Records; 33/17/D, a single record.
Diagnosis: Article 1 of antenna 1 dorsally carinate and produced, accessory
THE AMPHIPODA OF SOUTHERN AFRICA 147
Fig. 14. Lepidepecreum clypeatum Chevreux, 1888
Male, 5 mm: A-—antenna 1; B—antenna 2; C—mandible with molar enlarged;
D—maxilla 1; E—gnathopod 1 with palm enlarged; F—gnathopod 2 with articles 6 and 7
enlarged; G—pereiopod 5; H—profile of pleon segments 3-5; I, J—uropods 1,3; K—telson.
148 ANNALS OF THE SOUTH AFRICAN MUSEUM
flagellum of two or three minute articles; eyes absent; article 2 of pereiopod 5
greatly elongate, postero-distally produced to tip of article 5; pereon segments
dorsally ridged; pleon segments 1-3 each postero-dorsally produced into an
acute tooth, segment 4 with an erect hook-like dorsal process; third pleonal
epimeron postero-distally quadrate, without oblique ridge.
Distribution: West coast of South Africa.
Remarks: The above record is the first of a female of this species and the only
record other than that of J. L. Barnard (1962c). The single specimen differs from
Barnard’s male only in regard to sexual characters of antenna 1 and uropod 3.
Lepidepecreum twalae Griffiths, 1974
Lepidepecreum twalae Griffiths, 1974c: 310, fig. 14.
Records: 32/17/D, a single record.
Distribution: Endemic, Mossel Bay to Lambert’s Bay.
Lysianassa ceratina (Walker, 1889)
Lysianassa cubensis: K. H. Barnard, 1916: 120.
Lysianassa ceratina: Chevreux & Fage, 1925: 42, fig. 23.
Records: 29/16/I, 32/18/I,T,O to 34/18/FB/I, T, O, where it is one of the
most abundant shallow-water species.
Distribution: Atlantic, Indian Ocean, Mediterranean.
Lysianassa variegata (Stimpson, 1855)
Lysianassa variegata: Stebbing, 1888: 682, pl. 23.
Records: 32/18/O, D to 34/18/FB/T, O, abundant.
Distribution: Africa south of the equator.
Microlysias
The genus Microlysias was originally instituted by Stebbing (1918) for his
M. xenoceras from Durban Bay. Although his description was fully figured he
failed to illustrate structure of the epistome and upper lip. K. H. Barnard (1937)
subsequently erected a second species, M. indica, said to differ from M. xenoceras
in the shape of the epistome and upper lip. However, Barnard, who did not have
access to Stebbing’s types, based his concept of the mouth parts of M. xenoceras
on specimens he himself collected and identified (reported K. H. Barnard 1940),
Griffiths (1973) later identified and figured further specimens of ‘M. indica’,
using Barnard’s (1937) figures as a basis for identification.
A recent examination of Barnard’s (1940) ‘M. xenoceras’ from Plettenberg
Bay by the author has revealed that they in fact represent Orchomene plicata and
not Microlysias. As a result of this error the fabric of Barnard’s argument for
the creation of M. indica collapses, since its erection was based on comparison
with Orchomene mistaken to be M. xenoceras. It thus appears that all existing
THE AMPHIPODA OF SOUTHERN AFRICA 149
records of M. indica are referable to M. xenoceras, the profile of the epistome
and upper lip of which is figured as M. indica by Barnard (1937). Microlysias is
best distinguished from Orchomene by the structure of antenna 2 3, both genera
having branchiae pleated on both sides and article 4 of pereiopods 3—5 expanded.
Orchomene plicata (Schellenberg, 1925)
Orchomenopsis chilensis Schellenberg, 1925: 119, fig. 3. K. H. Barnard, 1925: 330.
Microlysias xenoceras (non Stebbing, 1918): K. H. Barnard, 1940: 441.
Orchomenella plicata: K. H. Barnard, 1940: 440.
Records: 29/16/I, 32/18/T, O to 34/18/FB/I, T, O, abundant.
Distribution: Cosmopolitan.
Remarks: For discussion of synonymy see Microlysias above.
Phoxostoma algoense K. H. Barnard, 1925
Phoxostoma algoense K. H. Barnard, 1925: 323, pl. 34 (fig. 2).
Records: 33/18/T. 34/18/FB/O, a few records.
Distribution: Endemic, Algoa Bay to Table Bay.
Procyphocaris induratus (K. H. Barnard, 1925) new synonymy
Uristes induratus K. H. Barnard, 1925: 333, pl. 34 (fig. 3).
Procyphocaris primata J. L. Barnard, 1961: 49, fig. 18.
Remarks: This species does not strictly fall into the scope of the present paper,
since it is restricted to depths exceeding 1 000 m. However, I have included it
here since its taxonomy is in need of revision.
In his original description of ‘Uristes’ induratus K. H. Barnard (1925)
failed to appreciate the taxonomic significance of the reduced first and second
coxae, an error brought to light by J. L. Barnard (1962c) who rejected the species
from a revised list of Uristes species. J. L. Barnard had, however, inadvertently
already redescribed the same species as the type of a new genus (Procyphocaris
primata J. L. Barnard, 1961). K. H. Barnard’s type specimen agrees almost
exactly with J. L. Barnard’s figures, the only differences being in the posterior
margin of article 2 of pereiopod 3, which is smooth in the South African specimen
and bears three weak serrations in the Australian one. (K. H. Barnard’s state-
ment that the flagellum of antenna 2 is bi-articulate is incorrect, there are nine
segments in an unbroken flagellum.)
The holotype of the resultant species, correctly known as Procyphocaris
induratus, is South African Museum number SAM-—A4545.
Schisturella adversicola (K. H. Barnard, 1925)
Lakota adversicola K. H. Barnard, 1925: 327.
Chironesimus adversicola: Schellenberg, 1926a; 219, fig. 13. J. L. Barnard, 1962c: 22, fig. 2.
Schisturella adversicola: J. L. Barnard, 1967: 71.
Records: 35/18/VD, a single record.
150 ANNALS OF THE SOUTH AFRICAN MUSEUM
Diagnosis: Eyes absent, coxa | reduced, partially concealed by 2; gnathopod |
subchelate, palm almost transverse; third pleonal epimeron postero-distally
acutely produced; inner ramus of uropod 2 strongly constricted medially; outer
ramus of uropod 3 bi-articulate; telson about 50% cleft.
Distribution: Endemic to deep waters off the Cape Peninsula.
Socarnes septimus sp. nov.
Fig. 15
Description of male (2,5 mm): Head hardly longer than pereon segment 1,
lateral lobes strongly produced, apically subacute, eyes (Fig. 15A) composed of
about 30 separate red ocelli, antenna 1 about as long as head plus two pereon
segments, peduncle very stout, article 1 greatly exceeding 2 plus 3 (Fig. 15B),
flagellum 5-articulate, bearing aesthetascs, accessory flagellum 3-articulate;
antenna 2 (Fig. 15C) only about 1,5 times length of 1, article 5 of peduncle half
as long as 4, flagellum 6-articulate; upper lip produced lobately beyond epistome;
mandible (Fig. 15D) with weakly toothed incisor, spine row of two small spines,
molar large, not strongly ridged, palp 3-articulate, article 2 naked, more than
twice length of 1, article 3 with two terminal setae; maxilla 1 (Fig. 15E) with
bi-articulate palp, palp article 2 distally serrate, bearing only a single minute
apical seta, outer plate with seven strong serrate spines; maxilliped (Fig. 15F),
outer plate with seven medial nodules, palp 4-articulate, greatly exceeding outer
plate.
Pereon segments with a few scattered dorsal setules; coxae 1-4 considerably
longer than their body segments, | not concealed by 2, 4 excavate posteriorly;
gnathopod | (Fig. 15G) simple, articles 5 and 6 subequal, 6 medially constricted,
dactyl with accessory tooth; gnathopod 2 (Fig. 15H) chelate, article 6 about
60% length of 5; article 2 of pereiopods 3—5 subcircular, posterior margin
crenulate with minute setae, anterior margin lined with strong spines, article 4
almost as wide as long, lobed posteriorly, 5—7 slender.
Pleonal epimera | and 3 postero-distally rounded, 2 produced into a minute
tooth; uropod 1 slightly exceeding 2 and 3, peduncle with a single dorsal spine,
outer ramus 70% length of inner, bearing two dorsal spines and one apical
spine, inner ramus with one dorsal and one apical spine; peduncle of uropod 2
(Fig. 15I) latero-dorsally keeled, outer ramus with a single dorsal spine, inner
ramus with two dorsal spines; uropod 3 (Fig. 15J) with outer margin of peduncle
dorsally keeled, outer ramus with small article 2, inner ramus smooth, telson
twice as long as broad, 60° cleft, lobes divergent (Fig. 15K), each terminating
in two small setae, one plumose.
Female: Similar to male, ovigerous at 2 mm, usually carries only a single
enormous ovum.
Holotype: SAM-A13466, male, 2,5 mm.
Type-locality: 34°12'S/18°37'E, 15 May 1961, depth 48 m, substrate khaki sand
and shell.
THE AMPHIPODA OF SOUTHERN AFRICA isi
Fig. 15. Socarnes septimus sp. nov.
Male, 2,5 mm: A—head; B—antenna 1; C—antenna 2; D—mandible; E—maxilla 1;
F—maxilliped; G—gnathopod 1; H-—articles 5-7 of gnathopod 2; I—uropod 2 (outer
aspect); J—uropod 3 (medial aspect); K—telson.
52 ANNALS OF THE SCUTH AFRICAN MUSEUM
Relationships: The minute size of this species alone distinguishes it from most
other members of the genus. In addition the rounded third pleonal epimeron
differs from those of Socarnes unidentatus Schellenberg and S. bidentata (Bate).
S. dissimulantia Imbach lacks eyes, while other members of the genus have
considerably more slender and longer first antenna than S. septimus sp. nov.
Material: 34/18/FB/T, O, fairly common.
Socarnopsis crenulata Chevreux, 1910
Socarnopsis crenulata: K. H. Barnard, 1916: 124. Chevreux & Fage, 1925: 49, figs 31-32.
Records: 33/17/O, 34/18/FB/O, a few records.
Distribution: Mediterranean, Atlantic, South Africa.
Stomacontion capense K. H. Barnard, 1916
Stomacontion capense K. H. Barnard, 1916: 109, pl. 28 (figs 27-28); 1937: 140, fig. 1.
Records: 32/17/D, 34/18/FB/O, two records.
Diagnosis: Head almost entirely obscured by triangular coxa 1; gnathopod 1
slender, simple, dactyl minute; dactyl of gnathopod 2 inserted at middle of distal
margin of article 6; article 2 of pereiopod 5 greatly lobed posteriorly, lobe
extending to tip of article 5 and postero-distally rectangular; pleon segment 4
with rounded dorsal protuberance; uropod 3 without rami or rami vestigial.
Distribution: Endemic, False Bay to Lambert’s Bay.
Stomacontion prionoplax Monod, 1937
Stomacontion prionoplax Monod, 1937: 6, figs 1-6. Griffiths, 1974c: 313, fig. 15.
Records: 34/18/FB/O, a single record.
Distribution: Suez Canal, South Africa.
Trischizostoma paucispinosum K. H. Barnard, 1916
Trischizostoma paucispinosum K. H. Barnard, 1916: 107, pl. 26 (fig. 1).
Records: 34/18/D, two records.
Diagnosis: Rostrum inconspicuous; coxa | almost completely concealed by 2;
gnathopod | powerful, article 6 truncated oval, palm subequal to hind margin,
sparsely spinose, dactyl smooth, slightly exceeding palm, articles 4 and 5 of
pereiopods 3-5 slender; telson 50% cleft.
Distribution: Endemic to deep waters off the Cape Peninsula.
Trischizostoma remipes Stebbing, 1908
Trischizostoma remipes Stebbing, 1908b: 61, pi. 34. K. H. Barnard, 1925: 321.
Records: 30/16/D to 34/18/FB/O, fairly common.
Distribution: Endemic, Natal to Hondeklip Bay.
THE AMPHIPODA OF SOUTHERN AFRICA 153
Trischizostoma serratum K. H. Barnard, 1925
Trischizostoma serratum K. H. Barnard, 1925: 320, pl. 34 (fig. 1).
Records: 34/18/FB/T.
Distribution: Endemic, Natal to False Bay.
Uristes sulcus Griffiths, 1974
Uristes sulcus Griffiths, 1974c: 315; fig. 16.
Records: 34/18/FB/O, a few records.
Distribution: Endemic, Plettenberg Bay to False Bay.
Family Ochlesidae
Ochlesis lenticulosus K. H. Barnard, 1940
Ochlesis lenticulosus K. H. Barnard, 1940: 447, fig. 23.
Records: 34/18/FB/T, O, a few records.
Distribution: Endemic, Natal to False Bay.
Family Oedicerotidae
Halicreion ovalitelson K. H. Barnard, 1916
Halicreion ovalitelson K. H. Barnard, 1916: 165, pl. 27 (fig. 4).
Records: 34/18/VD, a single record.
Diagnosis: Rostrum extending to apex of article 1 of antenna 1; eyes absent;
article 5 of gnathopods 1 and 2 produced posteriorly into a lobe guarding
posterior margin of article 6; uropod 2 extending only to tip of peduncle of
uropod 3; telson elongate-oval; entire.
Distribution: Endemic, the above record is the only one to date.
Monoculodopsis longimana Ledoyer, 1973
Monoculodopsis longimana Ledoyer, 1973: 79, figs 22-24.
Records: 32/18/T, O to 34/18/FB/T, O, numerous records.
Diagnosis: Primary cutting edge of mandible projecting, toothed, molar
represented by a small hump bearing a few spines; inner lobes of lower lip
separate; gnathopods dissimilar, both wide posterior lobe of article 5 fully
guarding posterior margin of article 6; article 6 of gnathopod | broad, expanding
distally, palm as long as hind margin, evenly convex, defined by a single spine;
article 6 of gnathopod 2 four times as long as broad, palm oblique, convex,
defined by a small spine; uropod 2 reaching apex of uropod 3; telson apically
emarginate.
Distribution: Madagascar, South Africa.
154 ANNALS OF THE SOUTH AFRICAN MUSEUM
Remarks: This species has not previously been recorded from South Africa. The
above specimens differ from these of Ledoyer (1973) only in that the mandibular
molar is represented by four spines rather than two (gnathopods 1 and 2 are
transposed in Ledoyer’s figure 22).
Oediceroides cinderella Stebbing, 1888
Oediceroides cinderella Stebbing, 1888: 850, pls 62-63.
Records; 32/18/T, 32/16/D, two records.
Diagnosis: Eyes present; rostrum not strongly deflexed, tapering evenly to an
acute point just beyond tip of article 1 of antenna 1; gnathopods similar,
article 5 about half length of 6, produced posteriorly as a rounded lobe pro-
jecting at right angles, not protecting posterior margin of article 6, palm
oblique, convex.
Distribution: Falkland Islands, west coast of South Africa.
Perioculodes longimanus (Bate & Westwood, 1868)
Perioculodes longimanus: Chevreux & Fage, 1925: 162, figs 163-164. Ledoyer, 19725: 775-781,
figs 2-3.
Records: 32/18/T, O, 32/17/D to 34/18/FB/T, O, abundant.
Distribution: Mediterranean, Atlantic, Indian Ocean.
Perioculodes pallidus sp. nov.
Fig. 16
Description of female (2,5 mm): Head about as long as two pereon segments,
rostrum extending to apex of article 1 of antenna 1, eyes absent; antenna 1
about 30% length of pereon, articles 1-3 subequal, flagellum 4-articulate;
accessory flagellum absent; antenna 2 marginally longer than 1, flagellum
4-articulate; mandible (Fig. 16B) with incisor not strongly toothed, molar
represented by a spinose hump, palp 3-articulate, article 2 the longest, article 3
with three strong terminal setae; lower lip without inner lobes; maxilla 1
(Fig. 16C) with bi-articulate palp terminally bearing six setae, outer plate with
seven strong apical spines; inner plate of maxilla 2 with four setae, outer plate
with six (Fig. 16D); inner plate of maxilliped with two long apical setae, outer
plate reaching to centre of article 2 of 4-articulate palp.
Coxae 1-4 successively larger, 4 with a strong seta at postero-distal corner;
gnathopods subchelate, similar, article 5 posteriorly guarding article 6; article 6
of gnathopod | (Fig. 16F) shorter and stouter than that of gnathopod 2
(Fig. 16G), palm defined by a small spine and with submarginal rows of minute
setae, dactyl slightly exceeding palm, constricted just before apex, pereiopods |
and 2 strongly setose, dactyl small (Fig. 16H), surrounded by setae; pereiopod 3
much smaller than 4, article 2 as wide as long, 5 with two strong anterior spines;
article 2 of pereiopod 4 (Fig. 161) strongly lobed posteriorly, anteriorly and
THE AMPHIPODA OF SOUTHERN AFRICA 155
Fig. 16. Perioculodes pallidus sp. nov.
Female, 2,5 mm: A-—lateral aspect; B—mandible; C—maxilla 1; D—maxilla 2;
E—maxilliped; F—articles 5-7 of gnathopod 1; G—articles 5-7 of gnathopod 2; H—articles
6 and 7 of pereiopod 1; I—pereiopod 4; J—uropod 3. Male, 2,5 mm: K—uropods 2 and 3
and telson.
156 ANNALS OF THE SOUTH AFRICAN MUSEUM
medially strongly setose, articles 4-6 strongly setose on both margins; articles 5-7
of pereiopod 5 missing but articles 2-4 much smaller than those of pereiopod 4.
Pleonal epimera 1-3 smoothly rounded; uropod | extending beyond 2 and 3,
rami smooth, lanceolate; uropod 2 slightly exceeding 3, peduncle with two small
distal spines, rami smooth; uropod 3 very short, outer ramus 60% length of
inner, both smooth (Fig. 16J); telson entire, with two small apical concavities,
each containing two setae, on either side of a small medial bump.
Male: Flagellum of antenna | hirsute; flagellum of antenna 2 7-articulate, palm
of gnathopod 2 transverse; article 2 of pereiopods 3 and 4 not as strongly lobed
as in 2; uropod 3 considerably longer (Fig. 16K) than that of 9.
Holotype: SAM-A13232, ovigerous female, 2,5 mm.
Type-locality: 34°13’S/18°31'E, 22 May 1961, depth 39 m, substrate sand.
Relationships: Perioculodes pallidus sp. nov. differs from other members of the
genus by reason of its large coxae, reduced third (and fifth?) pereiopods, short
antennae in both sexes and naked uropods.
Remarks: The most notable feature of the holotype is the small size of the
proximal articles of pereiopod 5, indicating a probable total length considerably
less than that of pereiopod 4. This would be in conflict with the normal condition
in the genus, or indeed the family Oedicerotidae, which characteristically has
pereiopod 5 larger than 4. However, pereiopod 5 was found only on one side of
one of the individuals collected, being lost in all other cases—a characteristic
often associated with great length. Thus, until the condition of pereiopod 5 is
verified by undamaged specimens, I have placed this species in the genus
Perioculodes on the basis of other characters, particularly the structure of the
gnathopods which in the @ are practically identical with those of P. aequimanus
(Kossmann). Should the reduced condition of pereiopod 5 be confirmed by
further material this would be strong evidence for the creation of a new genus.
Material: 34/18/O, two records.
Synchelidium tenuimanus Norman, 1871
Synchelidium tenuimanus: Oldevig, 1933: 131, fig. 2.
Records: 34/18/FB/D, 34/18/D, a few records.
Diagnosis: Rostrum strong, reaching almost to apex of article 1 of antenna 1;
eyes of moderate size; gnathopod 2 extremely slender and elongate, the chela
occupying only about 15% of its length.
Distribution: Atlantic, South Africa.
Westwoodilla manta Griffiths, 1974
Westwoodilla manta Griffiths, 1974c: 318, fig. 17.
Records: 31/18/O, D to 34/18/FB/D, fairly common.
Distribution: Endemic, Plettenberg Bay to Lambert’s Bay.
THE AMPHIPODA OF SOUTHERN AFRICA 157
Family Paramphithoidae
Epimeria longispinosa K. H. Barnard, 1916
Epimeria longispinosa K. H. Barnard, 1916: 172, pl. 27 (fig. 6).
Records: 34/18/D-VD, a single record.
Diagnosis: Pereon dorsally smooth; coxae 1-3 acutely pointed below, 4 crescent-
shaped, much longer than 3, 5 postero-distally produced into a narrow spiniform
process which extends to end of pleon segment 1; pleon segments 1-4 feebly
‘carinate, carinae on segments 2-4 posteriorly toothed; pleonal epimera quadrate,
without accessory teeth.
Distribution: Endemic, the above record is the only one to date.
Epimeria semiarmata K. H. Barnard, 1916
Epimeria semiarmata K. H. Barnard, 1916: 171, pl. 27 (fig. 3).
Records: 34/18/D, a single record.
Diagnosis: Pereon segments dorsally smooth; coxae 1-3 acutely pointed below,
4 somewhat longer than 3, 5 pentagonal, not postero-distally produced; pleon
segments 1 and 2 dorsally smooth, 3 with a faint carina ending in a minute
tooth, 4 depressed basally, distally humped; pleonal epimera 2 and 3 slightly
produced postero-distally, lacking accessory teeth.
Distribution: Endemic to deep waters off the Cape Peninsula.
Family Pardaliscidae
Halice anacantha K.. H. Barnard, 1925
Halice anacantha K. H. Barnard, 1925: 347, pl. 34 (fig. 12)
Pardisynopia anacantha: J. L. Barnard, 1969: 400. Griffiths, 1974c: 320.
Halice anacantha: Karaman 1974: 13.
Records: 32/17/D, 34/18/D, a few records.
Distribution: Endemic, Plettenberg Bay to Lambert’s Bay.
Remarks: In revising the family Pardaliscidae Karaman (1974) has amalgamated
Pardisynopia with Halice on the basis that component species can no longer be
separated into discreet groups, but provide a full range of variability between
the two previous generic definitions.
Family Phoxocephalidae
Heterophoxus cephalodens sp. nov.
Fig. 17
Description of female (3,5 mm): Head (Fig. 17A) as long as first three pereon
segments, rostrum medio-dorsally keeled, lateral margins with small cornified
ridges, apex acute, reaching beyond apex of article 3 of antenna 1, lower corner
158 ANNALS OF THE SOUTH AFRICAN MUSEUM
of head produced into a strong acute tooth, eyes small, composed of 4-5
individual ocelli; antenna 1 (Fig. 17B) with 7-articulate flagellum, accessory
flagellum 6-articulate; antenna 2 (Fig. 17C) with basal ensiform process,
flagellum 7-articulate; mandibular incisor strongly toothed, spine row of seven
spines, molar represented by a small process bearing two serrate spines, palp
elongate, article 2 with three distal setae, article 3 as long as 2, falciform,
bearing nine apical setae; maxilla 1 with bi-articulate palp exceeding outer plate;
article 4 of maxillipedal palp (Fig. 17D) with two small marginal setae and one
long terminal seta.
Gnathopods 1 and 2 subequal, subchelate, palm oblique, defined by a
distinct step; pereiopods 1 and 2 normal; coxae 1-4 distally setose, coxa 4
excavate posteriorly; article 2 of pereilopod 3 (Fig. 17E) slender, hardly wider
than article 3, articles 4-6 lined on both margins with plumose setae, article 7
spiniform; pereiopod 4 larger than 3 or 5; pereiopod 5 (Fig. 17F) with article 2
postero-distally produced to tip of article 4, posterior margin cut into about
seven teeth, a minute seta in each notch; antero-distal corner of article 2 with
four long plumose setae, articles 3 and 4 each with two plumose setae anteriorly.
First pleonal epimeron (Fig. 17G) minutely produced postero-distally,
second with five distal setae, postero-distal corner subacute; third pleonal
epimeron with an oblique row of setae leading to a small seta-bearing notch
above rounded postero-distal corner; uropods 1-3 extending about equally;
uropod | (Fig. 17H) with outer ramus spiniform, 60% length of inner, inner
ramus with six dorsal and two apical spines; uropod 2 (Fig. 171) with outer
margin of peduncle dorsally keeled, bearing six dorsal spines, rami equal, outer
with eight dorsal and two terminal spines, inner with three dorsal and two
terminal spines; peduncle of uropod 3 (Fig. 17J) ventrally spinose, outer ramus
with second article 25% length of first and terminating in two plumose setae,
inner ramus as long as article | of outer; telson (Fig. 17K) short, extending only
to tip of peduncle of uropod 3, 80% cleft, each lobe bearing two small brush
setae and a minute subapical spine.
Holotype: SAM—A13468, ovigerous female, 3,5 mm.
Type-locality: 34°19'S/18°29’E, 13 July 1967, depth 51 m, substrate khaki sand.
Relationship: The reduced spiniform outer ramus of uropod 2 distinguishes this
species from other members of the genus. The produced post-antennal corner
of the head is unusual, being shared only by H. opus sp. nov. (below).
Material: 34/18/FB/O, 32/18/T, D, VD, a few records.
Heterophoxus opus sp. nov.
Fig. 18
Description of female (4 mm): Head as long as first two pereon segments,
rostrum dorsally smooth, apically rounded, extending to apex of article 3 of
antenna 1, post-antennal corner of head acutely produced (Fig. 18A), eyes
THE AMPHIPODA OF SOUTHERN AFRICA 159
A
D F
E SON
SSS SSS ee
=4@P7)
ID 4 Gy 7; tS Vi
Ug 14 Ny i ff /
U
t
!
|
i
Fig. 17. Heterophoxus cephalodens sp. nov.
Female, 3,5 mm: A—head; B—antenna1; C—antenna 2; D-—articles 3 and 4 of maxillipedal
palp; E—pereiopod 3; F—pereiopod 5; G—pleonal epimera 1-3; H, I, J—uropods 1, 2, 3;
K—telson.
160 ANNALS OF THE SOUTH AFRICAN MUSEUM
composed of about eight individual ocelli; article 1 of antenna | (Fig. 18B) with
four posterior brush setae, articles 2 and 3 much smaller than 1, flagellum
6-articulate, accessory flagellum 4-articulate; antenna 2 (Fig. 18C) with ensiform
process, flagellum 7-articulate; mandibular incisor with distinct teeth, spine row
of eight spines, molar represented by a process bearing two serrate spines, article
2 of palp naked, article 3 with three terminal setae; maxilla | (Fig. 18D) with bi-
articulate palp terminating in seven spines, outer plate with eleven apical spines,
inner plate with three simple setae and one brush seta; article 4 of maxilli-
pedal palp (Fig. 18E) with two small lateral setae and one strong terminal seta.
Gnathopods subchelate, palm oblique, defined by a small lobe, gnathopod 2
slightly the larger; perelopods 1 and 2 normal; coxae 1-4 ventrally setose;
pereiopod 3 (Fig. 18F) with article 2 hardly wider than 3, articles 4-6 setose,
4 and 5 with a long postero-distal plumosa seta; pereiopod 4 longer than 3 or 5,
dactyl elongate, spiniform; pereiopod 5 (Fig. 18G) with article 2 greatly
expanded posteriorly, distally produced to middle of article 4, posterior margin -
with 11 minute serrations each bearing a setule, antero-distal corner of article 2
with a single plumose seta, articles 3 and 4 without plumose setae.
First pleonal epimeron with two distal plumose setae, postero-distally
quadrate, second pleonal epimeron with four distal setae, postero-distally
quadrate; third pleonal epimeron (Fig. 18H) with two distal spines, postero-
distally rounded with two minute setulose notches on posterior margin;
uropod | slightly exceeding 2, peduncle with four dorsal spines, rami subequal
(Fig. 181), outer with three dorsal spines, inner dorsally smooth; uropod 2
(Fig. 18J) with six dorsal and one terminal peduncular spines, rami subequal,
outer with four dorsal spines, inner dorsally smooth; uropod 3 (Fig. 18K) with
peduncle apically spinose, outer ramus with article 2 50°% length of 1, bearing a
long apical spine, inner ramus 80° length of outer, naked; telson (Fig. 18L)
short, 80% cleft, each lobe with a dorsal brush seta, and small apical spine.
Holotype: SAM-—A13469, female, 3,5 mm.
Type-locality: 34°18'S/18°29’E, 13 July 1967, depth 51 m, substrate khaki sand.
Relationships: Because of its eyes this species falls into the genus Heterophoxus,
although the third uropods are more closely allied to those of Harpinia, which
is an eyeless genus as presently defined. However, recent findings indicate that
the presence or absence of eyes is a dubious generic character, since species
possessing both oculate and anoculate forms have recently been described. If
eyes were to be disregarded as generic characters a considerable revision of the
group would be required, a task I am not in a position to carry out. I have thus
allocated this species to Heterophoxus provisionally so as to at least maintain
the status quo.
Heterophoxus opus sp. nov. may be distinguished from H. cephalodens
sp. nov. by the rami of uropod 1, which are subequal, and from other species in
the genus by the produced post-antennal corner of the head.
Material: 34/18/FB/O, two records.
THE AMPHIPODA OF SOUTHERN AFRICA 161
| AE | P
D sor
Fig. 18. Heterophoxus opus sp. nov.
Female, 4mm: A—head; B—antenna 1; C—antenna 2; D—maxilla 1; E—articles 3 and 4
of maxillipedal palp; F—pereiopod 3; G —pereiopod 5; H-—pleonal epimera 1-3;
I, J, K—uropods 1, 2, 3; L—telson.
162 ANNALS OF THE SOUTH AFRICAN MUSEUM
Mandibulophoxus stimpsoni (Stebbing, 1908)
Pontharpinia stimpsoni Stebbing, 19085: 75, pl. 11.
Mandibulophoxus stimpsoni: J. L. Barnard, 1957: 436, figs 3-4.
Records: 32/18/T, 34/18/FB/T, O, D, abundant in False Bay but uncommon
elsewhere.
Distribution: West and South Africa.
Paraphoxus oculatus Sars, 1891
Paraphoxus oculatus: J. L. Barnard, 1960: 240-243, pls 27-28.
Records: 32/17/D, 32/16/D, VD to 34/18/FB/T, O, 34/18/D. numerous
records.
Distribution: Circumboreal.
Platyischnopus herdmani Walker, 1904
Platyischnopus capensis K. H. Barnard, 1925: 338, pl. 34 (figs 13-14).
Platyischnopus herdmani: Rabindranath, 1971la: 521, figs 1-2.
Records: 32/18/T to 34/18/FB/T, O, numerous records.
Distribution: India, South Africa.
Pseudharpinia excavata (Chevreux, 1887)
Harpinia excavata: K. H. Barnard, 1925: 340. J. L. Barnard, 1962c: 47, figs 37-38.
Records; 32/17/D to 34/18/FB/O, 34/18/D, a few records.
Diagnosis: Head with small post-antennal tooth, eyes absent; article 2 of
pereiopod 5 with several antero-distal setae, posterior margin with 10 small
serrations; third pleonal epimeron with small postero-distal tooth.
Distribution: Atlantic, Pacific.
Family Podoceridae
Laetmatophilus purus Stebbing, 1888
Laetmatophilus purus Stebbing, 1888: 1198, pl. 132.
Records: 32/18/O, 32/16/D to 34/18/FB/T, O, 34/18/D, numerous records.
Distribution: Endemic, Mocgambique to South West Africa.
Laetmatophilus tridens K. H. Barnard, 1916
Laetmatophilus tridens K. H. Barnard, 1916: 275, pl. 28 (fig. 22).
Records: 33/18/I, 34/18/FB/T, O, a few records.
Distribution: Endemic, Mocgambique to Saldanha Bay.
THE AMPHIPODA OF SOUTHERN AFRICA 163
Podocerus africanus K. H. Barnard, 1916
Podocerus africanus K. H. Barnard, 1916: 278, pl. 28 (figs 24-25); 1937: 176, fig. 19.
Records: 33/17/T, 34/18/T, 34/18/FB/I, T, O, a few records.
Distribution: Arabia, Natal to South West Africa.
Podocerus inconspicuus (Stebbing, 1888) new synonymy
Podocerus palinuri K. H. Barnard, 1916: 277, pl. 28 (fig. 23).
Podocerus cristatus (non Thompson, 1879): K. H. Barnard, 1916: 276. Griffiths, 1973: 298;
1974a: 202; 1974c: 323.
Records: 29/16/I, 31/16/D to 34/18/FB/T, O, 34/18/O0, D, numerous records.
Distribution: Indian Ocean, west coast of South Africa.
Remarks: K. H. Barnard (1916, 1940) and Griffiths (1973, 1974a, c) have
previously recognized two South African species—Podocerus inconspicuus
(= palinuri) and ‘P. cristatus’—distinguishing them on the basis of degree of
dorsal carination. As more material has been collected, however, it has become
obvious that the two forms represent the extremes of a continuous range of
variation. All previous records of these two forms have thus been combined
under the name P. inconspicuus. This form is probably not synonymous with the
original P. cristatus of Thompson, which lacks a defining tooth on the palm of
gnathopod 2 J, a feature present in all southern African specimens. P. incon-
spicuus has now been recorded in every condition from totally lacking dorsal
carinae (Stebbing 1888), through various intergrades (e.g. Pirlot 1938) to fully
carinate (Barnard 1916—as P. palinuri).
P. inconspicuus should be rediagnosed as follows: Body showing variable
carination, carinae at first developing on pereon segments 6 and 7 and pro-
gressing forwards until in adult specimens the head has a low rounded keel and
all pereon segments show distinct carinae, these being largest on pereon segments
5-7 and on pleon segments | and 2, small subdorsal processes may also develop
on the posterior pereon segments; palm of gnathopod 1 ¢ oblique, dactyl
serrate; palm of gnathopod 2 3 defined by a small lobe and with two strong distal
teeth; palm of gnathopod 1 @ transverse, minutely serrate; palm of gnathopod
2 2 defined by a small lobe and with two small distal teeth; uropods moderately
spinose; telson with 6-8 dorsal spine setae.
P. inconspicuus may be distinguished from P. hystrix Stebbing and P. danae
Stebbing by the presence of a defining lobe on the palm of gnathopod 2 and the
absence of a strong upstanding head process.
Podocerus multispinis K. H. Barnard, 1925
Podocerus multispinis K. H. Barnard, 1925: 367, pl. 34 (fig. 18).
Records: 33/17/O, 34/18/O, 34/18/FB/O, a few records.
Distribution: Endemic, Natal to Saldanha Bay.
164 - ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 19. Podocerus pyurae sp. nov.
Male, 7 mm: A-—lateral aspect; B—articles 5-7 of gnathopod 1; C-—articles 4~7 of
gnathopod 2; D—urosome (dorsal aspect).
Podocerus pyurae sp. nov.
Fig. 19
Description of male (7 mm): Body without dorsal carinae, tubercles or spines;
head slightly longer than first pereon segment, eyes round, red, not reaching
margin of head; antenna | shorter than peduncle of 2, articles 1 and 3 of
peduncle subequal, slightly shorter than 2, flagellum 5-articulate, accessory
flagellum of a single article; antenna 2 considerably stouter than 1, article 5
longer than 4, flagellum of two large articles and one small article.
THE AMPHIPODA OF SOUTHERN AFRICA 165
Coxa | antero-distally produced forwards, following coxae subquadrate;
gnathopod 1 (Fig. 19B) subchelate, articles 5 and 6 subequal, palm oblique,
undefined, dactyl cut into several strong teeth; gnathopod 2 powerfully sub-
chelate (Fig. 19C), article 2 anteriorly keeled, not strongly lobed distally,
article 4 not strongly produced distally, hind margin of article 6 almost as long
as palm, palm oblique, defined by a strong tooth and with a smaller triangular
tooth distally and then a strong rectangular castellate process near finger-hinge,
dactyl strongly curved, subequal to palm; pereiopods 1-2 considerably shorter
than 3-5, article 2 four times as long as wide, 4 somewhat produced antero-
distally, 6 longer than 5, dactyl large; article 2 of pereiopods 3-5 posteriorly
keeled, keel external on pereiopod 3, medial on pereiopods 4 and 5, article 4
somewhat produced postero-distally.
Pleonal epimera 1-3 postero-distally rounded; uropod | (Fig. 19D) much
longer than 2, peduncle with two dorsal rows of slender spines, outer ramus 60%
length of inner and considerably narrow than it, with five dorsal and two
terminal spines, inner ramus with a comb-like row of close-set slender spines
dorsally and four larger spines apically; uropod 2 similar to 1 but considerably
shorter and with fewer spines; uropod 3 consisting of a small lobe with two
minute terminal setae; telson with about six distal setae on each margin.
Female: \ndistinguishable from the male except for the presence of brood plates.
Holotype: SAM-A13480, male, 7 mm.
Type-locality: Branchial cavity of ascidian (Pyura stolonifera) collected inter-
tidally at Kalk Bay, on the shores of False Bay, by Miss R. J. Imrie, 10 December
1973.
Relationships: This species has close affinities with P. hanapepe J. L. Barnard,
P. mangarevae Chevreux and P. zeylanica Walker but differs from all these
species in possessing comb-like rows of spines on the dorsal surfaces of the
inner rami of uropods | and 2.
Material: Seven specimens from the type-locality.
Family Sebidae
Seba saundersi Stebbing, 1875
Fig. 20
Paravalettia chelata K. H. Barnard, 1916: 112, pl. 26 (figs 2-3).
Seba saundersii: K. H. Barnard, 1957: 7, fig. 4.
Records: 33/18/I, 34/18/FB/O, a few records.
Diagnosis: Gnathopod 1 developing from chelate and lacking in palmer teeth,
through subchelate with palm transverse and moderately toothed to subchelate
with palm oblique and strongly toothed; article 5 of gnathopod 2 shorter than 6;
article 2 of pereiopods 4 and 5 subcircular, article 4 weakly expanded in juveniles,
greatly expanded and strongly produced postero-distally in adults; pleonal
epimera postero-distally with a small tooth.
166 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 20. Seba saundersi Stebbing, 1888
Male, 2,5 mm: A-—lateral aspect; B—mandible; C—maxilla 1; D—mazxilliped;
E—uropod 3; F—telson. Female, 2mm: G—gnathopod 1. Male, 1,4mm: H-—gnathopod 1;
I—pereiopod 5. Male, 2,4 mm: J—gnathopod 1; K-—pereiopod 5. Male, 3:0 sammie:
L—gnathopod 1; M—pereiopod 5.
THE AMPHIPODA OF SOUTHERN AFRICA 167
Distribution: Southern Atlantic.
Remarks: The current material includes a sample of 24 individuals from a single
station and this opportunity has been utilized to provide figures of growth
stages (Fig. 20). The variability of the structure of gnathopod 1 and of pereiopod
5 is remarkable and, as these characters are often used for identification, it seems
likely that when developmental stages of other species are known a revision of
the genus will become necessary.
Family Stegocephalidae
Stegocephaloides attingens K. H. Barnard, 1916
Stegocephaloides attingens K. H. Barnard, 1916: 131, pl. 26 (fig. 5). J. L. Barnard, 1961:
60, fig. 29.
Records: 34/18/VD, two records.
Diagnosis: First pereon segment tumid; eyes absent; coxae very large, forming a
continuous shield; coxa 4 with inferior margin very short, evenly curved;
article 2 of pereiopod 5 apically acute, reaching to apex of article 5, hind margin
weakly serrate; third pleonal epimeron postero-distally subquadrate.
Distribution: Angola to Cape Point.
Stegocephaloides australis K. H. Barnard, 1916
Stegocephaloides australis K. H. Barnard, 1916: 129, pl. 28 (fig. 29).
Records: 32/18/O to 34/18/FB/O, 34/18/D, VD, a few records.
Distribution: Endemic, Plettenberg Bay to Lambert’s Bay.
Family Stenothoidae
Proboloides rotunda (Stebbing, 1917)
Metopa rotundus Stebbing, 1917: 39, pl. 7A.
Proboloides rotunda: K. H. Barnard, 1940: 444.
Records: 33/17/O, 34/18/D, 34/18/FB/O, fairly common.
Distribution: Endemic, Natal to Saldanha Bay.
Stenothoe adhaerans Stebbing, 1888
Stenothoe adhaerans Stebbing, 1888: 748, pl. 39.
Records: 32/18/T, 33/17/O, 34/18/?, a few records.
Diagnosis: Antennae subequal, half body length; palm of gnathopod | defined
by five spines, pectinate throughout, dactyl pectinate; gnathopod 2 3 fairly
large, palm occupying 70% length of hand, defined by two spines and irregularly
dentate throughout with one larger tooth near finger-hinge; article 4 of
168 ANNALS OF THE SOUTH AFRICAN MUSEUM
pereiopods 3-5 considerably expanded, postero-distally produced almost to tip
of article 5; uropod 3 uniramous, article 2 of ramus straight.
Distribution: Endemic, Cape Agulhas to Lambert’s Bay.
Remarks: This species was previously known only from the female.
Stenothoe valida Dana, 1853
Stenothoe valida: J. L. Barnard, 1970b: 250, fig. 165.
Records: 32/18/T, 33/17/O, a few records.
Distribution: Cosmopolitan in tropical and temperate seas.
Family Synopiidae
Tiron australis Stebbing, 1908
Tiron australis Stebbing, 19085: 79, pl. 38.
Records: 34/18/FB/T, O, 34/19/O, a few records.
Distribution: Endemic, Natal to False Bay.
Superfamily TALITROIDEA
Family Ceinidae
Afrochiltonia capensis K. H. Barnard, 1916
Chiltonia capensis K. H. Barnard, 1916: 224, pl. 27 (figs 38-40).
Afrochiltonia capensis: K. H. Barnard, 1955: 93.
Records: 34/18/FB/E to 31/18/E, a few records.
Distribution: Endemic, Kosi Bay to Olifants River, a brack-water species.
Family Talitridae
Hyale diastoma K. H. Barnard, 1916
Hyale diastoma K. H. Barnard, 1916: 232, pl. 28 (fig. 8).
Records; 33/18/I, 34/18/FB/I, a few records.
Distribution: False Bay to South West Africa.
Hyale grandicornis (Kro6yer, 1845) new synonymy
Allorchestes inquirendus K. H. Barnard, 1940: 477, fig. 34b—c. Griffiths, 1974a: 202; 1974c: 328.
Hyale grandicornis: Hurley, 1957: 904, figs 1-29.
Records: 26/16/I to 34/19/I, numerous records.
Distribution: Cosmopolitan in tropical and temperate seas.
Remarks: Allorchestes inquirendus was erected by K. H. Barnard (1940) solely
on the basis of the lobe on article 5 of gnathopod 2 3, a feature supposedly
absent in the otherwise identical Hyale grandicornis. The transitory or variable
THE AMPHIPODA OF SOUTHERN AFRICA 169
nature of such processes has, however, been known for some time, thus Hurley
(1957) writes: ‘Although this diagnostic character is unmistakable in the adult ¢
of Allorchestes the generic distinction is slurred over by the development of a
similar process in the juveniles of Hyale; a process which does not always
disappear in the adult males . . . being well exemplified by Hyale grandicornis.’
Reid (1951) has noted similar changes in H. perieri and a continuous range of
variation is found amongst H. grandicornis from South Africa. Allorchestes
inquirendus is thus an invalid species and falls to Hyale grandicornis.
Hyale hirtipalma (Dana, 1852)
Hyale hirtipalma: K. H. Barnard, 1916: 234. Hurley, 1957: 922, figs 118-146.
Hyale macrodactyla (non Stebbing, 1899): K. H. Barnard, 1916: 235.
Records: 29/16/I to 34/18/FB/I, a few records.
Distribution: Pacific, South Atlantic.
Hyale maroubrae Stebbing, 1899
Hyale maroubrae: Hurley, 1957: 913, figs 51-71.
Records: 33/18/I, 34/18/FB/I, a few records.
Distribution: Widespread in Southern hemisphere.
Hyale plumulosa (Stimpson, 1853)
Hyale plumulosa: Bousfield, 1973: 155, pl. 44 (fig. 2).
Records: 34/19/I, a single record.
Diagnosis: Article 1 of antenna | not postero-distally lobed; article 5 of antenna 2
and flagellum covered with a dense growth of fine plumose setae; coxae 1-4 with
posterior triangular processes; palm of gnathopod 1 transverse; palm of
gnathopod 2 ¢ oblique, evenly convex, subequal to hind margin; article 2 of
pereiopods 1-5 weakly crenulate posteriorly; uropod | with long spine on inner
distal margin of peduncle.
Distribution: Atlantic and Pacific coasts of North America, South Africa.
Remarks: The above specimens, the first of this species recorded from Africa,
agree perfectly with the description given by Bousfield (1973).
| Hyale saldanha Chilton, 1912
Hyale saldanha Chilton, 1912: 509, pl. 2 (figs 24-29).
Records; 29/16/I to 34/19/I, numerous records.
Distribution: Endemic, East London to South West Africa.
170 ANNALS OF THE SOUTH AFRICAN MUSEUM
Orchestia ancheidos (K. H. Barnard, 1916)
Talorchestia ancheidos K. H. Barnard, 1916: 221, pl. 27 (figs 35-36); 1940: 470, fig. 31. Ruffo,
1947: 121, figs 3-5.
Orchestia ancheidos: Ruffo, 1958: 43, figs 3-4.
Records: 32/18/E to 34/19/E, a few records.
Distribution: Madagascar, southern and West Africa, usually in brack waters.
Orchestia dassenensis (K. H. Barnard, 1916)
Parorchestia dassenensis K. H. Barnard, 1916: 227, pl. 28 (figs 1-2).
Records: 33/18/I, 34/19/I, a few records.
Diagnosis: Antenna | almost as long as 2, antenna 2 slender; articles 4-6 of
gnathopod 1| ¢ each with a posterior pellucid lobe; articles 2 and 3 of gnathopod
2 ¢ anteriorly lobed, article 6 oval, palm oblique, moderately spinose, a small
notch near finger-hinge and another near defining angle, dactyl slightly exceeding
palm, inner margin sinuous, tip averted; articles 4 and 5 of pereiopod 5 linear;
third pleonal epimeron quadrate with a minute postero-distal tooth; outer ramus
of uropod | smooth.
Distribution: Endemic, Cape Agulhas to Saldanha Bay.
Orchestia gammarella (Pallas, 1766) new synonymy
Orchestia gammarella: Chevreux & Fage, 1925: 274, fig. 284. Bousfield, 1973: 159, pl. 45 (fig. 1)
Talorchestia inaequalipes K. H. Barnard, 1951: 705, fig. 5a—b. Griffiths, 1974c: 330.
Records: 33/18/I, a few records.
Diagnosis: Articles 4 and 5 of antenna 2 3 not expanded; palm of gnathopod 2 ¢
evenly convex, smooth, subequal to hind margin, dactyl evenly convex, tip not
averted, slightly exceeding palm; articles 4 and 5 of pereiopod 5 of adult 3
strongly expanded, giving limb an oar-like appearance; outer ramus of uropod |
dorsally spinose.
Distribution: North Atlantic, South Africa.
Remarks: A re-examination of Barnard’s types of Talorchestia inaequalipes has
shown gnathopod 1 @ to be distinctly subchelate. This species thus should be
transferred to Orchestia where it appears to be synonymous with Orchestia
gammarella.
Orchestia platensis Kroyer, 1845
Orchestia platensis: Bousfield, 1973: 160, pl. 46 (fig. 2).
Records: 34/19/I, two records.
Diagnosis: Articles 4 and 5 of antenna 2 ¢ greatly inflated; palm of gnathopod
2 § convex, oblique, subequal to hind margin, a sharp notch near defining angle
followed by a small hump which is prolonged as a ridge running along medial
margin of hand, dactyl slightly exceeding palm, tip not averted; articles 4 and 5
THE AMPHIPODA OF SOUTHERN AFRICA tt
of pereiopod 5 ¢ strongly inflated but cylindrical (not oar-like as in O. gamma-
rella); outer ramus of uropod 3 smooth.
Distribution: Cosmopolitan in tropical and temperate seas.
Remarks: This species has not previously been recorded from South Africa.
Orchestia rectipalma K. H. Barnard, 1940
Orchestia rectipalma K. H. Barnard, 1940: 473, fig. 32.
Records: 31/18/E to 34/19/E, a few records.
Distribution: Endemic, Natal to South West Africa, a brack-water species.
Talorchestia australis K. H. Barnard, 1916
Talorchestia australis K. H. Barnard, 1916: 220, pl. 27 (figs 33-34); 1940: 470, fig. 30.
Records: 34/19/I, E to 33/18/I, a few records.
Distribution: Endemic, Mogambique to South West Africa.
Talorchestia capensis (Dana, 1853)
Talorchestia capensis: K. H. Barnard, 1916: 216; 1940: 470, fig. 28.
Records: 29/16/I to 34/19/I, a few records.
Distribution: Mediterranean, Atlantic, South Africa.
Talorchestia quadrispinosa K. H. Barnard, 1916
Orchestoidea fischerii (non Milne-Edwards, 1826): Stebbing, 1910a: 459.
Talorchestia quadrispinosa K. H. Barnard, 1916: 217, pl. 27 (figs 29-32); 1940: 470, fig. 29.
Records: 29/16/I to 34/18/FB/I, the most abundant sandy beach species in
this area.
Distribution: Endemic, False Bay to South West Africa.
Family Temnophliidae fam. nov.
Diagnosis: Talitroidea with body dorsally depressed, pereon segments produced
laterally as pleurae; coxae reduced; mandibular molar nontriturative; maxilla 1
without palp; palp of maxilliped bi-articulate; uropods | and 2 uniramous,
uropod 3 without rami; telson entire.
Type-genus: Temnophlias K. H. Barnard, 1916.
Remarks: It has long been recognized that the genus Temnophlias should be
removed from the Phliantidae, into which it was originally placed, since its body
Shape and reduced mouthparts and uropods are inconsistent with the norm for
that family (J. L. Barnard 1969, 19725). J. L. Barnard (19726) has suggested the
possibility of placing Temnophlias in the Eophliantidae; however my feeling is
that the depressed body form and entire telson of Temnophlias preclude this
172 ANNALS OF THE SOUTH AFRICAN MUSEUM
possibility and that the genus warrants its own family—the Temnophliidae. The
removal of Temnophiias enables the diagnosis of Phliantidae to be tightened to
read as follows:
Family Phliantidae: Talitroidea with body greatly depressed; pereon segments
not laterally extended as pleurae; coxae not reduced; mandibular molar
nontriturative; maxilla 1 with or without palp; palp of maxilliped 3 or
4-articulate; uropod | birarious, uropod 2 biramous (except Pereionotus),
uropod 3 uniramous or rami absent; telson entire. '
Temnophliidae would thus be distinguished from Phliantidae by the
presence of pleurae on the pereon, the reduced maxillipedal palp and uniramous
first uropod.
Temnophlias capensis K. H. Barnard, 1916
Fig. 21
Temnophlias capensis K. H. Barnard, 1916: 158, pl. 26 (figs 25-35).
Records: 29/16/I to 34/19/I, fairly common intertidally and in shallow waters.
Distribution: Endemic, Still Bay to South West Africa.
Remarks: This species has been refigured here (Fig. 21) in order to provide a
suitable reference for the new family Temnophliidae. Although the lateral
margins of the pereon segments are distinctly discontinuous it should be noted
that this is not as marked as would appear from K. H. Barnard’s (1916) figures.
The distinction in body shape between 7. capensis and T. hystrix is not in fact
very great—the latter merely having developed strong dorsal carinae and lateral
processes on the pereonites and coxae.
Temnophlias hystrix K. H. Barnard, 1954
Temnophlias hystrix K. H. Barnard, 1954: 130, fig. 8.
Records: 30/17/I to 34/18/I, a few records.
Diagnosis: Head with tridentate dorsal process; pereon segments each with a
pair of lateral processes, | with two medio-dorsal processes, 2—7 with a single
medio-dorsal process; coxae 1-4 bifid, 5—7 trifid; pereiopods all chelate in both
Sexes.
Distribution: Endemic, False Bay to Cape Hangklip, usually intertidal.
Suborder CAPRELLIDEA
Family Aeginellidae
| Eupariambus fallax K. H. Barnard, 1957
Eupariambus fallax K. H. Barnard, 1957: 9, fig. 6.
Records: 32/17/O, D to 34/18/FB/O, 34/18/D, numerous records.
Distribution: Endemic, Still Bay to Lambert’s Bay.
THE AMPHIPODA OF SOUTHERN AFRICA
ee or
so Ne
ZB
ae
8
y ©
6
G
Fig. 21. Temnophlias capensis K. H. Barnard, 1916
Female, 7 mm:
A—dorsal aspect; B—mandible;
C—maxilla 1; D-—maxilla 2;
E—maxilliped; F-—setae of brood lamellae; G—pleopod 3; H, I, J—uropods 1, 2, 3.
L735
174 ANNALS OF THE SOUTH AFRICAN MUSEUM
Orthoprotella mayeri K. H. Barnard, 1916
Orthoprotella mayeri: K. H. Barnard, 1916: 284; 1925: 372.
Records; 34/18/D, a single record.
Distribution: Indo-Pacific.
Paradeutella serrata Mayer, 1903
Paradeutella serrata Mayer, 1903: 47, pl. 2 (fig. 6), pl. 6 (figs 68-69).
Records: 34/18/FB/I, a single record.
Diagnosis: Head with dorsal spiniform process; pereon segments | and 2 also
with large medio-dorsal processes, remaining segments with small medio-dorsal
humps; antenna 2 about as long as peduncle of 1; article 2 of gnathopod 2
longer than pereon segment 2, bearing an acute distal process, palm poorly
defined with three small teeth near finger-hinge, dactyl simple; pereiopods | and 2
minute, uniarticulate; pereiopods 3—5 fully developed; abdomen of 3 with two
pairs of small lobes.
Distribution: Endemic, known only from the above record.
Pseudaeginella tristanensis (Stebbing, 1888)
Pseudaeginella tristanensis: Stephensen, 1949: 52, fig. 23.
Records: 33/18/T, a single record.
Distribution: Tristan da Cunha, South Africa.
Pseudoprotella phasma (Montagu, 1804)
Pseudoprotella phasma: Chevreux & Fage, 1925: 437, fig. 423.
Records: 29/14/D, a single record.
Diagnosis: Head with strong rostral projection; pereon segment | with a medio-
dorsal process posteriorly, 2 with a pair of processes medially and another
posteriorly (large specimens with further spinose processes on pereon segments 3
and 4); lateral spinose processes above insertions of gnathopod 2 and gills;
pereiopods | and 2 of a single minute segment; pereiopods 3-5 fully developed;
abdomen of J with two pairs of uniarticulate appendages.
Distribution: Mediterranean, eastern Atlantic.
Family Caprellidae
Caprella cicur Mayer, 1903
Caprella cicur Mayer, 1903: 75, 97, pl. 4 (figs 5—7), pl. 8 (figs 3-5).
Records; 32/18/T to 34/18/FB/I, T, O, numerous records.
Distribution: Endemic, Natal to Lambert’s Bay.
THE AMPHIPODA OF SOUTHERN AFRICA 17S
Caprella danilevski Czerniavski, 1868
Caprella danilevskii: McCain, 1969: 22-25, figs 10-11.
Records: 32/18/T, 33/18/I, 34/18/FB/I, a few records.
Distribution: Widespread, pantropical.
Caprella equilibra Say, 1818
Caprella equilibra: McCain, 1968: 25-30, figs 12-13.
Records: 32/18/T, O to 34/18/FB/T, O, numerous records.
Distribution: Cosmopolitan.
Caprella natalensis Mayer, 1903
Caprella acutifrons var. natalensis Mayer, 1903: 81, pl. 3 (figs 22-23).
Caprella natalensis: Laubitz, 1972: 47, pl. 9 (figs F—G), pl. 10 (figs F—K).
Records: 33/18/I, 34/18/T, two records.
Distribution: Pacific North America, Tristan da Cunha, South Africa.
Caprella laevipes Mayer, 1903
Caprella laevipes Mayer, 1903: 108, pl. 5 (fig. 2), pl. 8 (figs 14-16).
Records: 32/18/T to 34/18/FB/T, O, a few records.
Distribution: Endemic, Natal to Lambert’s Bay.
Caprella penantis Leach, 1814 new synonymy
Caprella falsa Mayer, 1903: 101, pl. 4 (fig. 15).
Caprella penantis: McCain, 1968: 33-40, figs 15-16. Laubitz, 1972: 41, pl. 9 (figs A-—E),
pl. 10 (figs A—E).
Records: 32/18/T to 34/18/FB/I, T, O, numerous records.
Distribution: Cosmopolitan in tropical and temperate seas.
Remarks: The maintenance of C. falsa, created by Mayer (1903) solely on the
basis of the inflated article 2 of antenna 1, appears unjustified, since similar
variations in the stoutness of antenna | are found within other species (e.g.
C. cicur of Mayer, 1903: pl. 4, figs 6-7).
Caprella scaura Templeton, 1836
Caprella scaura: McCain, 1968: 40-44, figs 17-18.
Records: 29/16/I, 32/18/T to 34/18/T, quite common.
Distribution: Cosmopolitan.
Hemiaegina minuta Mayer, 1890
Hemiaegina minuta: McCain, 1968: 61-64, figs 29-30.
Records: 34/18/FB/T, a single record.
Distribution: Cosmopolitan in tropical and temperate seas.
176 ANNALS OF THE SOUTH AFRICAN MUSEUM
Family Cyamidae
Cyamus balaenopterae K. H. Barnard, 1931
Cyamus balaenopterae: K. H. Barnard, 1932: 309, fig. 171.
Records; Ectoparasitic on Blue and Fin Whales, Saldanha Bay whaling station.
Distribution: Widespread on Fin and Blue Whales.
Cyamus boopis Lutken, 1873
Paracyamus boépis: K. H. Barnard, 1932: 312.
Cyamus boopis: Margolis, 1955: 124, figs 7-12.
Records: Ectoparasitic on Humpback Whales, Saldanha Bay whaling station.
Distribution: Widespread on Humpback Whales.
Cyamus erraticus Roussel de Vauzéme, 1834
Paracyamus erraticus: K. H. Barnard, 1932: 310, fig. 172.
Cyamus erraticus: Margolis, 1955: 132, figs 1-6.
Records: Ectoparasitic on Right Whales, Cape Town and Saldanha Bay whaling
stations. .
Distribution: Widespread on Right Whales.
Cyamus gracilis Roussel de Vauzéme, 1834
Paracyamus gracilis: K. H. Barnard, 1932: 312, fig. 173.
Records: False Bay, ectoparasitic on Right Whale.
Diagnosis: Palp of maxilliped present in juveniles, lost in adult; body parallel
sided; head fused to pereon segment 1; pereon segment 2 laterally rounded,
branchiae on segments 3 and 4 single, as long as segments 3-5 together, accessory
lobes in § double on both segments (in 2 absent); pereon segments of 3 without
ventral processes, 2 with a single pair of processes on pereon segment 5.
Distribution: Southern oceans, ectoparasitic on Right Whales.
Cyamus ovalis Roussel de Vauzéme, 1834
Cyamus ovalis: K. H. Barnard, 1932: 307, fig. 170.
Records: Cape Town, False Bay, Saldanha Bay, ectoparasitic on Right Whales.
Diagnosis: Maxillipedal palp present in juvenile and adult; body broadly oval;
pereon segment | distinguished from head by an oblique groove; pereon
segment 2 produced postero-distally into a hooked process which engages an
anterior process of pereon segment 3; branchiae on segments 3 and 4 each
consisting of two equal lobes as long as segments 2-6 together, accessory lobes
in 3 single on segment 3, double on segment 4; 3 pereon with one pair of ventral
processes on each of segments 6 and 7, 2 with a pair of blunt processes on
segment 5 and a pair of tubercles on each of segments 6 and 7.
Distribution: Widespread on Right Whale and North Pacific Whale.
THE AMPHIPODA OF SOUTHERN AFRICA iMeAgs
Isocyamus delphini (Guérin-Meéneville, 1836)
Isocyamus delphini: K. H. Barnard, 1932: 313-314. Stephensen, 1942: 454-455.
Records: ‘From dolphin’—exact location unknown.
Diagnosis: Body ovate, head completely fused to pereon segment 1; gills on
pereon segments 3 and 4 short and stout, accessory gills in 3 single, almost as
long as gill, an outward-directed process projecting from base of each gill;
dS pereon with a pair of ventral tubercles on each of segments 5-7, 2 with a pair
of inward-directed processes on segment 5 and a pair of tubercles on each of
segments 6 and 7.
Distribution: Widespread on dolphins, Pilot Dolphins and False Killer Whales.
Remarks: This species has not previously been recorded from southern Africa.
Neocyamus physeteris (Pouchet, 1888)
Paracyamus physeteris: Stephensen, 1942: 453.
Neocyamus physeteris: Margolis, 1955: 131, figs 21-23.
Records: SAM-—A12307—station data unknown.
Diagnosis: Body slender with pereon segments 3 and 4 half as wide as 5 and 6;
gills short, each divided into about 12 filiform appendages, accessory gills
absent.
Distribution: Widespread on Sperm Whales and rarely Ocean Dolphin.
Remarks: This is the first record of this species from southern Africa.
Family Phtisicidae
Caprellina longicollis (Nicolet, 1849)
Caprellina longicollis: McCain, 1969: 289, fig. 2.
Records: 29/16/I to 34/18/I, T, 34/18/FB/I, T, fairly common.
Distribution: Mediterranean, southern oceans.
Caprellina spiniger K. H. Barnard, 1916
Caprellina spiniger K. H. Barnard, 1916: 282, pl. 28 (fig. 35).
Records; 33/18/1, 34/18/FB/I, a few records.
Distribution: Endemic, Mossel Bay to South West Africa.
| Phtisica marina Slabber, 1769
Phtisica marina: Chevreux & Fage, 1925: 434, fig. 422. McCain, 1968: 91-97, fig. 46.
Records: 32/18/T, 32/17/D to 34/17/D, 34/18/FB/O, numerous records.
Distribution: Mediterranean, Atlantic, southern Africa.
178 ANNALS OF THE SOUTH AFRICAN MUSEUM
ACKNOWLEDGEMENTS
I should like to express my gratitude to Professor J. H. Day for his construc-
tive guidance throughout this project. This manuscript and those of previous
papers in this series have been kindly read by Dr J. Laurens Barnard. My thanks
also to Dr B. F. Kensley for the loan of material in the collections of the South
African Museum. I am indebted to the South African Council for Scientific and
Industrial Research for their financial support of this project.
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THE AMPHIPODA OF SOUTHERN AFRICA 179
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Day, J. H. 1970. The biology of False Bay, South Africa. — Trans. R. Soc. S. Afr. 39: 211-221.
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19: 67-83.
GRIFFITHS, C. L. 1973. The Amphipoda of southern Africa, Part 1. The Gammaridea and
Caprellidea of southern Mocambique.— Ann. S. Afr. Mus. 60: 265-306.
GRIFFITHS, C. L. 1974a. The Amphipoda of southern Africa, Part 2. The Gammaridea and
Caprellidea of South West Africa south of 20°S.— Ann. S. Afr. Mus. 62: 169-208.
GRIFFITHS, C. L. 19746. The Amphipoda of southern Africa, Part 3. The Gammaridea and
Caprellidea of Natal.— Ann. S. Afr. Mus. 62: 209-264.
GriFFITHS, C. L. 1974c. The Amphipoda of southern Africa, Part 4. The Gammaridea and
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—Proc. Linn. Soc. N.S.W. 4: 319-350.
Hotsincer, J. R. 1974. Comments on the newly proposed gammaridean amphipod families
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Hurtey, D. E. 1957. Studies on the New Zealand amphipodan fauna. No. 14. The genera
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Kamm Maka, R. A. 1969. Contribution a l’étude de quelque espéces, du genre Ampelisca
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KARAMAN, G. S. 1972. On some new or very interesting Amphipoda of the Adriatic Sea. —
Memorie Mus. civ. Stor. nat. Verona 20: 97-147.
KARAMAN, G. S. 1974. Revision of the family Pardaliscidae with diagnosis of genera, distribu-
tion of species and bibliography.— Acta Adriatica 15(7): 1-46.
KARAMAN, G. S. & RuFro, S. 1971. Contributo alla conoscenza della specie Mediterranee del
genere Maera (Crustacea Amphipoda).— Memorie Mus. civ. Stor. nat. Verona 19: 113-176.
Lausitz, D. R. 1972. The Caprellidae (Crustacea, Amphipoda) of Atlantic and Arctic Canada.
—Publs biol. Oceanogr. natn. Mus. nat. Sci. (Can.) 4: 1-82.
180 ANNALS OF THE SOUTH AFRICAN MUSEUM
LepoyeR, M. 1967. Amphipodes gammariens des herbiers de phanérogames marines de la
région de Tuléar (République Malgache). Etude systématique et écologique. — Annls Univ.
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MARGOLIS, L. 1955. Notes on morphology, taxonomy and synonomy of several species of
whale-lice (Cyamidae, Amphipoda).—J. Fish. Res. Bd Can. 12: 121-133.
Mayer, P. 1903. Die Caprellidae der Siboga Expedition. — Siboga Exped. 34: 1-158.
McCain, J. C. 1968. The Caprellidae (Crustacea: Amphipoda) of the western Northern
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McCain, J. C. 1969. New Zealand Caprellidae (Crustacea: Amphipoda).—N.Z. J] mar.
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McCain, J. C. 1970. Familial taxa within the Caprellidea (Crustacea: Amphipoda).—Proc.
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MILLARD, N. A. H. & Scott, K. M. F. 1954. The ecology of South African estuaries. Part 6:
Milnerton Estuary and the Diep River, Cape.— Trans. R. Soc. S. Afr. 34: 279-324.
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Ampeliscidae.—Limnol. Oceanogr. 16: 357-387.
Monop, TH. 1937. Missions A. Gruvel dans le canal de Suez. 1. Crustacés.—Mém. Inst.
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Myers, A. A. 1974. A first record of the genus Pseudomegamphopus Myers (Crustacea,
Amphipoda) from the Indo-West Pacific with a redescription of P. jassopsis (K. H. Barnard)
comb. nov.—Trans. roy. Soc. S. Afr. 41: 195-202.
NAGATA, K. 1965. Studies on marine gammaridean Amphipoda of the Seto Inland Sea. I. —
Publs Seto mar. biol. Lab. 13: 131-186.
OLDEVIG, H. 1933. Sveriges Amphipoder. — Meddn. Géteborgs Mus. zool. Avd. 62: 3-282.
PENRITH, M-L & KeENsLeEy, B. F. 1970. The constitution of the intertidal fauna of rocky
shores of South West Africa. Part 1: Liideritzbucht.— Cimbebasia (A) 1: 191-239.
PiLxal, N. K. 1957. Pelagic Crustacea of Travancore, III: Amphipoda.— Bull. cent. Res. Inst.
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PirLoT, J. M. 1938. Les amphipodes de l’expédition du Siboga. Deuxiéme partie: Les amphi-
podes gammarides. III: Les amphipodes littoraux. 2: Familles des Dexaminidae, Talitridae,
Aoridae, Photidae, Ampithoidae, Corophiidae, Jassidae, Cheluridae et Podoceridae. —
Siboga Exped. monogr. 33f: 329-359.
RABINDRANATH, P. 1971a. Haustoriid amphipods (Crustacea) from India. — Hydrobiologia 38:
521-539.
RABINDRANATH, P. 19716. On a collection of Isaeidae (Crustacea, Amphipoda) from the
southern Indian region. —Bijdr. Dierk 41: 67-93.
Reip, D. M. 1951. Report on the Amphipoda (Gammaridea and Caprellidea) of the coast of
tropical West Africa.—Aflantide Rep. 2: 189-291.
RurFro, S. 1969. Studi sui crostacei amfipodi LXVII. Terzo contributo alla conscenza degli
amfipodi del mar Rosso.— Memorie Mus. civ. Stor. nat. Verona 17: 1-77.
Sars, G. O. 1895. An account of the Crustacea of Norway, with short descriptions and figures of
all the species. 1. Amphipoda. Cristiania, Copenhagen: Cammermeyers.
SCHELLENBERG, A. 1925. Crustacea VIII: Amphipoda.—Beitr. Kennt. Meeresfauna Westafr.
3: 113-204.
SCHELLENBERG, A. 1926a. Die Gammariden der deutschen Siidpolar-Expedition 1901-1903. —
Dt. Siidpol.-Exped. 18: 235-414.
SCHELLENBERG, A. 1926). Amphipoda 3: Die Gammariden der deutschen Tiefsee-Expedition.
Dt. Tiefsee-Exped. 23: 195-243.
THE AMPHIPODA OF SOUTHERN AFRICA 181
SCHELLENBERG, A. 1953. Erganzungen zur Amphipodon-fauna Siidwest-Africas nebst
Bemerkungen tiber Brutraumbildund.— Mitt. zool. Mus. Berl. 29: 107-126.
Scott, K. M. F., Harrison, A. D. & MAcNAE, W. 1952. The ecology of South African
estuaries. Part 2. The Klein River Estuary, Hermanus, Cape.—Trans. R. Soc. S. Afr. 33:
283-331.
SEXTON, E. W. & REID, D. M. 1951. The life-history of the multiform species Jassa falcata
(Montagu) (Crustacea, Amphipoda) with a review of the bibliography of the species. —
J. Linn. Soc. Lond. (Zool) 42: 29-91.
SHOEMAKER, C. R. 1933. Amphipoda from Florida and the West Indies.—Am. Mus. Novit.
598: 1-24.
SIVAPRAKASAM, T. E. 1966. Amphipoda from the east coast of India. Part 1. Gammaridea. —
J. mar. biol. Ass. India 8: 82-122.
SIVAPRAKASAM, T. E. 1967. Leucothoid Amphipoda from the Madras coast.—J. mar. biol.
Ass. India 9: 384-391.
SIVAPRAKASAM, T. E. 1968. A new species and a new record of Amphipoda from the Madras
coast.—J. mar. biol. Ass. India 10: 274-282.
STEBBING, T. R. R. 1888. Report on the Amphipoda collected by H.M.S. Challenger during the
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STEBBING, T. R. R. 1904. Gregarious Crustacea from Ceylon.—Spolia zeylan. 2: 1-29.
STEBBING, T. R. R. 1908a. On two new species of northern Amphipoda.—J. Linn. Soc. Lond.
(Zool.) 30: 191-197.
STEBBING, T. R. R. 1908. South African Crustacea (Part IV).— Ann. S. Afr. Mus. 6: 1-96.
STEBBING, T. R. R. 1910a. General Catalogue of South African Crustacea (Part V of S.A.
Crustacea).— Ann. S. Afr. Mus. 6: 447-473.
STEBBING, T. R. R. 19100. Scientific results of the trawling expedition of H.M.C.S. ‘Thetis’ off
the coast of New South Wales in February and March, 1898. Crustacea, Part 5.
Amphipoda.— Mem. Aust. Mus. 4: 565-658.
STEBBING, T. R. R. 1917. South African Crustacea (Part [X).—Ann. S. Afr. Mus. 17: 23-46.
STEBBING, T. R. R. 1918. Some Crustacea of Natal.— Ann. Durban Mus. 2: 47-75.
STEPHENSEN, K. 1942. The Amphipoda of N. Norway and Spitzbergen with adjacent waters. —
Troms? Mus. Skr. 3: 363-526.
STEPHENSEN, K. 1949. The Amphipoda of Tristan da Cunha.— Results Norw. scient. Exped.
Tristan da Cunha 3: 1-61.
VADER, W. 1970. The status of Bathyporeia gracilis Sars (Amphipoda, Haustoriidae).—
Sarsia 43: 155-162.
WALKER, A. O. 1904. Report on the Amphipoda collected by Professor Herdman, at Ceylon
in 1902. In: Royat Society. Report to the government of Ceylon on the pearl oyste
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London: the Society.
WALKER, A. O. 1909. Amphipoda Gammaridea from the Indian Ocean, British East Africa,
and the Red Sea.—Trans. Linn. Soc. (2) (Zool) 12: 323-344.
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
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.
C. L. GRIFFITHS
THE AMPHIPODA OF SOUTHERN AFRICA
PART 5
THE GAMMARIDEA AND CAPRELLIDEA OF
THE CAPE PROVINCE WEST OF CAPE AGULHAS
VOLUME 67 PART 6 OCTOBER 1975 | ISSN 0303-2515
CAPE TOWN. |
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PLEISTOCENE MOLLUSCS FROM THE WEST AND SOUTH COASTS
OF THE CAPE PROVINCE, SOUTH AFRICA
By
R. N. KILBURN
Natal Museum, Pietermaritzburg
&
A. J. TANKARD
South African Museum, Cape Town
(With 22 figures)
[MS accepted 6 January 1975]
ABSTRACT
Nineteen species of Pleistocene to Recent littoral molluscs are discussed, belonging to
the families Trochidae, Stomatiidae, Potamididae, Cerithiidae, Calyptraeidae, Buccinidae,
Terebridae, Acteonidae, Nuculanidae, Ungulinidae, Lucinidae, Veneridae, Petricolidae,
Donacidae and Tellinidae. New species and subspecies are described for the genera Cerithidea
(Cerithidea), Crepidula capensis (subsp. nov.), Triumphis, Duplicaria, Pupa (Strigopupa),
Petricola (Claudiconcha), Donax, Gastrana. New synonyms, new combinations, new records,
and cases of revised status are discussed. These include Cantharidus suarezensis suarezensis
(Fischer, 1878), C. s. fultoni (Sowerby, 1889), Pseudostomatella orbiculata (A. Adams, 1850),
Cerithium scabridum rufonodulosum E. A. Smith, 1901, Nuculana (Lembulus) bicuspidata
(Gould, 1845), Felania diaphana (Gmelin, 1791), Loripes (Microloripes) liratula (Sowerby,
1889), Venerupis dura (Gmelin, 1791), Macoma (Heteromacoma) tricostata (ROmer, 1872),
Leporimetis (Leporimetis) hanleyi (Dunker, 1853).
CONTENTS
PAGE
FPROCUCHONUMy Ee ee So awe. ce ete Ge ee oe 69 SS
Systematic descriptions
GlasstGastropodas Mey 8k gM eg OS) UV ES6
GlassuBivadl vias Meee Pete On co ps, Ok ee BA.) 206
FAGKMOWIEMSCIMCMESE NS) 5. ay csc oes Oe er gate 223
INclerenceSmeerme rt Re hoe Se Le re ie ag, 22
INTRODUCTION
Numerous exposures of Pleistocene marine sediments are preserved on the
wave-cut platforms adjacent to the present South African coast, and in estuaries
and lagoons. Radio-carbon measurements show that these deposits are beyond
the range of the “C dating technique (Tankard, in press). Generally the
Pleistocene faunas of the South African coastlands are poorly known. More
than 160 species of invertebrate fossils have so far been identified in this project.
The west and south coasts of the Cape Province have been tectonically
1am
Ann. S. Afr. Mus. 67 (6), 1975: 183-226, 22 figs.
184 ANNALS OF THE SOUTH AFRICAN MUSEUM
unstable in the Tertiary and possibly through the Early and Middle Pleistocene.
Between Cape Town and St Helena Bay (Fig. 1) Neogene marine sediments are
found at lower elevations than their chronostratigraphic equivalents on the Cape
south coast, or the South West African coast. Furthermore, between Cape Town
and St Helena Bay the highest observed Pleistocene beach deposits are at
10 m a.s.l. (above mean sea level) (Tankard, in press). Carrington & Kensley
(1969) record a series of transgressive complexes up to 90 m a.s.l. on the
Namaqualand coast, while Davies (1971) has identified Pleistocene raised
beaches at 60 m a.s.l. on the south coast. We envisage that intermittent sagging
took place in the area between the Olifants River and the Cape Peninsula
throughout most of the Cenozoic.
In the Saldanha—Langebaan area there are several exposures of Early
Pleistocene shelly sands. Just north of Langebaan (1,5 km) in a shallow quarry
at 9,5 ma.s.l. there is a shelly deposit overlying Tertiary limestone. The fauna is
characterized by Fissurella robusta Sowerby, Cerithidea sp. nov., Triumphis
sp. nov., Purpura praecingulata (Haughton), Petricola sp. nov., and large
Perna perna (Linnaeus) (= Mytilus tomlini Haughton). Similar deposits are
exposed on either side of the Hoedjiespunt peninsula where they lie either
directly on a quartz porphyry platform or on Miocene phosphorite. Behind the
Sea Harvest factory on the Hoedjiespunt peninsula the littoral deposit includes
wave-generated beach boulders. Here the horizon is exposed up to 8 ma.s.l. and
is composed largely of Patella spp., although Fissurella robusta, Petricola sp. nov.,
and large Perna perna (Linnaeus) are common. These deposits are assigned to
the Early Pleistocene, rather than the Late Pleistocene, because the mollusc
fauna is more primitive than any of the Late Pleistocene sites so far examined,
and does not contain any of the warm-water fauna that characterizes deposits
from other Late Pleistocene embayments and estuaries and is poorly preserved
when compared with these younger deposits.
Generally the mollusc fauna from the Early Pleistocene sites at Saldanha
and Langebaan is near-shore in character. At the Langebaan site the barnacle
Balanus amphitrite Darwin encrusts the Tertiary limestone. B. amphitrite is today
an inhabitant of the lower intertidal to infratidal zones. This would imply that
the strand line should really be recorded at about 10 m a.s.l.
The mollusc fauna, particularly Fissurella robusta, Triumphis sp. nov.
Purpura praecingulata, large Perna perna, and Petricola sp. nov. suggests
correlation with Haughton’s (1932) Zone-D of the Namaqualand coast, and
the 45-50 m transgression complex of Carrington & Kensley (1969). If this
correlation is correct the 10 m shoreline in the Langebaan-Saldanha area
would be of Early Pleistocene age.
In contrast, extensive Late Pleistocene beach deposits occur up to 7,0 ma.s.1.
on the west coast (e.g. Velddrif) and 7,2 m a.s.]. on the south coast (e.g. Coega
River mouth). The mollusc faunas that characterize these beach deposits have a
distinctly modern aspect and are identical with the present and adjacent open-
coast faunas. But preserved in Pleistocene estuarine and lagoonal deposits are
185
PLEISTOCENE MOLLUSCS FROM THE CAPE PROVINCE, SOUTH AFRICA
‘SOIJOU Ul OBULI [epl} JUSOId 0} JOJO SolIeNjso UI PU SUT[JseOd oY} SuOTe sjeIouINNY ‘deur AypeooT “[ “317
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186 ANNALS OF THE SOUTH AFRICAN MUSEUM
thermally anomalous fossil mollusc faunas. The intertidal molluscs and
ostracods all suggest a 6,5 m strandline.
The most characteristic feature of the fossil molluscs from the lagoonal and
estuarine facies is that they constitute mixed cool-water/warm-water assem-
blages. This mixed assemblage of taxa with mutually exclusive modern
geographic ranges is characterized by populations of warm-water molluscs that
existed in the last interglacial far south of their known present day geographic
range end-points. The cool-water element is similar to that of present day
open-coast faunas. The occurrence of the warm-water molluscs in the sheltered
environments is probably the result of a brief extension of their southerly range
during a relatively warmer part of the marine climatic cycle. Only in the sheltered
environments of the estuaries, far removed from the effect of cold oceanic
circulation, were the warm-water taxa able to establish reproductive populations.
Higher water temperatures than at present found in these sheltered environments
was probably the result of increased solar radiation (Tankard 1975). We
believe that this warm period coincides with the well documented period of
higher palaeotemperature at 120 000 B.P., the substage 5e of Shackleton (1969)
and Shackleton & Opdyke (1973). .
In the present paper the taxonomy of a number of problematical molluscs,
encountered during studies on Pleistocene deposits, are discussed. While some
have proved to be referable to species still extant elsewhere, several are clearly.
undescribed species.
Throughout this study height above sea level (a.s.l.) is referred to mean
sea level. Tidal ranges are shown in Figure 1. In the following text Natal Museum
has been abbreviated to N.M. and South African Museum to S.A.M. Where
possible, the elevation of sample localities is listed.
SYSTEMATIC DESCRIPTIONS
Class GASTROPODA
Family Trochidae
Cantharidus (Jujubinus) suarezensis (Fischer, 1878)
In analysing the systematics of this species, five names must be considered:
(1) Trochus suarezensis Fischer, 1878. This trochid appears to have remained
unknown to English workers, even though Dautzenberg (1929) implied it to be a
common Malagasy species. This is evidently the result of an error in Pilsbry’s
1889 translation of Fischer’s description, to which Dr Harald Rehder has kindly
drawn our attention. The word ‘geminatis’ (‘twinned’) of the original description
was evidently misread as ‘gemmatis’, which was rendered as ‘slightly granose’ in
Pilsbry’s text, and ‘distinctly granose’ in his comments. As a consequence the
number of spiral lirae was also incorrectly cited, the duplication of these
producing a total of twelve in the type, instead of only seven.
(2) Trochus fultoni Sowerby, 1889. Although this name appears in Sowerby’s
PLEISTOCENE MOLLUSCS FROM THE CAPE PROVINCE, SOUTH AFRICA 187
text, in the caption to his original plate he used the name Trochus stenomphalus.
Tomlin (1931: 419), acting as first reviser, selected T. fultoni as senior synonym.
This name was based on supposedly recent shells from Port Elizabeth. However,
like a number of other species described from there (e.g. Monilea ponsonbyi
(Sowerby, 1888), Loripes liratula (Sowerby, 1889) and Cerithium scabridum
rufonodulosum E. A. Smith, 1901), it does not appear to live in that region, and
the type material may be presumed to have been derived from a raised beach.
To this day specimens of most of these species, including ‘7.’ fultoni, are often
washed up on the shore on either side of the mouth of the Swartkops River,
which is here designated as the restricted type locality for ‘T.’ fultoni.
(3) Calliostoma farquhari Sowerby, 1892. While this was described as
coming from Port Elizabeth, a note has been left by H. C. Burnup to the effect
that John Farquhar had personally informed him that his shells were in reality
collected in Durban Bay. Thus the three Durban examples from the Ponsonby
collection in the British Museum, mentioned by Tomlin (1931), are possibly
syntypes. Similar shells, in more or less fresh condition, are common in Durban
Bay, although no living specimens have as yet come to the authors’ attention.
(4) Calliostoma bisculptum E. A. Smith, 1906. Described from a single
specimen from Durban.
(5) Calliostoma mosselense Tomlin, 1926. Described from Quaternary
deposits on the Klein Brak River.
Sowerby (1889) did comment on the resemblance of his Trochus fultoni to
T. suarezensis, but no direct comparison has hitherto been made. The inter-
relationships of the other taxa were discussed by Tomlin (1931) and Barnard
(1963a). The former synonymized T. farquhari with T. fultoni, but provisionally
accepted Calliostoma bisculptum as a valid species. Barnard treated Calliostoma
bisculptum, Calliostoma farguhari and Calliostoma mosselense all as synonyms of
Cantharidus fultoni. However, examination of large series from various localities
indicates that two morphologically distinguishable populations, isolated, as far
as can be determined, both temporally and geographically, can be distinguished.
These should be given subspecific rank. One population (fultoni) is restricted to
Pleistocene deposits of the southern Cape shoreline, the other is Recent,
occurring living from Natal to Tanzania and Madagascar. Specimens from the
Malagasy Republic (suarezensis), lent to us by Dr H. Rehder, agree well in all
characters (including dentition) with material from Durban and Mocgambique
(farquhari and bisculptum), and are clearly conspecific; Trochus suarezensis is
thus the earliest nomen applicable to this taxon.
Macnae & Kalk (1969: 127) and apparently Spry (1968: 6) have utilized the
name ‘Calliostoma interrupta (Wood) for examples of Cantharidus suarezensis.
Cantharidus (Jujubinus) interruptus (Wood, 1828), judging by Western Australian
specimens in the N.M., is indeed superficially similar, but has a completely
different colour pattern, is markedly narrower, and has more feebly pliculate
interstices.
Barnard’s figure (1963a: fig. 14i) of the radula of the present species is
188 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 2. Rachidian and lateral plates of Cantharidus suarezensis suarezensis (Fischer, 1878), with
intermediate plate of preceding row drawn in situ.
inaccurate, and the rachidian and lateral teeth are here refigured (Fig. 2).
In general the dentition closely resembles that of C. (J.) strigosus (Gmelin, 1791),
as figured by Fischer-Piette & Gailliard (1959: 59, fig. 1). As in many trochids
there is a degenerate, non-cuspidate plate between the laterals and marginals.
While it seems to be conventional to follow Troschel (1879) in regarding this as
an inner marginal tooth, it should be noted that this plate not infrequently bears .
traces of an alate side lamella, which is a characteristic of the lateral series of
teeth, not the marginals. In C. suarezensis and others this intermediate plate
seems to act as a buttress for the cutting head of the outermost lateral in the
succeeding row.
We are here relegating Jujubinus Monterosato, 1884, to subgeneric rank
under Cantharidus Montfort, 1810. Although maintained as a full genus by some
recent authors, Jujubinus appears to differ only in its more angular body whorl
and narrower spire angle.
Cantharidus suarezensis suarezensis (Fischer, 1878)
Fig. 3
Trochus suarezensis Fischer, 1878: 63; 1879: 378, pl. 115 (figs 2-2a).
Cantharidus suarezensis: Pilsbry, 1889: 130, pl. 45 (fig. 55). Dautzenberg, 1929: 332.
Calliostoma farquhari Sowerby, 1892: 43, pl. 2 (fig. 42). Syn. nov.
Caliiostoma bisculptum E. A. Smith, 1906: 54, pl. 8 (fig. 4). Syn. nov.
Cantharidus fultoni (partim): Barnard, 1963a: 281, figs 141, 19.
Calliostoma interrupta (non Wood, 1828): Spry, 1968: 6. Macnae & Kalk, 1969: 127.
Diagnosis
Distinguished by the thin, flat-topped spiral lirae, frequentiy arranged in
pairs, particularly medially; these lirae generally number 11-16 on the pen-
ultimate whorl, more rarely as few as 6 (fide Barnard 1963a). Intervals with
delicate, oblique axial plicules, which barely cross the intervening spirals.
PLEISTOCENE MOLLUSCS FROM THE CAPE PROVINCE, SOUTH AFRICA 189
Colour variable, often light green with articulated darker green dots on the lirae,
which may, by confluence, form oblique dark green lines or flames; in others the
ground colour is greenish-yellow or greenish-white with dark grey or olive-brown
axial flames; these often bifurcate at the basal periphery, which may bear red
marks; the intervals between the spiral lirae are characteristically orange-red,
although an occasional interval may lack this colour.
Dimensions
Holotype: 18 x 13 mm. B/Ht 0,72.
Durban Bay: 11,9 x 8,8 mm; 11,0 x 7,8 mm; 10,9 x 8,2 mm. B/Ht range
0,68-1,19.
inaaca, Island: 17,8 x 12,7 mm: 17.6 x 13,8 mm; 17,3 x 13,0 mm.
B/Ht range 0,63-1,07.
Malagasy: 11,7 x 9,5 mm; 11,5 x 8,8 mm; 11,5 x 8,6 mm. B/Ht range
0,74-0,81.
Fig. 3. Cantharidus suarezensis suarezensis (Fischer, 1878). B. Scanning electron photomicro-
graph showing sculpture on the base. (Inhaca Island.)
Distribution records
Natal: Durban Bay (N.M. ef auct.). Mocambique: Inhaca Island and
Delagoa Bay (N.M. and S.A.M.); Maxixe and Inhambane (S.A.M.); Bazaruto
and Benguera Islands and off Inhagondo region (N.M.); Mocambique Island
(Barnard and N.M.); Porto Amelia (N.M.). Tanzania: Dar-es-Salaam (N.M.).
Malagasy Republic: Nossi Bé (S.A.M.); Tulear and 13°23’S, 48°13’E (United
States National Museum); numerous other localities (Dautzenberg 1929).
190 ANNALS OF THE SOUTH AFRICAN MUSEUM
Barnard’s (1963a) Isipingo record is doubtful as there are no suitable
modern habitats nor raised beach deposits at that locality; and the specimen
dredged off Cape Morgan at 47 fathoms cannot be positively identified in the
South African Museum collection.
Habitat
On the sheltered mudflats on the west side of Inhaca Island and elsewhere
in Mocgambique C. s. swarezensis lives in abundance on the leaves of the marine
angiosperm Cymodocea ciliata (Forsk.) Ehrenb. ex Aschers., which forms
extensive beds along the infratidal fringe. Dautzenberg also records the species
as living ‘dans les Cymodocées’ in the Malagasy Republic.
Cantharidus suarezensis fultoni (Sowerby, 1889)
(Revised status)
Fig. 4
Trochus (Calliostoma) fultoni Sowerby, 1889: 153; 1892: 43, pl. 2 (fig. 43).
Trochus stenomphalus Sowerby, 1889: pl. 3 (fig. 7).
Calliostoma mosselense Tomlin, 1926: 81.
Cantharidus fultoni (partim): Barnard, 1963a: 281.
Diagnosis
Superficially smooth, but under magnification with fine declivous spiral:
lirae, never as strong as in the nominate subspecies, sometimes almost obsolete,
although always distinct in juveniles; these lirae are also always well developed
on the base. Oblique growth lines present, occasionally strong enough in places
A
Smm imm
Fig. 4. Cantharidus suarezensis fultoni (Sowerby, 1889). B. Scanning electron photomicrograph
showing sculpture on the base. (Swartkops River mouth.)
PLEISTOCENE MOLLUSCS FROM THE CAPE PROVINCE, SOUTH AFRICA 19]
to form fine plicules, which, however, always override the spiral sculpture,
instead of forming interstitial plicules such as characterize C. s. suarezensis.
Colour buff with oblique reddish or yellowish brown axial lines or spiral rows
of dots.
Dimensions
Swartkops River: 17,0 x 12,7 mm; 16,9 x 13,1 mm; 16,7 x 12,0 mm.
B/Ht range 0,72-0,94.
Kdeme Brak River: 12,1 x 8597 mm; 11,8 <x 8,3 mm; 11,3 x 7,8 mm.
B/Ht range 0,65—1,00.
Seageneld: 10/8 <x 7,5 mm; 955 < 7,3 mm; 9,0 x 7,6 mm. B/Ht range
0,69-0,85.
Distribution records
Type locality: Port Elizabeth, here restricted to the Swartkops River beds.
The known range of the subspecies is from Algoa Bay (Coega) to the Klein Brak
River, in beds of the 7 m level, and 4-5 m level. Swartkops River mouth (N.M..,
S.A.M.); Redhouse (N.M.); Coega River mouth (N.M., S.A.M.); Knysna
4,6ma.s.l. (S.A.M.); Sedgefield, 5 ma.s.l. (S.A.M.); Groot Brak River (S.A.M.);
Klein Brak River, 3,3 m a.s.l. (S.A.M.); Mossel Bay (N.M.).
Remarks
The diagnostic characters that serve to separate C. s. suarezensis and
C. s. fultoni have already been cited. In Figures 5A and 5B height is compared
with the ratio breadth/height for these two subspecies. It must be stressed,
however, that sampling was non-random; specimens were selected to present an
even distribution of size. As these graphs show, separation of the two subspecies
on shell dimension is not possible. The marked differences even within a single
subspecies may possibly be the result of habitat. For instance, as stated, the
Inhaca Island sub-population of C. s. swarezensis lives on the broad bladed ‘sea
grass’ Cymodocea ciliata (Forsk.). In Durban Bay Cymodocea is replaced by
Zostera, which is the probable habitat of the local C. s. suarezensis population.
By comparison with Cymodocea, Zostera is thin-leafed and much less robust.
Arguably, Zostera could not support broad specimens such as live on Cymodocea.
Figure 5 shows that the same difference in dimensions exists within C. s. fultoni,
but unfortunately the fossil record provides no indication as to which marine
angiosperms inhabited the Late Pleistocene estuaries and lagoons.
The differences in dimensions within each subspecies could equally be due
to the tendency for the development of local demes. Pelagic larval stages are
absent in European Cantharidus spp. and if this is applicable also to
C. suarezensis, it would contribute towards the isolation of sub-populations.
For example, there could be little gene exchange between the living Inhaca and
Durban Bay sub-populations, and between the Late Pleistocene Swartkops
River and Klein Brak River ones.
192 ANNALS OF THE SOUTH AFRICAN MUSEUM
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44. Cantharidus suarezensis fultoni— Swartkops
Fig. SA
Fig. 5. Comparison of height with breadth/height for Cantharidus suarezensis suarezensis and
C. s. fultoni. A. Scatter diagram. B. Calculated best-fit lines through the individual scatter
diagram populations.
20
Height mm
PLEISTOCENE MOLLUSCS FROM THE CAPE PROVINCE, SOUTH AFRICA
©
05 07 09
Breadth Height
e |. Cantharidus suarezensis suarezensis — Inhaca
02. Cantharidus suarezensis suarezensis— Durban Bay
x 3. Cantharidus suarezensis fultoni— Klein Brak River
44. Cantharidus suarezensis fultoni — Swartkops
Fig. 5B
193
194 ANNALS OF THE SOUTH AFRICAN MUSEUM
Family Stomatiidae
Pseudostomatella orbiculata (A. Adams, 1850)
Fig. 6
Stomatella orbiculata A. Adams, 1850: 31; 1854: 837, pl. 174 (figs 23-24). Sowerby, 1874: pl. 4
(fig. 23). Tomlin, 1923: 50.
Stomatella sp: Barnard, 1963a: 245, fig. 12a.
Distribution records
Knysna, 4,6 m a.s.l. (S.A.M.); Sedgefield, 5 m a.s.l. (S.A.M.); Klein Brak
River, 3,3 m a.s.l. (S.A.M.).
Remarks
Specimens from Pleistocene raised beaches (S.A.M.) in the Mossel Bay-—
Algoa Bay area agree well with Recent specimens (N.M.) from Mocambique
Island, Bazaruto and Santa Carolina Islands (Mogambique) and from Dar-es-
Salaam. The species was recorded from Algoa Bay by Tomlin (1923), but this
example was almost certainly washed out of a raised beach, as there is no other
record of Recent specimens from South African waters. There are in fact
numerous specimens from the Swartkops beds in the S.A.M. collection. The type
locality was Mocambique and it has been recorded from as far north as Ceylon
(Robertson 1969).
Barnard (1963a) described Pleistocene specimens of P. orbiculata in detail.
Fig. 6. Pseudostomatella orbiculata (A. Adams, 1850). A. Fossil (Knysna). B. Living (Dar es
Salaam).
PLEISTOCENE MOLLUSCS FROM THE CAPE PROVINCE, SOUTH AFRICA 195
Family Potamididae
Cerithidea (Cerithidea) bifurcata sp. nov.
Fig. 7
?Cerithium cf. guinaicum (non Philippi): Haughton, 1932: 45.
Type material
Holotype: SAM-K4496.
Paratypes | and 2: NM-A 80/T 1821.
Paratype 3: SAM-K4563.
Diagnosis
Apex decollate, later part of spire cyrtoconic, sutures narrowly canaliculate;
sculpture of fine axial riblets and well-developed subsutural crenules, crossed by
fine spiral lirae.
Description
Adult acuminately pupoid, with flaring aperture and decollate apex;
whorls gently convex, periphery situated at base of each whorl; sutures very
narrow but canaliculate. Aperture rounded quadrate, angular posteriorly, with
a small parietal denticle in the angle, labium evenly curved, margin of columella
callus free, but not erect; labrum evenly rounded, gently opisthocline; siphonal
canal short, very oblique, truncate, and deeply channelled. Axial sculpture
consists of fine, close, opisthocyrt riblets, about 35-41 in number on penultimate
whorl, rather weak towards back of lip, developing below the suture into strong
crenule-like nodules, 17—20 on the penultimate whorl. Spiral sculpture is present
anterior to these nodules, consisting of three narrow and sometimes very
shallow grooves on the penultimate whorl; their intervals are flattened, rarely
forming nodules at the intersections with axial ribs; spiral grooves are more
closely spaced on base of body whorl. One specimen retains traces of brown
coloration in the spiral grooves.
Adults (two ‘complete’ examples available) retain 53 whorls; no perfect
juveniles have been seen, but the only two immature specimens examined show
7 and 84 whorls respectively (their apices seem to be broken, however). The
exact stage at which decollation occurs cannot at present be estimated.
Dimensions
Holotype: adult, 20,6 x 10,1 mm.
Paratype |: adult, 21,6 x 10,7 mm.
Paratype 2: juvenile, 25,8 x 10,8 mm (labrum broken).
Paratype 3: juvenile, 19,9 x 7,7 mm.
Distribution records
Early Pleistocene beach deposits at 9,5 m a.s.l., 1,5 km north-east of
Langebaan in quarry (type locality).
196 ANNALS OF THE SOUTH AFRICAN MUSEUM
Remarks
The naturally decollate apex distinguishes C. bifurcata from most members
of the genus Cerithidea Swainson, 1840, certainly from all Recent Atlantic
species. The closest ally to C. bifurcata appears to be C. decollata (Linnaeus,
1758), a Recent Indo-Pacific species common in mangrove swamps in Natal, but
living among salt marsh vegetation in estuaries as far west as the Gamtoos River
(approx. 25°05’E). C. decollata differs in possessing regular suture-to-suture
axial ribs, a shallower siphonal canal, a definite posterior apertural angle, and
non-channelled sutures.
The pupoid shape and subsutural crenules of Cerithidea bifurcata may have
led to Tomlin Gn Haughton 1932) misidentifying it as the Recent tropical
Atlantic Cerithium guinaicum Philippi, 1849; in complete specimens shape and
sculptural details are very dissimilar.
8mm
Fig. 7. Cerithidea (Cerithidea) bifurcata sp. nov. Holotype. (Quarry 1,5 km north of
Langebaan.)
PLEISTOCENE MOLLUSCS FROM THE CAPE PROVINCE, SOUTH AFRICA 197
Family Cerithiidae
Cerithium scabridum rufonodulosum E. A. Smith, 1901
(Revised status)
Fig. 8
Cerithium mediterraneum (non Deshayes): Sowerby, 1892: 35. Turton, 1932: 125.
Cerithium rufonodulosum E. A. Smith, 1901: 108, pl. 1 (fig. 8). Barnard, 1963: 131.
Cerithium vulgatum (non Bruguiére): Bartsch, 1915: 116. Turton, 1932: 125.
Distribution records
4-5 m and 7 m levels: Coega River mouth (S.A.M.); Redhouse (S.A.M.);
Knysna, 4,6 m a.s.l. (S.A.M.); Sedgefield, 5 m a.s.l. (S.A.M.); Groot Brak River
(S.A.M.); Klein Brak River, 3,3 m a.s.l. (S.A.M.); Arniston (S.A.M.).
Remarks
As in the case of several other Cape Pleistocene molluscs, C. rufonodulosum
from Port Elizabeth was described as a Recent species. These specimens were
no doubt washed out of raised beaches along the banks of the Swartkops River.
The Port Alfred specimens recorded as C. vulgatum and C. mediterraneum were
also presumably derived from local deposits. Cerithium rufonodulosum closely
resembles Sowerby’s figure of his C. nigropunctatum (1855: 860, pl. 180, fig. 97)
from an unknown habitat, as was indicated by Smith. Unfortunately the original
description of C. nigropunctatum contains insufficient detail, and from Smith’s
phraseology it would appear that the types are lost. Reeve (1865a) omitted the
Fig. 8. A. Cerithium scabridum rufonodulosum E. A. Smith, 1901 (Algoa Bay). B.C. s. rufono-
dulosum E. A. Smith, 1901 (Sedgefield). C. C. s. scabridum Philippi, 1848 (Muscat, Gulf of
Oman).
198 ANNALS OF THE SOUTH AFRICAN MUSEUM
species, and Kobelt (1898: 242, pl. 42, fig. 10) merely copied Sowerby’s descrip-
tion and figure. C. nigropunctatum must therefore be rejected as a nomen
dubium, at least until the type (if extant) is located. The possibility does exist,
nevertheless, that it may indeed be based on a discoloured specimen of the
present species.
Of Recent species C. rufonodulosum appears to be allied only to Cerithium
scabridum Philippi, 1848. This species is recorded from the Red Sea, Persian
Gulf and India, south to the Mascarene Islands and Quirimba Island (12,4°S,
40,7°E), but has not yet been reported from farther south. The morphological
differences between C. scabridum and C. rufonodulosum are very small and we
believe that the relationship should be regarded as a subspecific one. Comparison
with specimens of C. s. scabridum from the Gulf of Oman (N.M.: F. Luther)
shows it to differ from C. s. rufonodulosum only in having less prominent
tubercles. While fresh examples of C. s. scabridum are pale greyish, sometimes
zoned with orange-brown, and with rows of articulated black and white marks
crossing the rows of tubercles, an old, dead specimen is light brownish-buff
with dark brownish-orange tubercles, exactly as in most South African
C. s. rufonodulosum (some examples are tinged violet), suggesting that the
coloration to be seen in specimens of the latter is secondary.
A syntype of C. s. rufonodulosum (NM-1041/T512) measures 22,5 x
8,7 mm. It is here designated as lectotype; although the labrum is broken it is
otherwise in good condition.
Family Calyptraeidae
Crepidula capensis praerugulosa subsp. nov.
Fig. 9
Crepidula rugulosa Dunker, 1846 (partim). Barnard, 1963: 72, fig. 9f.
Type material
Holotype: SAM-K4564.
Paratypes 1-8: NM-9229/1825.
Paratypes 9-11: SAM-K4580.
Diagnosis
A Pleistocene chronosubspecies of Crepidula capensis Quoy & Gaimard,
1835, differing in its marginal apex and larger size, and in the absence of rugose
sculpture.
Description
Outline curved-pyriform, breadth variable, apex spirally coiled, situated on
postero-lateral margin, protoconch frequently retained in moderately large
specimens; outer surface with regular, well-marked growth lines. Septum as in
Crepidula capensis, i.e. margin sigmoid, shallowly concave on right, with a
prominent lobe to the left of the midline, and a deep, narrow sinus on the left side.
PLEISTOCENE MOLLUSCS FROM THE CAPE PROVINCE, SOUTH AFRICA 199
Dimensions
Holotype: 27,1 x 21,7 mm.
Paratypes: 27,7 <x 18,2 mm;
35,6 x 31,0 mm.
Distribution records |
Late Pleistocene: found extensively in exposures of 7 m beach adjacent to
present open coast between Elands Bay and Saldanha Bay; Kruispad, 4,6 m
a.s.l. (S.A.M.); Velddrif West, 4-7 m a.s.l. (S.A.M.) (type locality); Milnerton
lagoon (S.A.M.).
Remarks
Although many species of Crepidula are notoriously variable in shell shape,
the numerous specimens of C. c. praerugulosa show its characters to be relatively
constant, save for some variation in proportions as discussed below. While it is
always distinguishable from C. capensis Quoy & Gaimard, 1835 (syn. C. rugulosa
Dunker, 1846, cf. Kilburn, 1974), the form of the septum and general shape of
C. c. praerugulosa agree very closely with that species. C. capensis does not
occur in the same deposits, and it seems advisable to rank C. c. praerugulosa as
a chronosubspecies rather than as a full species.
The regularly curved ventral margin of C. c. praerugulosa suggests that it
lived attached to mussel shells, a common habitat of Crepidula porcellana
Lamarck, 1801. C. c. capensis appears to live entirely on the undersides of rocks.
Indeed the wide range of proportions found in C. c. praerugulosa (breadth
ranging between 0,66 and 0,87 of length) somewhat parallels that of C. porcellana.
In the latter species the broader form generally occurs in individuals attached to
the wider, flatter, posterior part of the bivalve surface, while the narrower form
generally lives on the strongly curved anterior or umbonal part. Possibly the
same may have applied to C. c. praerugulosa. One paratype even shows a series
i@mm 10mm
Fig. 9. Crepidula capensis praerugulosa subsp. nov. Holotype. (Velddrif West.)
200 ANNALS OF THE SOUTH AFRICAN MUSEUM
of xenomorphic ridges, strongly suggesting attachment to the ribbed mussel
Aulacomya ater (Molina, 1782).
In addition to the type series of C. c. praerugulosa, a set of eight specimens
from an older deposit 1,5 km north of Langebaan (9,5 m a.s.l.) has been
examined. These are in a very poor state of preservation, but are morpho-
logically most interesting. Not only is the apex in these specimens more terminal
than in typical C. c. praerugulosa, and the shells on the average more compressed,
but the left side of the septum appears to be less lobate. This might be construed
as indicating an origin from a porcellana-type ancestor. Much more and better
material is needed, however.
Family Buccinidae
Triumphis dilemma sp. nov.
Fig. 10
Type material
Holotype: SAM —K4565.
Paratype 1: NM—A1217/T1827.
Diagnosis
Anal canal well developed; shell covered by thin spiral lirae with feeble
axial folds on the spire.
Description
Ovate-fusiform, aperture nearly twice length of spire, whorls moderately
convex, aperture constricted, labrum thickened, expanded, anal canal trough-
like, rendering the lip shouldered; labrum with ten internal ridges, terminating
in denticles of which the anterior two are partially fused; labium with a
5mm 8mm
Fig. 10. Triumphis dilemma sp. nov. A. Holotype. B. Paratype 2. (Quarry 1,5 km north of
Langebaan.)
PLEISTOCENE MOLLUSCS FROM THE CAPE PROVINCE, SOUTH AFRICA 201
moderately wide, flattened callus, which is continuous with that of the labrum;
paries with one or two denticles, columella with three basally; fasciole weak.
Sculptured by tabulate spiral cords, with sharply incised intervals; cords 5—6 in
number on penultimate whorl, 17—18 on body whorl, median ones on last whorl
mostly shallowiy bisected; intervals between cords with fine axial plicules; spire
with obscure traces of axial folding. Apex worn, four teleoconch whorls
remaining.
Dimensions
Holotype: 16,3 x 10,4 mm.
Paratype 1: 14,8 x 9,3 mm.
Paratype 2: 24,9 x 16,0 mm.
Distribution records
Early Pleistocene at Langebaan, 9,5 m beach (type locality).
Remarks
Although the thickened labrum and fully-developed apertural characters
suggest that the type specimens are adult, two larger specimens from the same
deposit are probably conspecific. The latter are, however, in poor condition
(one being fragmentary), and there are no intermediate specimens to connect
the two extremes. They are therefore excluded from the type material.
Although similar in general shape, proportions and sculpture, these larger
individuals show reduced development of the anal canal, less constricted
apertures and reduced lip denticles. Thus the labrum is lirate internally, without
denticles, the columella pustules are weak, and the paries bears only one low
tubercle. On the body whorls of these two specimens there are 25-34 spiral
ridges. The more complete specimen (four teleoconch whorls remaining, apex
worn) measures 25 x 16,3 mm.
The affinities of Triumphis dilemma are not clear. In some respects it
resembles species of the genus Cantharus Roding, 1798, but in apertural
characters is closer to Triumphis Gray, 1857. It differs from the Recent Panamic
T. distorta (Wood, 1838) in shape, in the presence of sculpture over the whole
surface, and in the less developed anal canal.
Family Terebridae
Duplicaria otiosa sp. nov.
Fig. 11
Type material
Holotype: NM-8185/T1832.
Paratype: SAM-K4567.
Diagnosis
Small (15,0 mm), with 11 strong, rounded axial ribs with sloping sides,
shallowly cut by a subsutural groove, demarcating a series of oblong subsutural
nodules; no other spiral sculpture.
202 ANNALS OF THE SOUTH AFRICAN MUSEUM
Description
Acuminate, 7 teleoconch whorls; whorls strongly convex; spire about twice
length of aperture; siphonal fasciole strongly twisted and carinate, columella
thinly calloused, aperture elongately ovate-rhombic. Subsutural groove situated
one-third of whorl length from upper suture, shallow, incising axial ribs, but
barely cutting interstices; only visible from the 4th whorl onwards. Axial ribs
extending from suture to suture, feebly opisthocyrt, rounded with gently sloping
sides, narrower than intervals, strongly developed throughout, 11-12 in number
on first teleoconch whorl, 11 on penultimate whorl; posterior part of each rib
(cingulum), where cut off by the sulcus, is axially oblong, not noduliform.
No sign of spiral striae, but growth lines conspicuous. Protoconch worn,
bluntly conical, of about 24 whorls.
Dimensions
Holotype: 14,5 x 4,5 mm.
Paratype: 16,4 x 4,6 mm, labrum broken.
5mm
Fig. 11. Duplicaria otiosa sp. nov. Holotype. (Coega salt works.)
PLEISTOCENE MOLLUSCS FROM THE CAPE PROVINCE, SOUTH AFRICA 203
Distribution records
Pit at Coega salt works, 4,5 m a.s.l. (type locality).
Remarks
Closely allied to Strioterebrum (Partecosta) wilkinsi Dance & Eames, 1966,
from the Recent Hammar Formation of Iraq. Duplicaria wilkinsi differs in its
more nodular subsutural band, in the development of the subsutural groove on
the early whorls and in its convex whorl profile. Cernohorsky (1969: 221) lists
D. wilkinsi as a synonym of the unfigured Terebra fuscobasis E. A. Smith, 1877.
However, Melvill & Standen (1917: 208) noted the presence of spiral microstriae
in the latter, these being apparently absent in both Duplicaria wilkinsi and
D. otiosa.
Of species recorded from South Africa D. otiosa resembles only D. evoluta
(Deshayes, 1859) from Natal and the Indo-Pacific; this is a much larger species
(30-40 mm), with a bigger protoconch and a finely punctate subsutural groove,
with the axial ribs shouldered where they border the latter.
The radula of D. otiosa will never be known, and we are referring it to the
genus Duplicaria Dall, 1908, on account of its general resemblance to Recent
species such as D. evoluta and D. fictilis (Hinds, 1845).
Family Actaeonidae
Pupa (Strigopupa) daviesi sp. nov.
Fig. 12
Actaeon (Solidula) suturalis (non A. Adams): Sowerby, 1892: 52.
?Actaeon pudica (non A. Adams): Turton, 1932: 2.
Solidula sulcata (non Gmelin, 1791): Barnard, 1962: 192 (partim); 1963a: 316.
?Solidula suturalis: Barnard, 1963a: 316.
Type material
Holotype: SAM-K4568.
Paratypes: NM-—A1218/T1828;
NM-A1219/T1827;
NM-8186/T1824.
Diagnosis
Narrowly pupoid, lip flattened, left side of base obliquely truncate, columella
with a single bifid fold, no parietal tubercle, sculpture of low, tabulate spiral
ridges; axial colour lines visible under ultraviolet light.
Description
Narrowly pupiform, with acute, cyrtoconical spire, sutures moderately deep
but not channelled, whorls gently convex; profile of labrum somewhat flattened
in middle, basally initially sharply rounded, becoming obliquely truncate on left
204 ANNALS OF THE SOUTH AFRICAN MUSEUM
side, which lies in plane of curvature of left side of body whorl profile. Columella
with a single strong fold, shallowly bisected by a groove, paries fairly straight
above, strongly curved below, basal parietal tubercle absent. Surface sculptured
by low, flat-topped spiral ridges separated by narrow grooves; first teleoconch
whorl with three spiral cords, increasing to 7-11 on the fourth whorl; in the
middle of the body whorl these ridges tend to be bisected by median grooves,
and on the base split into thinner and more widely spaced lirae. Growth lines
regular and rather coarse, crossing spiral cords and forming fine plicules in the
intervals, rendering these superficially foveolate.
Shell always bleached, but under U.V. illumination pigments fluoresce to
produce a negative pattern of numerous thin, wavy axial lines, the posterior
ends of which tend to fuse below the suture.
Dimensions
Holotype: 14,6 x 6,8 mm.
Paratypes: 21,6 x 10,0 mm;
20,8 x 9,4 mm;
20,8 < 9,3 mm.
Distribution records
Coega salt works, 4-7 m a.s.l. (N.M.); Redhouse, 7 m a.s.l.; The Creek, '
5,7 m a.s.l.; Knysna, 4,6 m a.s.l. (S.A.M.) (type locality); Sedgefield, 5 m a.s.1.
(S.A.M.); Klein Brak River, 3,3 m a.s.l. (N.M., S.A.M.). See Davies (1972) for
site information.
Remarks
Pupa daviesi was initially reported from South Africa as the Philippine Pupa
suturalis (A. Adams, 1855), as indicated by N.M. material originally from the
Crawford collection, on specimens from which Sowerby based his 1892 record.
Some of these were subsequently sent by Henry Burnup to E. A. Smith as
P. ‘ ?suturalis’, and were returned as P. ‘solidula var.’. J. R. le B. Tomlin, who saw
the same material, disagreed with both identifications, but did not suggest an
alternative. We agree that neither P. suturalis nor P. solidula is at all similar to
the present species. Adams (1855: 61) in his original description of P. suturalis
stressed the conspicuously channelled suture, this being shallow in P. daviesi.
P. solidula (Linnaeus, 1758) is a large (20-30 mm), globose species with very
different columella folds and a rounded base. P. daviesi was probably the species
that was recorded from Port Alfred beach drift as Acteon pudica (A. Adams,
1855) by W. H. Turton. Judging by Reeve’s figure (1865a: pl. 3 fig 13) this is a
true Acteon (with simple columella pleat); otherwise it does resemble Pupa
daviesi in form and sculpture, but has a wider aperture and a rounded base.
P. daviesi is referable to the subgenus Strigopupa Habe, 1958, which includes
only a few species, all characterized by their obliquely truncate base and
PLEISTOCENE MOLLUSCS FROM THE CAPE PROVINCE, SOUTH AFRICA 205
strigate colour pattern. Other species referable here are P. strigosus (Gould,
1859), P. affinis (A. Adams, 1855) and P. fumata (Reeve, 1865). P. strigosus from
Japan appears to have a far more prominent double columella fold than any of
the others; it shows a well-developed parietal denticle, and has the groove
separating this from the columella fold markedly reduced; spiral sulci are fewer
than in P. daviesi. P. affinis (of which P. fumata should be regarded as a synonym)
is even closer to P. daviesi, differing only in possessing a parietal tubercle and a
more strongly bisected columella fold; although generally slightly narrower than
P. daviesi, there is some overlap in this respect. P. affinis at the present day lives
as far south as Durban Bay (it is the species misidentified by Barnard (1963a:
316) as Solidula sulcata (Gmelin, 1791)). In all probability Pupa daviesi originated
as an isolated deme of the P. affinis population, but in view of the structural
differences we believe that it should be given full species status. This species has
been named in honour of Professor O. Davies, collector of much of the Natal
Museum Pleistocene material used in this study.
5mm
Fig. 12. Pupa (Strigopupa) daviesi sp. nov. Holotype. (Knysna.)
206 ANNALS OF THE SOUTH AFRICAN MUSEUM
Class BIVALVIA
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Fig. 13
Nucula (Leda) bicuspidata Gould, 1845: 37.
Nucula largillierti Philippi, 1851: 87.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (figs 8a—b).
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110 (further references).
Description
Cape Pleistocene material: nuculaniform, rostrum conspicuous but short,
very truncate; shell thin; umbo anterior to midline, antero-dorsal and anterior
margins well rounded, ventral margin gently convex, postero-dorsal margin long
5mm
Fig. 13. Nuculana (Lembulus) bicuspidata (Gould, 1845). (Kruispad, Velddrif.)
PLEISTOCENE MOLLUSCS FROM THE CAPE PROVINCE, SOUTH AFRICA 207
and slightly concave. Surface sculpture of regular, oblique, sharply incised ribs,
becoming concentric anteriorly, growth lines faint. Posterior end with a promi-
nent broad sulcus, bordered in front by a sharp umbonal ridge, and behind by a
broader dorsal ridge, rendering posterior margin bicuspidate; this sulcus is
crossed by lunulate, comarginal ridges, which form small squamose denticles
along the crest of the umbonal ridge, and two similar rows on the dorsal ridge,
separated by a shallow furrow. Anterior end bearing a low, narrow umbonal
ridge, bordered behind by a shallow concavity, rendering the margin sinuous at
this point; surface anterior to the ridge sometimes with 1 or 2 feeble radial lirae.
Resilifer small, triangular; hinge teeth chevron shaped, at shell length of 18 mm
with 25 teeth posteriorly, 21 anteriorly. Lunule concave, lanceolate, extending to
posterior end, bearing growth lines only. Muscle scars and pallial line very
weakly impressed, pallial sinus short, ascending, not confluent with pallial line.
Dimensions
ie < 9:5 mm, 17,1 x 9,4 mm, 16;6 x 9,3 mm.
Distribution records
Redhouse (S.A.M.); Kruispad, 5,1 m a.s.l. (S.A.M.); Milnerton (S.A.M.).
Remarks
Pleistocene material agrees well with published figures and with a Recent
N.M. specimen from Senegal. The species at the present day ranges from
Mauritania to Angola (Nicklés 1950). A closely allied species, NV. gruveli Nickles,
1952, from the Quaternary of Gabon, differs in having a weaker posterior
sulcus, a more central umbo, and in being less truncate posteriorly.
The only two comparable species known from South Africa are N. lamellata
Sowerby, 1904, and N. gemmulata Sowerby, 1904, both Recent species from
moderately deep water off Natal/Zululand. In these two the posterior sulcus is
traversed by a median ridge or ridges, and there is no anterior umbonal ridge.
Family Ungulinidae
Felania diaphana (Gmelin, 1791)
Fig. 14
Venus diaphana Gmelin, 1791: 3292.
Lucina adansoni Reeve, 1850 (non Orbigny): pl. 9 (fig. 51).
Lucina senegalensis Reeve, 1850: appendix.
Felania diaphana: Recluz, 1851: 71. Chavan, 1962: 5.
Felania rosea Recluz, 1851: 72, pl. 2 (figs 10-12).
Diplotodon [sic] cf. senegalensis: Smith in Rogers, 1906: 294.
Diplodonta (Felania) diaphana: Lamy, 1920: 371 (references). Fischer-Piette, 1942: 317,
pl. 14 (fig. 7) (holotype). Nicklés, 1950: 188, fig. 351.
? Diplodonta (Felania) agulhasensis Thiele & Jaeckel, 1931 (partim): 219. Barnard, 1964: 468.
Diplodonta cf. senegalensis: Van Hoepen, 1940: 191. Barnard, 1962: 185; 1964: 463.
Ungulina alba (non Rang): Fischer-Piette in Davies, 1972: 254.
Distribution records
Cenozoic: Swartkops River (S.A.M.); Coega salt works, 7,1 m a.s.l;
Redhouse, 7,2 m a.s.l.; Cradock, 13,5 m a.s.].; Deal Party, 5,1 m a.s.I.; Knysna,
208 ANNALS OF THE SOUTH AFRICAN MUSEUM
10mm 10mm
Fig. 14. Felania diaphana (Gmelin, 1791). (Mossel Bay.)
4,6 m a.s.l. (S.A.M.); Sedgefield, 5 m a.s.l. (S.A.M.); Klein Brak.River, 3,3 m
a.s.l. (S.A.M.); Mossel Bay (N.M.); Cape Cross Bay, S.W.A. (N.M.). See Davies
(1972) for site descriptions.
Remarks
This Recent West African species is fairly common in raised Pleistocene
beaches along the south Cape coast..Such specimens agree closely with figures
of Felania diaphana, and with valves from the Cuanza River mouth, 75 km south
of Luanda, Angola (N.M.: B. R. Stuckenberg). A rather fresh right valve from
Cape Cross Bay, South West Africa (N.M.: O. Davies) may indicate the Recent
occurrence of the species within South African limits. However, Thiele &
Jaeckel’s record (1931: 219) of the species (as Diplodonta (Felania) rosea) from
deep water on the Agulhas Bank requires confirmation. The species at present is
known to range from Mauritania to Angola (Nicklés 1950).
Family Lucinidae
Loripes (Microloripes) liratula (Sowerby, 1889)
Fig. 15
Lucina (Divaricella) liratula Sowerby, 1889: 155, pl. 3 (fig. 5); 1892: 61, pl. 2 (fig. 63).
Lucina liratula: Haughton, 1932: 35.
Lucina contempta (non Cossman): Turton, 1932: 235, pl. 62 (fig. 1641).
Divaricella liratula: Barnard, 1964: 479.
Description
Subcircular, moderately compressed; sculptured by growth lines and fine
acentric lirae, which are rather irregular, but tend to be somewhat oblique
across the middle, distinctly undulating at each end. Lunule small, ovate,
PLEISTOCENE MOLLUSCS FROM THE CAPE PROVINCE, SOUTH AFRICA 209
concave, wider in right valve than left. Resilifer deep, internal, oblique. Hinge
plate narrow; right valve with a strong, elevated anterior lateral tooth, a
ridge-like posterior lateral, partly overlying distal end of resilifer, and a single
oblique cardinal; left valve with anterior and posterior ridge-like laterals and a
cuneiform anterior cardinal with a low posterior cardinal ridge adjacent to it.
Anterior adductor scar flexuous. Internal margin crenulate.
Dimensions
ie ti mm, 16:2 < 16,5 mm, 15,2 x 15,4 mm.
Distribution records
Swartkops River (S.A.M.); Cradock, 13,5 m a.s.l.; Coega salt works,
7,1 m a.s.l. (S.A.M.); Knysna, 4,6 m a.s.l. (S.A.M.); Sedgefield, 5 m a.s.l.
(S.A.M., N.M.); Groot Brak River (S.A.M.); Klein Brak River, 3,3 m a.s.l.
(S.A.M.); Verlorevlei, 4 m a.s.l. (S.A.M.); Kruispad, 5,1 m a.s.l. (S.A.M.);
Bloemendal (S.A.M.); Geelbek, 4,5 m a.s.l. (S.A.M.); Churchhaven, 1,5 m and
4,3 maz.s.l. (S.A.M.).
Remarks
The present species was described as a Divaricella, but its oblique internal
ligament shows it to be a Loripes. We follow Chavan (1937) in treating Micro-
loripes as a subgenus of Loripes, in preference to his 1969 system under which it
is transferred to the genus Parvilucina. In Parvilucina the ligament is sunken, but
not internal, and lies parallel to the dorsal margin.
Descriptions and figures strongly suggest that Loripes liratula is a synonym
of the Recent West African Loripes (Microloripes) contrarius (Dunker, 1846).
However, as no comparative material is available, we are forced to retain
Sowerby’s name pro tem. Although described and recorded as a Recent species,
L. liratula appears to be extinct in South Africa, beach specimens being derived
from Pleistocene deposits.
5mm
Fig. 15. Loripes (Microloripes) liratula (Sowerby, 1889). (Kruispad, Velddrif.)
210 ANNALS OF THE SOUTH AFRICAN MUSEUM
Family Veneridae
Pitar (Lamelliconcha) madecassina (Fischer-Piette & Delmas, 1967)
Cytherea manillae (non Sowerby, 1851): Sowerby, 1897: 24.
Pitaria manillae Barnard, 1964: 503. Boshoff, 1965: 164, pl. 10 (fig. 2).
Dosinia (Sinodia) madecassina Fischer-Piette & Delmas, 1967: 12, pl. 3 (figs 17-19).
Pitar (Lamelliconcha) madecassina: Fischer-Piette, 1968: 789, pl. 1 (figs 3-4).
Distribution records
Cenozoic: Cradock, at head of Papkuils estuary, 15,2 m a.s.]. (N.M.).
Remarks
Fischer-Piette (1968) compared Pitar madecassina with P. manillae, and
recorded the former from various localities in Mocgambique and Natal. He did
not, however, suggest that this was the species that had previously been reported
from South Africa as P. manillae. P. madecassina ranges from the type locality,
Madagascar, south to Pondoland (Mzamba, N.M.; Port St Johns, S.A.M.).
Although articulated specimens are rarely washed up on the shore, the species
lives in abundance at depths of 12-50 fathoms off Natal.
Barnard (1962: 185; 1964: 503) erred in regarding ‘Chamelea’ schwarzi
Newton, 1913 (syn. ‘C.’ rogersi Newton, 1913, first reviser Barnard, 1962) as
being based on worn P. madecassina. This late Cenozoic species differs con-
spicuously from P. madecassina in its compressed valves, oblong-ovate shape,
moderately curved umbo and weak umbonal ridge; also the anterior and median |
cardinals of the left valve only intersect at the very dorsal margin. P. madecassina
is tumid, ovate-trigonal, posteriorly subrostrate with a strong umbonal ridge,
umbo strongly prosogyrate, left anterior and median cardinals intersecting some
distance below the dorsal margin. ‘Chamelea’ krigei Newton, 1913, is even more
different in form and dentition.
Venerupis dura (Gmelin, 1791)
Fig. 16
Venus dura Gmelin, 1791: 3292.
Tapes dura var. simili [sic] (? non similis Deshayes, 1853): Schwartz, 1910: 116.
Venerupis (Polititapes) dura: Nicklés, 1950: 203, fig. 387.
Macrocallista lilacina (non Lamarck, 1818): Barnard, 1964: 505.
Venerupis dura: Fischer-Piette & Meétivier, 1971: 11 (references and synonymy).
Venerupis rufiscensis Fischer-Piette & Métivier, 1971: 13, pl. 3 (figs 2-7). Syn. nov.
Distribution records
Cenozoic: Redhouse (S.A.M.); Knysna, 4,6 m a.s.l. (S.A.M.); Geelbek
(S.A.M.); Kruispad, 5,1 m a.s.l. (S.A.M.); Verlorevlei, 4 m a.s.l. (S.A.M.).
Remarks
Venerupis dura is a West African species, living as far south as Luanda
(Angola), but also abounding in Late Pleistocene sediments of the south-western
and southern Cape. This extinct deme differs morphologically from the Recent
PLEISTOCENE MOLLUSCS FROM THE CAPE PROVINCE, SOUTH AFRICA in
25mm
25mm
Fig. 16. Venerupis dura (Gmelin, 1791). (Kruispad, Velddrif.)
form in its weak to almost obsolete concentric sculpture; while in most specimens
shallowly incised concentric grooves are present, in some examples coarse
growth lines are the only external surface feature. In addition the pallial sinus
is usually somewhat narrower and sharper than in Recent Angolan V. dura.
Mr C. P. Nuttall, who kindly examined specimens from Verlorevlei, has
suggested (in /itt. 26 August 1971) that these may be comparable with ‘Callisto-
tapes vetulus (Basterot) var. plioglabroides Sacco from the Pliocene (Astian) of
Piedmont’, Italy.
The S.A.M. specimens recorded by Barnard as Macrocallista lilacina
(Lamarck, 1818) [= Callista spuma (Réding, 1798)] are referable to Venerupis
dura.
We believe Venerupis rufiscensis to be based merely on coarsely-ribbed
examples of V. dura. The range of supposed differences between the two is no
greater than that found in the present material. Two Recent Angolan shells
(N.M. coll.) even appear to combine the ridge number of V. dura with the ridge
profile of V. rufiscensis.
We are following Fischer-Piette & Métivier (1971) in referring this species
to Venerupis Lamarck, 1818. However, despite the gradually ascending pallial
212 ANNALS OF THE SOUTH AFRICAN MUSEUM
sinus, its true affinities (as suggested by shape, sculpture, colour and hinge
structure) probably lie not with Venerupis but with Paphia Réding, 1798.
Differences between Paphia, Venerupis and Tapes Von Mihlfelt, 1811, are,
however, small and possibly artificial. The species at present ranges from
southern Morocco to Angola (Nicklés 1950).
Family Petricolidae
Petricola (Claudiconcha) prava sp. nov.
Fig. 17
Type material
Holotype: SAM-K4569.
Paratypes 1-3: 1 right, 2 left valves, NM—A79/T1820.
Paratype 4: right valve, SAM-K4570.
Paratype 5: right valve, SAM-K4571.
Diagnosis
Shell relatively large (40-50 mm), inequivalve, always deformed and
irregular; sculptured by fine radial lirae, often only visible posteriorly, and
coarse, irregular, non-lamellar growth lines; pallial sinus very deep.
Description
Shape very irregular and always more or less deformed, often somewhat
rostrate posteriorly with the ventral margin concave; valves thick and very
convex; umbo orthogyrate, strongly curved, varying in position from submedian
to less than one-third of distance from anterior end. Sculpture of fine radial
lirae, often only visible posteriorly, and coarse, irregular growth lines, sometimes
elevated posteriorly, but not lamellar.
Right hinge with two peg-like cardinal teeth, of which the posterior one
may be shallowly bisected by a groove. Left hinge with three cardinals of which
the anterior is reduced to a small denticle, the central is peg-like, triangular in
cross-section, and medially concave below (but not bifid), and the posterior one
is an oblique lamella partly fused to the nymph. Nymphs are very large, some-
times massive, more or less projecting and spathulate, separated from valve
margin by a deep groove. Pallial sinus free, apically rounded, very deep,
extending well below midline.
Dimensions
Holotype: left valve, 49,9 x 37,2 mm;
right valve, 52,0 x 37,4 mm;
breadth of valves together (inflation), 31,3 mm (posterior end
chipped).
Paratypes: single left valve, 51,0 x 36,5 mm;
single right valve, 49,3 x 39,9 mm.
PLEISTOCENE MOLLUSCS FROM THE CAPE PROVINCE, SOUTH AFRICA 213
20mm
10mm
Fig. 17. Petricola (Claudiconcha) prava sp. nov. A. Holotype. B. Paratype 4. A (lower), B,
C show variation in shell form. (Quarry 1,5 km north of Langebaan.)
214 ANNALS OF THE SOUTH AFRICAN MUSEUM
Distribution records ,
Smitswinkelbaai (Saldanha), behind Sea Harvest factory, 6,5-8 m a.s.1.;
quarry 1,5 km north of Langebaan, 9,5 m a.s.l. (type locality). Early Pleistocene.
Remarks
The markedly inequivalve shell of Petricola prava shows it to be referable
to the subgenus Claudiconcha Fischer, 1887. Of described members of the
taxon, Petricola japonica Dunker, 1882, from Japan, P. quadrasi (Hidalgo, 1886)
from the Philippines and Indonesia, P. cumingi Deshayes, 1853, from South
Australia, and P. chinensis Deshayes, 1853, from China and the Philippines,
differ in possessing strong concentric sculpture without radials. On the other
hand, while P. monstrosa (Gmelin, 1791) from Nicobar and the Ryukyu Islands,
and P. madreporica (Jousseaume, 1895) from the Red Sea, do show dominant
radial lirae as in P. prava, in these the lirae are stronger and more uniformly
developed, and shell size is markedly smaller. The closest species is probably
P. robusta Sowerby, 1834, from tropical west America, which is usually regarded
as a Rupellaria, but has the unequal valves of Claudiconcha; this differs from
Petricola prava in its somewhat weaker radials and much shallower pallial sinus.
The distinctly inequivalve shell and large size distinguish P. prava from the
three Recent South African species, Petricola (Rupellaria) bicolor Sowerby, 1854,
P. (Petricola) ponsonbyi Sowerby, 1892, and P. (P.) divergens (Gmelin, 1791).
The latter two also differ in their divaricate sculpture. P. bicolor is the most.
similar, as not only are some individuals very slightly inequivalve, but a rock-
boring morph which occurs in South West Africa is frequently deformed in
much the same manner as P. prava. However, the stronger radial sculpture and
shallower pallial sinus of P. bicolor, together with the characters cited above, are
diagnostic.
Family Donacidae
Donax sanctuarium sp. nov.
Fig. 18
Type material
Holotype: right valve, SAM—K4572.
Paratypes: 1 right, 1 left valve, NM—A1216/T1826;
2 right, 2 left valves, SAM—K4573.
Diagnosis
Moderately large (40-60 mm); oblong-trigonal, posteriorly truncate,
umbonal ridge feeble; smooth, posterior end with concentric ridges, and a series
of interstitial radials just behind the umbonal ridge; inner margin smooth.
Nymphs very prominent, lateral teeth present, except for the right anterior one,
which may be absent.
Description
Oblong-trigonal, length 2,1 x height, umbo approximately one-third
distance from posterior end; antero-dorsal margin straight, anterior end well-
PLEISTOCENE MOLLUSCS FROM THE CAPE PROVINCE, SOUTH AFRICA 215
15mm
Fig. 18. Donax sanctuarium sp. nov. A. Holotype. B. Paratype. (Churchhaven.)
rounded, postero-dorsal margin convex, declivous, posterior end sharply
rounded, ventral margin gently and evenly curved, ends narrowly gaping,
posterior end with a very slight umbonal ridge, but no distinct angle, posterior
face moderately truncate. Greater part of shell smooth, except for growth lines,
posterior face corrugated by fine, wavy, concentric ridges, with a narrow area
adjacent to the umbonal ridge also traversed by interstitial radial riblets; inner
ventral margin of valves smooth. Hinge of left valve with two narrow, subequal,
divergent cardinals, a low ridge-like anterior lateral tooth situated close to the
cardinals, and a rather remote and stronger ridge-like posterior lateral. Hinge of
right valve with a somewhat trigonal posterior cardinal, somewhat variable in
shape, but always bisected by a shallow groove, occasionally weakly bifurcate;
anterior cardinal long, oblique and ridge-like, anterior lateral feeble or absent,
posterior lateral strong and elongate, bordered dorsally by a trough. Nymph
prominent, variable, but usually large and projecting conspicuously beyond
dorsal margin. Pallial sinus extending to midline, lower margin fused with
pallial line, not markedly ascending, shape variable, but usually straight dorsally,
roundly truncate anteriorly.
Dimensions
Holotype: 46,4 <x 22,9 mm.
Paratypes: 59,5 x 28,2 mm;
52,4 x 24,7 mm.
Distribution records
Churchhaven, 1,5 m a.s.l. (N.M., S.A.M.) (type locality).
216 ANNALS OF THE SOUTH AFRICAN MUSEUM
Remarks
Donax sanctuarium appears to be quite distinct from any other member of
the genus. It may perhaps be compared with D. (Capsella) variegata (Gmelin,
1791), from the Mediterranean, which, however, is rounded and smooth
posteriorly, and has a weaker nymph. D. haughtoni Carrington & Kensley, 1969,
from the Late Cenozoic of Namaqualand, is a larger, deeper and more ovate
species, with a more central umbo and a smooth posterior face. D. oweni
Hanley, 1843, from West Africa is smaller (25 mm) and has a very curved
ventral margin.
D. sanctuarium (whose specific name is a literal translation of the type
locality Churchhaven) does not agree with any of the described subgeneric units
(cf. Keen 1969) of the genus Donax. As the current system of classification is
most unsatisfactory, we have not erected a new taxon for its reception.
Family Tellinidae
Gastrana fibrosa sp. nov.
Fig. 19
Type material
Holotype: right and left valves (articulated shell), SAM—K4574.
Paratypes: 1 articulated shell, 1 right valve, 2 left valves, NM—A1220/
T1830; ;
2 right valves, 2 left valves, SAM-K4581.
Diagnosis
Large (60-70 mm), very solid, posterior end produced but not rostrate;
sculptured medially by radial threads, more or less replaced at each end by
lamellose concentric threads. Hinge plate massive, left anterior cardinal
projecting and spathulate, not bifid.
Description
Shell very solid, moderately compressed, with a narrow gape at each end.
Umbo situated two-fifths of total length from anterior end; anterior end broadly
rounded, posterior end somewhat tapering, with a very slight umbonal ridge;
between the umbonal ridge and the postero-dorsal margin there is a faint
indication of a ridge, visible more as an alteration in the direction of the con-
centric sculpture than as a prominence; postero-dorsal margin straight or
slightly convex, ventral margin evenly convex or with a suggestion of a sinuosity
posteriorly.
Sculpture tripartite, with regular thread-like concentric lamellae posteriorly,
another weaker series anteriorly, and the median half or two-thirds bearing fine,
uneven, radial threads, crossed by growth lines which may render them almost
cancellate; in odd individuals the concentric threads may cross the whole
surface, although becoming irregular medially.
Hinge complex massive, two cardinal teeth in each valve, no laterals. Right
PLEISTOCENE MOLLUSCS FROM THE CAPE PROVINCE, SOUTH AFRICA 217
25mm
Fig. 19. Gastrana fibrosa sp. nov. Holotype. (Naval Academy, Saldanha.)
anterior cardinal almost perpendicular to hinge axis and diverging from
posterior cardinal at an angle of 40-45°; both teeth are simple and ridge-like,
not, or very slightly, bisected, posterior cardinal longer than anterior one. Left
anterior cardinal large and spathulate, projecting conspicuously from hinge
plate, tip rounded and simple, with only a trace of a median groove; posterior
cardinal a rather low, triangular ridge. Nymphs well developed. Adductor
muscle scars and pallial line well impressed, pallial sinus deep, largely confluent
with pallial line, end rounded to subacute.
Dimensions
Holotype: 66,5 x 43,5 mm;
total breadth 25,3 mm.
Paratypes: 72,4 x 46,6 mm;
68,8 <x 46,0 mm.
Distribution records
Naval Academy, Saldanha (1,7 m a.s.l.) (type locality) and Churchhaven
(1,5 m a.s.l.).
Remarks
Gastrana fibrosa is very closely allied to the poorly known G. multangula
(Gmelin, 1791) (syn. Tellina polygona Réding, 1798, non Gmelin, 1791) from
218 ANNALS OF THE SOUTH AFRICAN MUSEUM
Gambia, and may prove to be only a subspecies of that. Judging by R6mer’s
description and figures (1872: 272, pl. 51 figs 5-7), G. multangula differs in
possessing a subcentral umbo, very fine interstitial radial lirae, continuous
concentric threads which traverse the middle, and a stronger umbonal ridge.
Unfortunately details of the shape of the left anterior cardinal are unknown,
Romer referring to this merely as ‘subfisso’, which barely applies to G. fibrosa.
It should be noted, too, that while ROmer’s figure of G. multangula shows a
wholly free pallial sinus, his text contradicts this.
G. fibrosa closely approaches G. rostrata Carrington & Kensley, 1969, from
the Lower Pleistocene of Namaqualand in size and sculpture, but differs in its
non-rostrate posterior end and straight or convex dorsal margin, this being
concave in G. rostrata; hinge details also differ, notably the characteristic left
anterior cardinal, which in G. rostrata is short and distinctly bisected, rendering
the tip almost bifid.
G. fibrosa differs widely from the remaining two comparable species of
Gastrana, G. matadoa (Gmelin, 1791) (syn. Tellina abildgaardiana Spengler,
1798) from South and West Africa, and G. fragilis (Linnaeus, 1758) from the
Mediterranean, both of which are of Recent occurrence. It differs in its much
larger size, greater solidity, in outline, non-bifid left anterior cardinal, and
different sculpture, the radial element being well developed and the concentric
one irregular or restricted to the ends; in G. matadoa and G. fragilis the concen-
tric lamellae are well developed over the whole surface, and radial sculpture is —
reduced to very fine interstitial striae.
Tellina (Eurytellina) madagascariensis Gmelin, 1791
Fig. 20
Tellina madagascariensis Gmelin, 1791: 3237. Romer, 1871: 64, pl. 17 (figs 4-7). Dautzenberg.
1912: 102. Nicklés, 1950: 224, fig. 435. Paes da Franca, 1960: 36.
Tellina rosea (non Spengler, 1798): Haughton, 1932: 37.
Tellina (Eurytellina) madagascariensis: Boss, 1969: 122, pl. 10 (fig. 3).
Description
Elongate-elliptical, compressed, umbo peaked, slightly anterior to middle,
posterior end strongly flexed to right, narrowly gaping; right valve less convex
than left; anterior margin broadly rounded, postero-dorsal margin declivous,
shallowly concave behind umbo, then straight or slightly convex, posterior end
narrowly rounded or feebly truncate, ventral margin evenly curved. External
surface with fine growth striae and very faint, scratch-like radial striae; right
valve with a low, blunt umbonal ridge, corresponding to a shallow depression
in the left valve. Right anterior cardinal tooth simple, posterior one bifid as is
the left anterior cardinal; right posterior cardinal thin, ridge-like and simple.
Anterior lateral teeth close to cardinals, posterior ones distant, situated at end
of nymph; in right valve anterior lateral is strong, posterior one feeble; both left
laterals are weak. Nymphs well developed, each with a ridge on its inner surface.
PLEISTOCENE MOLLUSCS FROM THE CAPE PROVINCE, SOUTH AFRICA 219
Fig. 20. Tellina (Eurytellina) madagascariensis Gmelin, 1791. (Churchhaven.)
220 ANNALS OF THE SOUTH AFRICAN MUSEUM
Adductor muscle scars well impressed; two faint cruciform scars just below
posterior end of pallial line. Pallial sinus not ascending, lower margin wholly
confluent with pallial line, extending almost to anterior adductor.
Dimensions
69,5 x 41,3 x 17,3 mm, 66,8 x 44 x 19 mm.
Distribution records
Redhouse (S.A.M.); Deal Party, 12-23,8 m (N.M.); Cradock, 45,7 m
(N.M.); Klein Brak River (Boss 1969); Churchhaven, 1,5 m a.s.l. (S.A.M.);
Geelbek (S.A.M.); Verlorevlei, 4 m a.s.l. (S.A.M.); Cape Cross, S.W.A.
(S.A.M.).
Boss (1969) showed that previous Recent records of Tellina madagascariensis
from South Africa were based on Tellina alfredensis Bartsch, 1915. He did,
however, record the true West African 7. madagascariensis from a raised beach
at the ‘Klein’ (i.e. Klein Brak) River mouth. During the Pleistocene this species
was in fact abundant along much of the south and west coast of the Cape, and
in South West Africa. After its extinction along the South African coast, no
doubt due to climatic change, 7. madagascariensis was replaced by the endemic
T. alfredensis.
Tellina madagascariensis occurs living today from Baia dos Tigres (17°S) to
Sao Thome (0°) (Boss 1969).
Macoma (Heteromacoma) tricostata (R6mer, 1872)
Fig. 21
Tellina tricostata Romer, 1872: 235, pl. 49 (figs 10-12).
Description
Both valves rather inflated, right valve slightly deeper than left; outline
ovate-trigonal, umbones high, usually situated two-fifths of length from anterior
end, but sometimes nearly median; posterior end moderately rostrate, tapering
rapidly, end roundly truncate; postero-dorsal margin straight and evenly
declivous, rest of margin evenly rounded. Shell slightly flexed to right posteriorly,
right valve with a very weak umbonal ridge, followed by 1-2 very feeble ridges,
left one with a depressed umbonal ray, followed by a weak ridge near the dorsal
margin; surface otherwise sculptured by growth lines only. Pallial sinus very
deep, almost reaching anterior adductor scar, non-ascending, lower margin
completely fused with pallial line, upper margin feebly bilobate; sinus larger in
right valve than in left. Hinge of each valve with two cardinal teeth, of which the
posterior ones form thin, feeble ridges, the anterior ones being erect and more
peg-like, simple in the right, apically bifurcate in the left. Nymphs large, thick,
bordered by deep ligamental furrows.
Dimensions
33,6 < 43,1 x 23,0 mm; 46 x 36,3 x 18,7 mm; 47,2 x 383493:
41,3 x 35,0 mm.
PLEISTOCENE MOLLUSCS FROM THE CAPE PROVINCE, SOUTH AFRICA 2a
See ee ee at ne ee
a ed
20mm
Fig. 21. Macoma (Heteromacoma) tricostata (ROmer, 1872). Note variation in form.
(Verlorevlei.)
Distribution records
Verlorevlei, 4 m a.s.l. (S.A.M.).
Remarks
Specimens from Verlorevlei agree well with Recent valves from Angola,
except in being thicker. While these specimens resemble Nickleés’s figure (1950:
fig. 432) of the exterior of Tellina nymphalis Lamarck, 1818, they disagree with
R6mer’s excellent figures (1872: pl. 45, figs 1-4) of the same species. These show
a non-rostrate shell with more extensive, rounded pallial sinus. On the other hand
these specimens agree closely with R6mer’s T. tricostata from Gabon. There is,
however, some variation in shape in the present material, and one specimen
somewhat approaches R6mer’s concept of 7. nymphalis in its moderately
reduced posterior end. Not only does the relationship between the two require
229 ANNALS OF THE SOUTH AFRICAN MUSEUM
investigation, but re-examination of the holotype of 7. nymphalis is desirable in
order to confirm R6mer’s interpretation. For the present it is advisable to use
the name tricostata, which undoubtedly applies to the present material.
The only comparable South African Macoma is M. (Macoma) litoralis
(Krauss, 1848), a smaller, more compressed species with a lower umbo, stronger
hinge-teeth and inconspicuous nymphs.
Leporimetis (Leporimetis) hanleyi (Dunker, 1853)
Fig. 22
Tellina hanleyi Dunker, 1853: 53, pl. 10 (figs 4-6). Romer, 1871: pl. 14 (figs 7-9); 1872: 214.
Apolymetis orbicularis (partim non Sowerby); Barnard, 1964: 549.
Apolymetis papyracea (non Gmelin, 1791); Kilburn in Davies, 1972: 252.
Description
Ovate, with peaked umbones situated slightly posterior to middle; shell
flexed to the right and narrowly gaping posteriorly; antero-dorsal margin gently
convex, anterior end strongly rounded, postero-dorsal margin straight or slightly
15mm
Fig. 22. Leporimetis (Leporimetis) hanleyi (Dunker, 1853). (A. Knysna. B. Redhouse.)
PLEISTOCENE MOLLUSCS FROM THE CAPE PROVINCE, SOUTH AFRICA 223
convex, fairly steeply descending, posterior margin truncate to broadly rounded,
often showing a weak double sinuosity; ventral margin evenly curved, not
sinuated. Sculptured by fine growth lines, becoming coarse posteriorly; umbonal
ridge very feeble, often not visible in right valve, in left valve bordered behind by
a shallow trough, followed by a second feeble ridge. Hinge in each valve with two
ridge-like cardinals, of which the anterior ones are the stronger. Nymphs well
developed, moderately impressed into hinge-plate. Pallial sinus deep, extending
almost to anterior adductor scar, lower margin largely free from pallial line,
upper margin shallowly concave, end rounded to moderately truncate.
Dimensions
41,8 x 31,8 mm, 44,7 x 35,0 mm, 39,0 x 30,5 x 14,4 mm (complete
example).
Distribution records
Cenozoic: Redhouse, 7,2 m a.s.l. (N.M.); Knysna, 4,6 m a.s.l. (S.A.M.);
Sedgefield, 5 m a.s.l. (S.A.M.); Klein Brak River, 3,3 m a.s.l. (S.A.M.).
Remarks
Pleistocene specimens agree well with RoOmer’s detailed description and
figures of Tellina hanleyi from Luanda (the original description is not available
to us). While South African examples are rarely as rounded posteriorly as in
these figures there is much variation in this respect, and one example from
Redhouse is decidedly curved behind. The species does not appear to have been
discussed by recent authors.
The only comparable species is the Recent West African Leporimetis
(Florimetis) papyracea (Gmelin, 1791), which differs in its more elevated,
subcentral umbones, markedly more inflated valves, and distinct umbonal
ridge, median flexure and postero-ventral sinuosity. Although French workers
commonly utilize the name ‘/acunosa Schroter, 1788’ for L. papyracea, that
nomen was proposed in a non-binomial work, and was not validated until 1817
(by Dillwyn).
The fossil material recorded by Barnard (1964) as Apolymetis orbicularis is
actually Leporimetis hanleyi. The Recent Leporimetis (Florimetis) orbicularis
(Sowerby, 1889) from the False Bay—Port Alfred area is a strongly inflated,
suborbicular species with a well-developed umbonal ridge, and there are also
differences in details of the cardinal complex.
ACKNOWLEDGEMENTS
We wish to express our sincere thanks to Mr V. Branco for his help with the
drawings and Mr D. A. Gerneke for the photography. The two scanning
electron photomicrographs were printed by Mr R. Cross at Rhodes University.
One of us (A.J.T.) was supported by a research grant from the South African
Council for Scientific and Industrial Research.
224 ANNALS OF THE SOUTH AFRICAN MUSEUM
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Woop, W. 1828. Index testaceologicus. Supplement: 1-59. London: Wood.
6. SYSTEMATIC papers must conform with the International code of zoological nomenclature
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BuLLouGu, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
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FIscHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines.—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.
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 67 Band
October 1975 Oktober
Part ff Deel
THE MORPHOLOGY AND RELATIONSHIPS OF
A CROCODILIAN, ORTHOSUCHUS STORMBERGI,
nieomM THE UPPER TRIASSIC OF LESOTHO
By
DIANE S. NASH
Cape Town Kaapstad
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THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN,
ORTHOSUCHUS STORMBERGI, FROM THE UPPER TRIASSIC OF
LESOTHO
By
DIANE S. NASH
Birkbeck College, University of London*
(With 44 figures and 4 tables)
[MS. accepted 13 January 1975]
ABSTRACT
The skull, lower jaws and most of the postcranium of the type of Orthosuchus stormbergi
Nash, 1968 were collected in 1963, by an expedition of the South African Museum, from the
Upper Triassic Red Beds Formation of Lesotho. The skull and jaws of a smaller specimen,
collected from the same horizon but a separate locality, are considered to be congeneric.
The cranial table is flattened, and of typical crocodilian form and sculpturing. There is a
deep otic recess, and pronounced otic notch which lies open posteriorly. The quadrate and
quadratojugal are strongly inclined, and the quadrate sutures with the parietal and squamosal
within the superior temporal fossa. A short bony secondary palate is developed, and the
pterygoids bear prominent flanges characteristic of crocodilians.
The anterior dorsal ribs are flanged on both the leading and rear margins. The coracoid,
radiale and ulnare are elongated. The acetabulum is open, and the pubis excluded from the
acetabular margin. The ankle joint is crocodilian in type, and a paired row of dorsal scutes,
which are imbricated and sculptured, is developed over the trunk and tail.
Orthosuchus is closely related to Notochampsa, Erythrochampsa and Protosuchus, and
probably also to Stegomosuchus and Pedeticosaurus. It is also related to Hemiprotosuchus
though possibly less closely.
The thecodontian heritage of Orthosuchus can be seen in the presence of an antorbital
fenestra, small basipterygoid process, and in that the quadrate has a posterior contact with the
squamosal. The lower jaw includes a prearticular, and a retroarticular process is not developed.
The musculature of Orthosuchus probably differed little from that of living crocodiles.
As in the modern group, the hind-limb and tail musculature were probably powerful, while
there was a general reduction in the lower arm and hand musculature associated with great
mobility of the wrist joint, and use of the elongated carpus as an extra limb segment.
The presence of a soft secondary palate, which in life extended well back to the base of the
skull, suggests that Orthosuchus spent much of its time in water. The laterally orientated
nostrils and orbits, and the relatively longer proximal limb elements, indicate that Orthosuchus
was less well adapted to this environment than are living crocodiles.
CONTENTS
PAGE
Introduction ; : : ; ; : : : c : : : a 22S
Historical review ‘ : : : » 228
Taxonomic position, diagnosis and materials under consideration : : ; min geet 4
Description of Orthosuchus . : ‘ ‘ é : ; : ? : rn 25D
heskull>.. : : i ; ; ‘ : : 2 : : oa 235
The lower jaw. ; ‘ : : : : : ‘ | 2A6
The vertebral column and ribs : : : : : d : . . 248
The limb girdles and limbs. d i ‘ é ; : é : «© 254.
Body armour : : : : $ : : : ; : : a eaeOS
Myology of Orthosuchus . : : : B : : ‘ 4 ‘ . 264
* Present address: Bath College of Higher Education, Bath
227
Ann. S. Afr. Mus. 67 (7), 1975: 227-329, 44 figs, 4 tables
228 ANNALS OF THE SOUTH AFRICAN MUSEUM
Age variations in the skulls of recent Crocodilia, and an analysis of the differences
between the skulls (K409 and K4639) of Orthosuchus : ; P 5 «ye eee
Mode of life : : : : : < DZS
Relationship of Orthosuchus to thecodontians : ; : ‘ d : 95
Relationship of Orthosuchus to crocodilians . : : : 25, 298
Limb and limb girdle proportions in Crocodilia and Thecodontia : oe. 315
Relationship of the protosuchians to forms of questionable crocodilian affinity . ee |
Acknowledgements . : : . ; : : : : : : PaO
References : : : : ‘ : : : ‘ : : : 2”. 326
Abbreviations. 5 : ‘ : : : . ; : ; : eal 329
INTRODUCTION
In April 1963 an expedition of the South African Museum, led by Professor
A. W. Crompton (now Director of the Museum of Comparative Zoology at
Harvard University) collected reptilian material from the Upper Triassic Red
Beds Formation of the Stormberg Series of southern Africa. It soon became clear
that among this material was a nearly entire skeleton of a new type of crocodilian.
Pending further preparation and more detailed study, a preliminary description
was published (Nash 1968) in which the type was named Orthosuchus
stormbergi.
A second smaller skull had also been collected from the same ree oe but
from a separate locality. An analysis of this second specimen demonstrates that
this material is congeneric.
This paper is mainly concerned with the detailed description of the skull and
postcranial skeleton of Orthosuchus. The probable arrangement of the muscula-
ture of head and limbs is given, and some observations are made as to its mode
of life. The position of Orthosuchus in relationship to both thecodontians and
crocodilians is discussed.
Fig. 1. Orthosuchus stormbergi type skeleton (K409). Dorsal view of the skull and postcranium. Ns
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 229
oe i ee
Fig. 2. Orthosuchus stormbergi type skeleton (K409). Ventral view of the skull and
postcranium.
HISTORICAL REVIEW
Although a number of forms of Triassic age have recently been associated
with the Crocodilia, there are still only a limited number which bear a direct
relationship to this group. Notochampsa was the first of these to be described
(Broom 1904). On the basis of two specimens from the same locality (Barkly
East, Cape Province, South Africa) but different horizons, Broom proposed two
species within this genus which he included within the crocodilian suborder
Mesosuchia. Notochampsa istedana was collected from the Cave Sandstone of
the Stormberg Series, and the second, Notochampsa longipes, from near the top
of the underlying Red Beds. It is now generally considered that the Cave
Sandstone does not represent a separate formation, but is merely a facies of the
Red Beds.
Having further developed the material of Notochampsa istedana, Haughton
(1924) redescribed the type, and questioned its crocodilian relationship.
Haughton came to the conclusion that there were close resemblances between
Notochampsa istedana and Pedeticosaurus leviseuri Van Hoepen, 1915 and
provisionally placed the two genera in the same family, the Notochampsidae.
He considered them advanced pseudosuchians having affinities with the
Crocodilia.
At this time, Haughton also reconsidered the specimen described by
Broom as Notochampsa longipes. In his opinion Broom had attached too much
significance to the resemblances between the dorsal scutes of the two forms
Notochampsa istedana and longipes. Haughton considered the two types quite
distinct, and erected a new genus, Erythrochampsa, for Notochampsa longipes,
placing this type in the Crocodilia.
Von Huene (1925) disagreed with Haughton’s and Broom’s interpretations
230 ANNALS OF THE SOUTH AFRICAN MUSEUM
and placed both Notochampsa and Erythrochampsa in the Pseudosuchia. In his
view, the Pseudosuchia contained forms all of which indicate a link between this
group and the crocodilians. He postulated an evolutionary sequence with the
following pseudosuchians leading to the Crocodilia: Erpetosuchus, Aétosaurus,
Stegomosuchus, Sphenosuchus, and Pedeticosaurus. Presumably, Notochampsa
and Erythrochampsa form the final link in this series.
Broom (1927) reassessed the material of Sphenosuchus and returned to the
question of the taxonomic position of Notochampsa and Erythrochampsa. He
was now quite satisfied that their generic separation was valid, but included both
within the Crocodilia. Furthermore, he questioned the evolutionary sequence
proposed by Von Huene. He considered Erpetosuchus and Aétosaurus too
specialized; he believed Pedeticosaurus to be bipedal and not related to croco-
dilian ancestry, and Sphenosuchus to be closely allied to Pedeticosaurus. He was
of the opinion that Stegomosuchus might well occupy a position close to
crocodilian ancestry, though the type was too imperfectly known to be sure.
Broom was of the opinion that a group of pseudosuchians of a form similar to
Euparkeria left dry ground and inhabited the marshes, developing the com-
paratively feeble long limbs and flattened skulls characteristic of the crocodiles.
There is now an excellent redescription of Euparkeria from the Lower
Triassic beds of Aliwal North, South Africa, by Ewer (1965). Ewer considered
Euparkeria as the probable direct ancestor to advanced forms like Ornithosuchus —
and Hesperosuchus.
In 1930 and 1931 the first North American specimens of an ancestral
crocodile of Upper Triassic or basal Jurassic age were collected from the
Dinosaur Canyon Sandstone of Cameron, Arizona. Barnum Brown published a
preliminary notice in 1933 in which he named the type Archaeosuchus richardsoni
and established a new family, the Archaeosuchidae. Unfortunately, the name
Archaeosuchus was already preoccupied (Archaeosuchus cairncrossi Broom, 1905,
a titanosuchian), and in 1934 Brown renamed this important reptile Protosuchus
richardsoni, changing the family name to Protosuchidae.
In a revision of the classification and evolution of the Crocodilia, Mook
(1934) proposed a new suborder, the Protosuchia, to contain the single genus
Protosuchus richardsoni. No mention was made of either Notochampsa or
Erythrochampsa. Mook considered that Protosuchus should be placed very near
the line of direct ancestry of the mesosuchian crocodiles.
Subsequently Colbert & Mook (1951) redescribed Protosuchus, this time
relating it to both Notochampsa and Erythrochampsa. They remained firmly
convinced of the crocodilian nature of Protosuchus, but considered that the
relationships of Notochampsa were less certain. Consequently they suggested
that the family Protosuchidae be retained in preference to Notochampsidae.
In their opinion, Pedeticosaurus was less definitely crocodilian than either
Erythrochampsa or Notochampsa and should be placed among the pseudo-
suchians.
Other forms for which crocodilian affinity has at some time been suggested
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 931
include Platyognathus hsui Young, 1944, from the Dark Red Beds of the Lower
Lufeng Series, Yunnan Province, China. Additional material was described by
Simmons in 1965. Following Young, Simmons placed Platyognathus in a new
family of Pseudosuchia, the Platyognathidae. He believed Platyognathus to be a
specialized form, not on the direct line of descent toward Notochampsa and
Erythrochampsa. However Romer (19726) was of the opinion that this species
did not merit the erection of a separate family, and included Platyognathus
within the Protosuchia.
Microchampsa scutata, also from the Lower Lufeng Series, is generally
considered a protosuchian though of an essentially new type (Young 1951;
Simmons 1965).
A preliminary notice of another crocodilian, probably of Upper Triassic
age, from a fissure in the Carboniferous Limestone of Glamorgan, Wales, was
given by Kermack (1956). A complete description of this type is still awaited.
More recently (Bonaparte 1969, 1971) Hemiprotosuchus leali has been
described as a crocodilian closely related to Protosuchus. Hemiprotosuchus is
from the upper beds of the Los Colorados Formation, La Rioja Province,
Argentina.
In 1959 Reig described the skull of Proterochampsa barrionuevoi from the
late Middle or basal Upper Triassic Ischigualasto beds of San Juan Province,
Argentina. Sill reassessed this material in 1967, and agreed with Reig that it
represented an early crocodilian. Sill suggested that whereas both Notochampsa
and Proterochampsa strongly resemble one another, neither genus is very close
to Protosuchus. It therefore appeared to Sill that there were two lines of
crocodilian evolution during the Triassic. Proterochampsa could be regarded as
representing an earlier and more typical line of crocodilians, and Protosuchus as
representing an aberrant and more terrestrial group. In Sill’s view, Chasmato-
saurus probably most nearly represents the group of thecodontians from which
the Crocodilia arose.
Sill proposed a new suborder, the Archaeosuchia, an unfortunate choice in
view of Brown’s earlier experience. This was to contain two families, the new
Proterochampsidae, for the single genus Proterochampsa, and, provisionally, the
Notochampsidae, to contain Notochampsa and Erythrochampsa. He suggested a
separate suborder Protosuchia, to contain the Protosuchidae, with the sole
genus Protosuchus, and questionably the Sphenosuchidae, with Sphenosuchus,
Pedeticosaurus and Platyognathus.
Sill’s interpretation of Proterochampsa was strongly criticized by Walker
(1968). In his opinion the advanced crocodilian characters described by Sill are
actually phytosaurian in nature. Walker’s reinterpretation of the skull of
Protosuchus indicates its close relationship to Notochampsa and reaffirms the
position of Protosuchus as an ancestral crocodile.
In Walker’s view Stegomosuchus longipes is sufficiently well known to
establish its close relationship to Protosuchus. He has proposed grouping these
forms together with Erythrochampsa and Orthosuchus in one family, the
232 ANNALS OF THE SOUTH AFRICAN MUSEUM
Stegomosuchidae. Walker suggested Cerritosaurus Price, 1946, from the
Santa Maria Formation of Brazil, as a probable ancestor, pointing out that
since Cerritosaurus shows resemblances to Chasmatosaurus, both phytosaurs and
crocodiles appear to converge back on Chasmatosaurus.
More recently Romer (1971, 1972a) has demonstrated a close relationship
between Cerritosaurus and Proterochampsa, and two newly discovered thecodon-
tians, Chanaresuchus bonapartei and Gualosuchus reigi. Romer regards these four
types as forming a close family, the Proterochampsidae, which represents a
sterile offshoot of the primitive proterosuchian stock.
Walker (1970) has suggested a major regrouping of crocodiles and closely
allied forms within a proposed order Crocodylomorpha. The Stegomosuchidae
is placed as the sole family in the infraorder Protosuchia of the suborder
Crocodylia, which also includes the taxa Mesosuchia and Eusuchia, now ranking
as infraorders. A suborder Paracrocodylia is proposed to include the infraorders
Pedeticosauria, Baurusuchia and Hallopoda. The Pedeticosauria includes only
the Upper Triassic family Pedeticosauridae with the genera Pedeticosaurus,
Sphenosuchus, Saltoposuchus, Platyognathus and Hesperosuchus.
In view of this, Romer (19725) has advocated retaining the Protosuchia as a
suborder, but to include both ancestral forms and those others that are trending
toward the crocodiles from the typical thecodontian pattern. Protosuchus,.
Notochampsa, Erythrochampsa, Orthosuchus, Stegomosuchus and questionably
Platyognathus and Microchampsa are placed in the family Protosuchidae, while -
Sphenosuchus, Hemiprotosuchus and questionably Pedeticosaurus are grouped in
the Sphenosuchidae.
On the other hand, Bonaparte (1971) considers that the Crocodilia as a
group are too distinct for the inclusion of Sphenosuchus, and has suggested that
the allies of Sphenosuchus are found in Pseudhesperosuchus Bonaparte, 1969
and Hesperosuchus. These three forms he groups in the Sphenosuchidae within
the suborder Pseudosuchia.
TAXONOMIC POSITION, DIAGNOSIS AND MATERIALS
UNDER CONSIDERATION
TAXONOMIC POSITION
Orthosuchus has retained from thecodontian ancestors a number of
primitive features, notably the antorbital fenestra and prearticular. However,
many other characters present in Orthosuchus are diagnostic of the Crocodilia.
These are as follows:
(1) Sculpturing of the external surfaces of the bones of the skull and lower jaw.
(2) Forwardly sloping quadrate and development of an extensive otic notch.
(3) Midline fusion of the parietals.
(4) Some degree of secondary palate formation.
(5) Firm fusion of the pterygoid and quadrate with the braincase; the
pterygoid bears a flange with a guide facet on its external margin for the
lower jaw.
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 233
(6) Elongated coracoid, radiale and ulnare.
(7) The acetabulum is open; the pubis is excluded from the acetabulum by a
forward process of the ischium.
(8) Femur without marked fourth trochanter; no development of greater
trochanter.
(9) Ankle joint of crurotarsal pattern with well developed tuber on the
calcaneum.
(10) External surfaces of the dorsal scutes are sculptured.
Orthosuchus is closely related to Protosuchus, Notochampsa and Erythro-
champsa, and indeed probably also to Pedeticosaurus, Stegomosuchus and
Hemiprotosuchus. All these forms may reasonably be included within the same
family. Pedeticosaurus and Stegomosuchus are both insufficiently known to
establish the family on either type. Furthermore, although the relationships of
Notochampsa are no longer in any doubt, it seems more reasonable to retain the
family Protosuchidae because of its previous general acceptance.
Protosuchia Mook 1934
Protosuchidae Brown 1934
Pedeticosaurus Van Hoepen iS
Notochampsa Broom 1904
Stegomosuchus Von Huene 1922
Erythrochampsa Haughton 1924
Protosuchus Brown 1934
Orthosuchus Nash 1968
Hemiprotosuchus Bonaparte 1969
Type
Orthosuchus stormbergi Nash, 1968, SAM-—K409. An articulated skeleton,
nearly complete.
Horizon |
Upper Red Beds Formation of the Stormberg Series, Upper Triassic.
Locality
Orange River Valley in the Qacha’s Nek Province, Lesotho (formerly
Basutoland).
GENERIC AND SPECIFIC DIAGNOSIS
Skull bones ornamented; extensive slender preorbital region; skull table
flattened; external nares separate and terminal; temporal fenestrae approxi-
mately equal in size; upper temporal fenestra large and posteriorly situated on
the skull table; intertemporal area narrow, antorbital fenestra opening into a
sinuous groove posteriorly; tooth count 7—8/15—-18; pronounced lateral notch
lies on the premaxilla/maxilla suture; two supraorbital bones on each side;
frontal enters the superior temporal fenestra; midline fusion of parietals; a deep
otic notch formed by the forwardly sloping quadrate and quadratojugal and
234 ANNALS OF THE SOUTH AFRICAN MUSEUM
overhung by the postorbital and squamosal which together form a wide upper
temporal arch; quadrate fenestrated and with a posterior articulation with the
squamosal; secondary palate formed from the premaxillae and mazxillae,
primary palate vaulted; pterygoid and quadrate fused to the braincase, the
pterygoid bearing a prominent flange with external facet to guide the lower jaw
on closure; lateral and median eustachian openings lying anteriorly in the
basisphenoid; small basipterygoid process present. External mandibular
fenestra large; prearticular well developed; the surangular forming a horizontal
flange; no pronounced retroarticular process. Vertebrae amphicoelous; dorsal
ribs with both an anterior and posterior flange. The coracoid elongated, the
proximal expansion greater than the distal. Humerus not greatly expanded
proximally, deltopectoral crest prominent; radiale and ulnare elongated,
radiale longer than the metacarpals; a pisiform and two distal carpal elements
present; manus small. Iliac blade forming a pointed preacetabular process; the
ischium with a forward process that excludes the pubis from the acetabulum;
the astragalus and calcaneum large, the calcaneum bearing a prominent tuber;
four elongated metatarsals, the fifth reduced in length. A complete dorsal
cuirass of paired ornamented scutes, gastralia present.
MATERIALS UNDER CONSIDERATION
SAM-K409, the type, listed above.
SAM-K4639, skull and lower jaws in articulation. This specimen was
excavated from an elevation of 2115 metres on the slopes of Majubanek
Mountain facing the Kromme Spruit River. It is from the same stratigraphic
level as the type specimen.
Preservation and preparation
The general condition of the type specimen, K 409, is excellent though there
is some distortion of parts attributable to post-mortem damage. Further, the
skull fractured during excavation, and the trunk was broken in two places. The
right femur was also broken. Major breaks were restored in the field. The
material was X-rayed before preparation was undertaken.
All preparation has been carried out by mechanical means, and this proved
a lengthy process. The vibro-tool was used extensively, but much of the work
was done with the use of a pin-vice. A few areas of bone had been eroded before
preservation, and these parts were restored with N.H.P. self-hardening Model
Plastic.
The preserved parts of specimen K409 include the skull and both rami of
the lower jaw, twenty-one presacral (three or possibly four mid-dorsals are
lacking), two sacral and eight caudal vertebrae, two cervical ribs, eleven dorsal
ribs and rib fragments, both scapulae, the right coracoid and part of the left
coracoid; the left fore-limb, carpus and manus, the proximal carpal elements
and distal portions of the right radius and ulna; the entire pelvis; both hind-
limbs, the tarsus and incomplete left pes, and the right astragalus and calcaneum ;
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 235
a double row of dorsal scutes covering the trunk and anterior caudal region;
several gastralia.
The second specimen, K 4639, is a smaller skull and articulated lower jaws
partially embedded in matrix. The bone is very delicate. The block is split into
two halves to expose the lateral surface of the skull. It was possible to prepare
the posterior region of the skull, but anteriorly it had been crushed during
preservation and work was not found to be profitable in this area.
DESCRIPTION OF ORTHOSUCHUS
THE SKULL
The skull of the type specimen is entire and fully prepared. It is in an
excellent state of preservation, although there has been slight post mortem
crushing and some consequent distortion of the cranial region. The smaller
specimen, K4639, lacks the extreme tip of the snout and occiput.
External features
The skull is flattened along its length. The cranial table is broad and
narrows to a slender snout with a slightly bulbous tip. The entire external skull
surface is sculptured in the form of an irregular pattern of shallow pits.
The antorbital fenestra is oval in shape. It opens posteriorly into a sinuous
groove which runs obliquely down towards the lateral orbital margin. The orbit
is large, and is directed outward and forward. Laterally, paired supraorbitals
together form a complete functional dorsal roof to the orbit. These elements are
triangular in shape, the anterior supraorbital being the larger of the two.
Sclerotic bones are not present. The temporal fenestrae are of approximately
equal size, and they approximate the size of the orbit.
Dermal roofing elements (Fig. 3)
Premaxilla
The premaxilla meets the tip of the nasal anteriorly on the snout, and
together these elements form a curved suture which passes back from the
posterior border of the external naris. The posterior margin of the premaxilla
lies within a deep backwardly directed notch on either side of the snout.
The premaxilla curves round to the palatal surface of the skull where it
makes a small contribution to the formation of a secondary palate. At the tip of
the snout, the premaxillae meet in the midline. Behind this, each borders a
premaxillary foramen which lies obliquely orientated to the long axis of the
sxull. The medial edge of the foramen is made by the maxilla. In life the
premaxillary foramina would have been closed by membrane, as in living
crocodiles. The premaxilla also extends inwards as a rounded area of bone
behind the premaxillary foramen, where it is sutured with the maxilla.
Each premaxilla carries four (previously described as six) discrete alveoli.
Two entire teeth are known, most of the remainder of the alveoli hold broken
236 ANNALS OF THE SOUTH AFRICAN MUSEUM
teeth. The teeth are conical and recurved, and the crowns bear longitudinal
striations. Internal to the alveolar border the premaxilla is perforated by
vascular and nervous foramina.
Nasal
The nasal is a large element forming most of the dorsal surface of the
rostrum. There is a distinct suture between the nasals in the midline. The tip of
the snout has been superficially weathered away, but the bony bar which
separates the nares must have been formed by the nasal, much as in the modern
alligator, Alligator mississippiensis.
ww
AMD A AWS jaa MY
Ge 55 YW4YY
LO,
4
mai a
Fig. 3. Orthosuchus stormbergi. Reconstruction of the skull (natural size). A. Dorsal view.
B. Lateral view.
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 237
The nasal unites laterally with the premaxilla and, behind the lateral notch,
with the maxilla to the level of the antorbital fenestra. Posteriorly the nasal
forms a wedge of bone which meets the prefrontal laterally, and the frontal
medially.
Frontal
The area previously shown as the prefrontal (Nash 1968) is no longer
regarded as a discrete element, but as part of the frontal. The frontal is a paired
element, as it generally is in mesosuchians, e.g. Alligatorellus and Goniopholis.
This is unlike the condition in the eusuchians, where both the frontals and
parietals are fused in the midline.
In Orthosuchus an anterior process of the frontal interposes between the
nasals in the midline. Laterally each frontal forms a curved suture with the
prefrontal. The frontal broadens considerably and extends postero-laterally to
form an oblique suture with the postorbital. A medial process extends posteriorly
to meet the parietal within the margin of the superior temporal fenestra. A part
of the orbital margin is formed by the frontal, as is the anterior margin of the
superior temporal fenestra. The laterosphenoid makes broad sutural contact
with the posterior medial process of the frontal on its ventral surface.
Parietal
This forms the greater part of a narrow temporal bar separating the two
superior temporal fenestrae. In specimen K4639, a median suture is discernible,
a juvenile condition, but in K409 the left and right parietals have fused together
to form a single element. There is no trace of a pineal foramen.
Only the anterior third of the superior temporal fossa* is open from below
for the passage of the m. adductor mandibulae externus profundus (= tempo-
ralis adductor). Laterally the quadrate and squamosal form part of the floor of
the fossa, while medially the remainder is formed by the parietal. The parietal
also forms the major part of the posterior wall of the fenestra, and meets the
squamosal in an interdigitating suture.
The posterior edge of the parietal is concave and is bent down to form the
superior edge of the occiput where it meets the supraoccipital ventrally. A small
foramen, visible on the left side, probably in life provided passage for the
temporal artery which ran through a canal into the superior temporal fossa.
Part of the roof of this canal is formed by the parietal and part by the squamosal,
while the floor is made by the supraoccipital and exoccipital.
Maxilla
Anteriorly the maxilla forms a knife-like edge which projects forwards
and conceals part of the lateral notch. Behind this it forms the lateral wall of the
rostrum and the anterior and lateral margins of the antorbital fenestra.
* The terms ‘temporal fenestra’ and ‘temporal fossa’ are not synonymous. The former is
used for the opening in the side of the skull; the latter refers to the space within the skull medial
to (for instance) a superior temporal fenestra.
238 ANNALS OF THE SOUTH AFRICAN MUSEUM
Unfortunately, the type is damaged on both sides at the anterior margin of the
orbits and a suture between the maxilla and jugal cannot be seen with any
certainty. However, it seems likely that the maxilla was excluded from the border
of the orbit by the lachrymal and jugal. By inference, the maxilla probably
excludes the jugal ventrally from the lateral margin of the post-palatine vacuity.
On the palatal surface, and at the level of the mandibular notch, the
maxillae extend towards one another, meeting in the midline to form a short
secondary palate. An anterior process from each maxilla runs forward between
the premaxillae and the premaxillary foramina. Paired, elongate choanae open
at the level of the first maxillary tooth, and extend back to the palato-maxillary
suture. The alveolar wall of the maxilla borders the choana and carries three or
four alveoli.
Jugal
An anterior process of the jugal forms the lateral margin of the orbit. This
process also extends inwards along its length so that it forms a shelf of bone
bordering the orbital region on the palatal surface of the skull. Although
sutures cannot be discerned, the jugal probably forms a short suture with the
lachrymal and, more posteriorly, with the maxilla on the palatal surface.
Behind this, the jugal meets the ectopterygoid in a straight suture.
An ascending process of the jugal meets the postorbital to form a superficial
postorbital bar. Although the postorbital bar is preserved on both sides of the:
skull in K 409, it is traversed by numerous cracks, and no clear suturing between
the jugal and postorbital is evident. The internal surface of the bar can be seen
on the right side of K4639, but again there is no suture visible along its length.
On balance, it seems likely that the jugal and postorbital together form an over-
lapping suture, as is the case in modern crocodiles. In this way the jugal reaches
up to around the halfway point on the external surface of the bar, while the
internal surface is formed by the postorbital.
A narrow, posterior process of the jugal forms the lateral margin of the
inferior temporal fenestra. Posteriorly it forms an oblique suture with the
quadratojugal.
Lachrymal
The element previously described as the lachrymal (Nash 1968) is now
interpreted as consisting of both lachrymal and prefrontal. On the left side of
the type the lachrymal is damaged at its extremities, and very little is preserved
on the right.
The lachrymal is triangular in shape, and is sutured medially with the
prefrontal. Its lateral border forms the medial wall of the antorbital fenestra and
groove. The lachrymal meets the maxilla both in front of the antorbital fenestra
and behind, where it forms the anterior margin of the orbit. This margin is
thickened, and penetrated by a foramen which in life provided passage for the
lachrymal duct.
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 239
Prefrontal
The prefrontal is bounded by the lachrymal laterally, and by the nasal and
frontal medially. It forms the antero-medial margin of the orbit. Two processes
of the prefrontal extend down from the internal orbital margin. The outer
process lies along the internal edge of the posterior margin of the lachrymal;
it is relatively short and tapers off halfway along the preorbital bar. Its extreme
tip is broken so that it is impossible to be sure how far it extended, but it is
doubtful that it reached the maxilla. In modern crocodiles a comparable
process is found, small and never reaching the maxilla. The inner descending
process passes medially and somewhat posteriorly, but again in Orthosuchus it is
incomplete. This process corresponds to that which articulates with the pterygoid
and palatine bones in living crocodiles. By inference it seems likely that in
Orthosuchus this process met its fellow in the midline and may also have met the
pterygoid ventrally.
Postorbital
This bone lies between the orbit and the temporal fenestrae and forms the
anterior corner of the cranial table. Medially the postorbital unites with the
postero-lateral process of the frontal in an oblique suture, while laterally it
extends to form the postorbital bar with the jugal. Posteriorly the postorbital
extends back a short distance between the superior and inferior temporal
fenestrae to make oblique sutural contact with the squamosal. This suture lies in
much the same position as it does in living crocodiles. The postorbital forms
little of the lateral margin of the superior temporal fenestra, and does not
contribute to its floor. Neither does it contribute to the dorsal margin of the
inferior temporal fenestra.
Squamosal
This is a very large bone which forms the major area of the lateral and
posterior margins of the cranial table. In K409 the borders of the superior
temporal fenestra are thickened and heavily sculptured. However, in K4639
there is no such obvious thickening, a feature that again might be due to an
age difference.
In the type specimen a longitudinally running groove delimits a thickened
inner border of the upper temporal arch from a thinner flange which slopes down
and overhangs a large, forwardly directed otic recess. In living crocodiles a
groove in this position marks the junction between scales covering the skull and
those covering the upper ear flap. It therefore seems probable that in life the ear
of Orthosuchus was similarly protected by ear flaps.
In living crocodiles the ear flaps are open anteriorly when the top of the
animal’s head is out of water, and closed when it submerges. This action does
not completely exclude water from the otic recess, but presumably it protects
the tympanic membrane from mechanical injury when the animal is submerged
(Shute & Bellairs 1955).
240 ANNALS OF THE SOUTH AFRICAN MUSEUM
In Orthosuchus the otic recess is closed anteriorly by the squamosal, which
extends down beneath the squamosal flange to unite with the quadratojugal
below. In this way, a narrow process of the squamosal interposes between the
postorbital and quadratojugal. Posteriorly the flange extends back as a tapering
process, the extremity of which is carried back significantly beyond the level of
the single basioccipital condyle. Posteriorly the squamosal contributes to the
occiput, meeting the exoccipital ventrally and the parietal medially. Its occipital
surface is not sculptured (Fig. 6).
Within the floor of the superior temporal fossa, the squamosal is sutured
obliquely to the palatal wing of the quadrate, and more posteriorly forms a
straight suture with the parietal. In this way the squamosal forms the external
portion of the floored area of the superior temporal fossa. Anteriorly a small
area of the squamosal is visible on the palatal surface, the squamosal at this
point being bent very sharply back on itself. However, this is probably the
result of dorso-ventral compression during preservation; this region would
have been more gently rounded in life.
Quadratojugal
The anterior third of the floor of the otic recess is formed by this element,
which lies with its longitudinal axis oblique to the long axis of the skull. In this
way, the quadratojugal forms the posterior margin of the inferior temporal
fenestra and makes an acute angle with the lower temporal arch. It is a thin,
unscuiptured and rather fragile element.
The quadratojugal extends back as a wedge of bone between the quadrate
and the jugal. It forms an oblique suture with the quadrate, and an overlapping
suture with the jugal. Anteriorly the quadratojugal meets the squamosal in a
horizontal suture which runs across the internal surface of the upper temporal
arch. Internally the quadratojugal is sutured to the palatal extension of the
squamosal. Medially it cannot be ascertained whether the quadratojugal meets
the postorbital.
Quadrate (Fig. 4)
This forms the greater part of the floor of the otic recess, and, like the
quadratojugal, it is unsculptured and lies in a strongly inclined position. Its
upper end meets the ventral surface of the squamosal along the lateral margin
of the superior temporal fenestra to form the otic recess.
The lateral surface of the quadrate is markedly fenestrated, and dorsal
contact with the squamosal is made by slender bars of bone. In recent croco-
dilians the tympanic cavity is linked to air passages within the quadrate and
supraoccipital, and by an elaborate system of cavities and tubes to the throat.
The posterior quadrate contact in Orthosuchus forms a somewhat expanded
head, socketed immediately beneath the rear end of the squamosal in typical
archosaurian fashion. The posterior margin of the quadrate within the otic
recess is curved, forming a distinct otic notch. This, together with the over-
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 241
hanging tip of the squamosal and paroccipital process, forms the bony housing
for the tympanic membrane (Fig. 4). The posterior quadrate contact is lost in
recent crocodiles, and indeed is also absent in teleosaurs such as Pelagosaurus.
In living crocodiles the squamosal, together with the opisthotic, extends
down to gain contact with the rear margin of the quadrate, so closing the notch
posteriorly. This can be seen in crocodilians of Upper Cretaceous age such as
Leidyosuchus. The closure of the otic notch in these forms has meant that the
tympanic membrane now adheres to bone all along its periphery.
Although the exoccipital in Orthosuchus is incomplete, it clearly overlaps
the major portion of the posterior margin of the quadrate. These two bones are
closely apposed, and the exoccipital blocks the major part of the primitive area
of the cranioquadrate passage.
X
yt
Mi)
// ") Vy
WMG
Fig. 4. Orthosuchus stormbergi. Reconstruction of the right
quadrate of the skull, seen in postero-lateral view. The lateral
flange of the squamosal has been removed to show the otic
notch and suggested position of the tympanum (x 14).
The palatal process of the quadrate passes forward and curves around the
anterior margin of the floored part of the superior temporal fossa. In this way
the quadrate forms a saddle-shaped area wedged in between parietal and
squamosal dorsally within the superior temporal fossa. Internally, the palatal
wing, as seen on the right side of the type specimen, meets the laterosphenoid in
a short suture which runs vertically down from the superior temporal fossa.
A semi-lunar foramen lies on this suture, and this no doubt gave passage to the
three divisions of the trigeminal nerve. Below the foramen, the quadrate is
sutured to the pterygoid, the suture continuing its vertical passage down to the
basicranium. The quadrato-pterygoid suture then runs back for a short distance
parallel to the midline before running obliquely out to the medial edge of the
condylar surface. The articular surface for the lower jaw consists of two
condyles separated by a shallow groove. The internal condyle is the larger of
the two.
242 ANNALS OF THE SOUTH AFRICAN MUSEUM
The quadrate is certainly firmly fused to the pterygoid and doubtless to the
prootic also. Unfortunately, very little of the medial end of the quadrate can be
made out, and it is in a poor state of preservation.
Pterygoid (Fig. 5)
The pterygoid is an extensive element. It probably reaches forward to the
choanae, and extends far back to the occiput. It forms a narrow plate of bone
on the floor of the basicranium where the pterygoids meet in the midline.
Posteriorly the pterygoids diverge to form a narrow wing on each side which
passes back to unite with the exoccipital. Medially the pterygoid wing overlaps
the basisphenoid, forming a curved suture. Laterally the pterygoid is sutured to
the palatal wing of the quadrate.
The quadrate ramus of the pterygoid extends vertically over the anterior
wall of the braincase, as in all crocodiles. Dorsally this ramus unites with the
laterosphenoid at the level of the foramen for the trigeminal nerve. Below the
quadrate ramus, the pterygoid extends in a latero-posterior direction as a
prominent pterygoid flange which forms an overlapping suture with the ecto-
pterygoid. The outer edge of the flange is made by the ectopterygoid ventrally
and by the pterygoid dorsally, much as in Gavialis. The pterygoid flange lies
farther forward in Orthosuchus, and although it is somewhat inclined ventrally
it does not descend as steeply as it does in living crocodiles. Each flange bears a
facet which guides the lower jaw on closure. ;
The rear edge of the pterygoid flange of Orthosuchus is considerably
thickened to form a horizontal ridge. Medially the ridge divides to run in both
directions along the basicranium parallel to the midline. In addition, the
pterygoids form a continuous median ridge which runs forwards from the
pterygoid-basisphenoid suture to become confluent with the vomerine
septum.
Immediately in front of the pterygoid flange, a triangular-shaped plate of
bone extends forwards on each side to the level of the antorbital fenestra. The
exact contribution of pterygoid and palatine to this area is problematical since
the bone is poorly preserved. It seems likely that the pterygo-palatine suture runs
transversely just in front of the pterygoid flange, the pterygoid extending
forwards in the midline to meet the vomer. However, there is no possibility of
verifying the presence or absence of this suture.
Ectopterygoid
This element meets the lateral edge of the pterygoid flange and then
extends laterally as a flat bar of bone. Externally it meets the jugal, and may well
also unite with an internal extension of the postorbital. It is unlikely that the
ectopterygoid has any sutural contact with the maxilla as it has in modern
crocodiles. The anterior margin of the ectopterygoid forms the major part of the
posterior border of the post-palatine fenestra. Its posterior margin forms the
outer half of the anterior border of the pterygoid fossa.
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 243
——
=
=
= ——
\
}
Fig. 5. Orthosuchus stormbergi. Reconstruction of the palate
(natural size).
Palate (Fig. 5)
The type specimen of Orthosuchus is unique among the protosuchians in
that the palate is well preserved. Although sutures are not entirely clear, there is
little doubt as to the extent of the various elements. Elongate choanae open
behind a short secondary palate made by the premaxillae and maxillae. The
choanae correspond to what Huxley (1877) called ‘the primitive posterior nares
of the Crocodilia’. In modern crocodiles these lie between the septum formed by
the vomers and the anterior processes of the pterygoids centrally, the palatines
behind and the maxillae in front and at the sides, and this is exactly the case in
Orthosuchus.
In Orthosuchus the median septum formed by the vomers is continuous with
a median pterygoid septum which runs to the rear of the skull. Laterally, each
vomer forms a deep channel which opens on to the pterygoid. In this way the
bony secondary palate opens into a pair of narial tubes formed by the vomers
244 ANNALS OF THE SOUTH AFRICAN MUSEUM
anteriorly and the pterygoids posteriorly. Orthosuchus differs from living
crocodiles in that these tubes are not floored by bone, though there can be little
doubt that they were covered by membrane in life. Hence the functional choana
opened on the rear margin of the pterygoid in much the same position as in
recent crocodiles.
The shape of the palatine is also suggestive of its incipient inclusion in the
formation of the secondary palate. Anteriorly the palatine forms an oblique
suture with the maxilla, posterior to the alveolar border. Behind this the
palatine twists along its length and its lateral border appears to be in the process
of rotating ventro-medially to attain the tubular shape that characterizes this
element in living crocodiles.
Occiput and braincase (Fig. 6)
The occiput of the type specimen is distorted because of the dorso-ventral
compression that has occurred in this region, and it would have been more
nearly vertical in the natural state. Telescoping of the right quadrate, together
with reduction in height of the foramen magnum, indicates the degree of
compression that has occurred.
Fig. 6. Orthosuchus stormbergi. Reconstruction of the occiput
(natural size).
Supraoccipital
This is an unpaired, triangular-shaped element which bears a median
vertical ridge. It meets the parietals above in a horizontally running suture, and
the exoccipitals below in an oblique suture. There is no evidence of a dermo-
supraoccipital bone.
Exoccipital
The exoccipital forms the superior margin and lateral boundary of the
foramen magnum, and also a small part of the occipital condyle, as in modern
crocodiles. On either side, the exoccipital fuses with the large paroccipital wing
of the opisthotic and divides laterally into two processes, as it does in the
teleosaur Pelagosaurus. The more extensive upper wing is sutured dorsally to the
squamosal, and the lower wing overlaps the posterior margin of the quadrate.
The main trunk of the facial nerve, the ceratohyal, the orbitotemporal artery
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 245
and the lateral head vein would then have passed back from the middle ear
region, probably along the grooved exoccipital-quadrate suture. In living
crocodiles this groove is no longer open, but is transformed into a narrow canal
by the much broader fusion of the quadrate with the upper paroccipital wing.
Ventrally the exoccipital unites with the basioccipital medially, and more
laterally with the pterygoid. Although the exoccipital has suffered some damage,
there appears to be evidence of a small foramen lying dorsal to the exoccipital-
pterygoid suture. This probably provided passage for the internal carotid artery
and may also have transmitted the vagus and hypoglossal nerves. In living
crocodilians twin foramina lie to either side of the foramen magnum. The inner
carries the hypoglossal nerve, while the glossopharyngeal, vagus and accessory
nerves and vein pass through the larger vagus foramen. Antero-ventrally to this
lies a foramen through which the internal carotid enters the middle ear.
Basioccipital
Although the major part of the occipital condyle is formed by the basi-
occipital, this element forms only a small median part of the ventral border of
the foramen magnum. The remainder of the margin is made by the exoccipital.
The condyle is clearly delimited and is oval in shape. Laterally the basioccipital
meets the exoccipital, and although this suture is difficult to discern, it must run
downwards and outwards to the lower boundary of the rear margin of the skull.
The basioccipital continues forwards on the ventral surface of the skull for only
a very short distance before uniting with the basisphenoid in a curved suture. In
this region there is a small basal tuber on either side for the ventral neck muscles.
Basisphenoid
The basisphenoid is seen as a half-moon-shaped element interposed
between the basioccipital and the pterygoids, although very probably it extends
far forwards above the pterygoids. Small basipterygoid processes lie behind the
basisphenoid-pterygoid suture. A very deep pit is located in the midline and
paired slit-like foramina can be seen lateral to each basipterygoid process. These
foramina lie within the basisphenoid, although the anterior margin of each is
formed by the pterygoid. There can be little doubt that they are the openings of
the eustachian system, although no connection between median and lateral pits
can be discerned. Within the median pit a narrow channel runs forwards above
the pterygoid, and a larger channel runs back into the body of the basisphenoid.
Living crocodiles have both lateral and a large median eustachian opening,
but they lie between the basisphenoid and basioccipital. They lead upward bya
complex system of tubes to the ear, and downward by three membranous tubes
to the throat, eventually forming one tube which opens almost at the level of the
choanae.
Laterosphenoid
As seen on the right side of the type specimen, the laterosphenoid has the
same form and relationship to other elements as found in living crocodilians.
246 ANNALS OF THE SOUTH AFRICAN MUSEUM
Antero-dorsally the laterosphenoid articulates with the internal surface of the
frontal, and also extends laterally and probably just touches the postorbital.
Postero-dorsally the laterosphenoid unites with the internal surface of the
parietal. The laterosphenoid-quadrate suture runs perpendicularly down to the
foramen which transmitted the trigeminal nerve. Ventrally the laterosphenoid
meets the quadrate ramus of the pterygoid.
Otic region
Nothing can be said of either the prootic or epiotic, since these elements are
concealed by the quadrate, parietal and supraoccipital. The opisthotic forms the
large, horizontally directed paroccipital process to which the exoccipital is fused.
Medially it fuses with the supraoccipital and dorsally it is sutured to the
squamosal. Internal to the quadrate, fragments of bone can be seen and very
probably the otic elements were crushed during preservation due to compression
of the skull in this region. The stapes is also unknown.
THE LOWER JAW (Fig. 7)
Both mandibular rami of the type specimen are well preserved although
slightly distorted. The middle area of the right has been restored, and the
articular region is firmly attached to the condylar surface of the quadrate. The
articular region of the left ramus is incompletely preserved. In K4639, the rear
halves of both rami are visible although the posterior margin is incomplete in .
both cases.
Dentary
This is the largest of the lower jaw elements and externally forms the lower
half of the mandible. It is sculptured in the form of numerous small, shallow pits
which grade into slit-like markings, more sparsely arranged, farther back. A
Fig. 7. Orthosuchus stormbergi. Reconstruction of the lower jaw (natural size). A. Lateral
view. B. Medial view.
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 247
careful comparison of the alveolar border of both rami suggests that there were
probably 15 dentary teeth, though up to 18 could have been present. The teeth,
like those of the upper jaw, are conical, longitudinally striated and somewhat
recurved. There is no differentiation of tooth shape and there appear to be no
marked differences in tooth size, apart from those reflecting the pattern of tooth
replacement.
The mandibular symphysis is short and stout, and is complete at the level
of the fourth mandibular tooth. Extending back from the middle of the
symphysis is a longitudinally running groove, the Meckelian canal. This carried
the mandibular artery, vein and nerves, and was almost certainly covered by the
splenial in life. This latter element may have entered the symphysis; on both
rami the anterior border of the splenial is incomplete. Posteriorly the dentary
extends to the anterior margin of a large, elongate external fenestra which lies
between the dentary, angular and surangular.
Splenial
This element lies wholly on the anterior inner surface of the mandible,
uniting with the dentary above and below. The exact position of its posterior
margin is uncertain, as suture lines are difficult to distinguish in this region.
Probably the splenial extended back to the mid-region of the ramus, where it
united with the coronoid and, below this, to a smaller extent with the angular.
Coronoid
The single coronoid element forms the anterior margin of an extensive
adductor fossa, and occupies much the same position as it does in recent
crocodiles. Posteriorly the coronoid has a rounded edge which overlaps the
surangular. The suture between these elements then runs forwards just below the
dorsal margin of the ramus on its internal surface.
Surangular
The surangular extends over the external mandibular fenestra as a narrow
flange, and in this respect the element differs from that of living crocodiles.
Externally the posterior region of the surangular forms an overlapping suture
with the angular. Internally it borders the adductor fossa dorsally, and behind
this a descending process meets the prearticular ventrally. Posteriorly the
surangular is overlapped medially by the articular.
Angular
This is a large postero-ventral element which forms the ventral border of
the external mandibular fenestra. The angular wraps round the rear margin of
the mandible, and is sutured internally to the articular. A retroarticular process
is not developed. On the inner surface, the upper margin of the angular is
bordered by the prearticular. These two elements diverge anteriorly to form a
slit-like foramen, of which the anterior boundary is formed by the coronoid. This
248 ANNALS OF THE SOUTH AFRICAN MUSEUM
corresponds to the Meckelian foramen which, in living crocodilians, lies between
the angular and splenial.
Prearticular
This element is not normally found in crocodilians, but it occurs in early
mesosuchians such as Pelagosaurus and Metriorhynchus. In Orthosuchus it is a
slender, elongate element which forms the ventral margin of the adductor fossa.
Anteriorly it is overlapped by the coronoid, and may have extended forward
medially to this element to contribute to the inner wall of the Meckelian canal.
The foramen which transmits the chorda tympani branch of the facial nerve,
and which is almost universally present on the inner surface of the prearticular,
cannot be discerned. However, there is a groove running along its inner surface.
Posteriorly the prearticular lies over the anterior edge of the articular.
Articular
On the right side this element is fused by matrix to the condylar surface of
the quadrate, while only fragments of the left articular are preserved. The
glenoid fossa cannot be seen, although from the contour of the articular area it
appears to consist of a large internal and smaller external articular surfaces.
THE VERTEBRAL COLUMN AND RIBS
General features
The presacral series is complete but for a gap which corresponds to three
mid-dorsal vertebrae, giving a presacral count of 24. Of these, probably eight —
may be regarded as cervicals. Two sacral vertebrae are in articulation with the
most anterior caudal vertebra. Seven other caudal vertebrae are known, of
which only one is in a reasonable state of preservation.
The vertebral column shows regional differentiation both in the progressive
Fig. 8. Orthosuchus stormbergi. Anterior (upper row), and posterior
(lower row) views of vertebrae (natural size). A. Cervical six.
B. Cervical five. C & D. Dorsal seventeen. E & F. Dorsal twenty-
four. G & H. Caudal four.
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 249
changes that occur in the rib articulation facets, and in the overall dimensions of
the vertebrae. The centra of the anterior cervicals are short, and a progressive
increase in length occurs through to the lumbar region, as shown in Table 1.
The vertebrae are spool-shaped and, unlike the procoelous vertebrae of
eusuchians, in Orthosuchus they are all amphicoelous. In the majority of the
vertebrae a clear sutural union of the neural arch on the centrum is visible. The
neural spines are not high in any region of the column, but are tallest in the
cervical region and decrease slightly through to the lumbar region. The articular
surfaces of the zygapophyses are more or less vertical on the cervicals, but in the
mid-dorsals they change to a more nearly horizontal direction. Posterior to this
they again become more nearly vertical in orientation (Fig. 8).
Table 1
Measurements of the vertebrae in mm
Length of centrum (L). Height of centrum posteriorly (H). Width across
posterior end of centrum (W). Height of neural spine (NH). Maximum
height of vertebra (VH).
Vertebra No. L H W NH VH
Cervical
Atlas : : == == == = 15,5
Axis . ; : Teal 7,9 Sy) 8,0 17,0
Sie, ‘ ; 6,4 8,5 6,0 — —
6,6 8,5 6,0 8,0 21,0
5 6,8 Tel 6,0 8,0 21,0
6 6,8 — 6,0 20,0
i 6,8 7,6 6,0 6,0 19,0
8 6,8 6,5 6,0 18,0
Dorsal
9 4 6,5 6,5 6,0 18,0
10 8,4 = — 6,0 18,0
11 8,4 a= == 6,0 18,0
12 8,8 3 6,5 6,0 18,0
13 8,8 7,3 6,0 6,0 18,0
14 a = = cms
15 — _— oes = —
16 nes == = — —
17 10,0 7,0 6,5 6,0 18,0
18 10,0 iS 6,5 6,0 18,0
19 10,5 7 6,5 6,0 18,0
20 10,5 7,5 6,0 6,0 18,0
21 10,0 8,0 6,7 5,0 19,0
22 9,6 8,5 — 5,0 19,5
23. ; ‘ 9,6 8,5 7,0 5,0 20,0
2a ; : 9,6 8,5 8,7 5,0 18,5
Sacral
ae : , 10,0 8,5 es 7,0 18,0
Dis : 10,0 8,0 7,0 7,0 18,0
Caudal
i. 4 : 8,0 7,0 6,2 —_ 19,0
Dey. : d 77 7,0 6,4 — —
Bx. ; . 7,7 15 6,0 9,0 20,0
vs i : 7,7 — 6,0 9,0 20,0
Mid-caudal 9,0 3,5 4,5 — —
(isolated vertebra)
250 ANNALS OF THE SOUTH AFRICAN MUSEUM
Atlas and axis (Fig. 9)
The intercentrum of the atlas is well developed, and in cross-section forms
a dumb-bell shape. A single-headed rib is in articulation with the intercentrum
postero-laterally, while dorso-laterally the intercentrum supports the stout base
of the pedicel of the neural half-arch on each side. Dorsally a narrow gap
separates the two neural half-arches. Presumably this gap was filled by cartilage
and overlapped by a pro-atlas in life, as in recent forms. Posteriorly the dorsal
portion of the neural arch is drawn back to form an almost horizontal post-
zygapophysis which overlaps the axis. The prezygapophysis of the axis with
which it articulates is not visible, but is probably quite small.
The centrum of the atlas, the odontoid process, can be seen lying between
the pedicel bases of the neural half-arches of the atlas. The anterior face of this
process articulates with the occipital condyle, as in living crocodiles. The
postero-lateral margin of the odontoid process together with the adjacent edge
of the axis centrum bears an articular facet for the second rib. This rib is closely
apposed to the first, and is also single-headed.
The axis centrum is very stout, and posteriorly is pulled down to form a
hypapophysis. The neural spine is long and low, and is pointed anteriorly where
it projects between the atlas neural half-arches.
Fig. 9. Orthosuchus stormbergi. Vertebrae
(natural size). A. Atlas seen in anterior view.
B. Atlas, axis and third cervical seen from the
left side. C. Anterior presacrals, vertebrae
eight to twelve inclusive, seen from the
right side.
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 2a
Cervical vertebrae 3 to 8
The most anterior vertebrae are strongly keeled and the centrum of each
bears both an anterior and a posterior hypapophysis. The keeling becomes
progressively less marked through to the posterior cervicals, and is absent on
vertebra eight. In modern crocodiles this keeling is extended to the anterior
dorsal vertebrae, and all bear a hypapophysis.
Laterally each centrum bears a prominent parapophysis for the articulation
of the capitulum. Each parapophysis is broad anteriorly and tapers to the mid-
region of the centrum. On the third vertebra, the parapophysis lies near to the
ventral surface, but through the succeeding vertebrae it gradually alters its
position so that on vertebra eight it is borne near to the neurocentral suture.
The neural arch has a pronounced diapophysis on each side for the tuber-
culum. The position and shape of the diapophysis also changes progressively
along the length of the cervical vertebrae. On the third the diapophysis is a
thickened ridge along the anterior half of the neurocentral suture. Through the
cervical series this becomes progressively raised on a transverse process.
Dorsal vertebrae (Fig. 9)
The progressive changes of the positions of the articular facets for the ribs
which occur along the cervical series continue through the anterior dorsal
vertebrae. The parapophysis continues to move dorsally up the anterior margin
of the centrum, coming to lie beneath the diapophysis on vertebra twelve.
Coupled with this movement, the parapophysis also becomes larger and more
round in shape. The diapophysis does not alter its position from that seen on the
eighth cervical, though both the transverse process and the diapophysis become
progressively broader through to the twelfth vertebra.
Behind this the parapophysis continues to migrate upward, coming to lie
on the same level, although still separate from the diapophysis on vertebra
twenty-two. Both the facets and the transverse processes then diminish in size
towards the pelvis so that on the last presacral the transverse process is quite
slender and short and the two facets fused. In modern crocodiles these facets are
confluent and the dorsal ribs single-headed on the eighteenth vertebra. In this
way the transverse processes become narrower earlier on in the series.
Sacral vertebrae (Fig. 10)
The centra of the sacrals are long, slightly exceeding the length of the
posterior dorsals. The transverse process of the first sacral vertebra is stout and
short, as is the first sacral rib. In addition to its articulation with the transverse
process, the rib also unites with the centra of both the last presacral and first
sacral vertebrae. An identical condition is found in living crocodilians. The
transverse process of the second sacral vertebra faces somewhat posteriorly.
Distally it articulates with a very broad rib, which, as in modern crocodiles, also
articulates with the posterior portion of the lateral surface of the centrum.
The sacral ribs form a very strong support for the ilium. The articular areas
252 ANNALS OF THE SOUTH AFRICAN MUSEUM
are triangular in shape in both cases, though that between the second sacral rib
and the ilium is the more extensive. Only a very small median portion of the
ilium is free.
Caudal vertebrae
Only eight caudal vertebrae are known. One of these is in articulation with
the sacral series, and three others are from the proximal region of the tail. The
remainder have small, elongate centra and must be mid-caudal vertebrae.
Probably the tail was long, of the order of 30 to 40 vertebrae.
The neural spines of the anterior caudals are taller than elsewhere in the
Fig. 10. Orthosuchus stormbergi. Sacrum, corrected for distortion
(natural size). A. Dorsal view with sacral ribs and internal surfaces of the
ilia. B. Ventral view with sacral ribs. C. First sacral vertebra seen in
anterior view.
column, and are directed slightly backwards. The zygapophyses slope so that
their facets meet more or less vertically. The first chevrons probably lie between
the third and fourth caudal vertebrae. The fourth bears a nearly entire caudal
rib, although elsewhere these are broken above their bases.
Ribs (Fig. 11)
The proximal ends of the atiantal and second rib are present on both sides.
They are slender and single-headed. The first has a large articular facet for union
with the intercentrum of the atlas, and the second rib has a smaller facet for
union partly with the odontoid and partly with the centrum of the axis. This
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 253
differs from modern crocodiles where the axial rib is double-headed, the second
facet lying on the odontoid.
Only one other cervical rib is complete, the fifth, and this is indistinguish-
able in type from that of living crocodiles. It is double-headed and has a short
shaft which runs parallel to the long axis of the vertebral column. The capitular
and tubercular processes rise at right angles to the shaft of the rib and diverge
as they pass upward to the centrum of the vertebra.
The shaft of the eighth cervical rib is transitional in type between that of
the cervicals and anterior dorsals. It more closely resembles that of the latter
though it is much more slender.
Behind this, on the left side, dorsal ribs nine to fourteen inclusive are
preserved in situ. Three right dorsal ribs were also associated with the material.
All the dorsal ribs are double-headed and their shafts long and strongly curved.
The articular facets for the union of rib with vertebra move further apart
passing from the sixth cervical back to vertebra nine, and are at their widest on
this vertebra. Posterior to this they come closer together. Further, in each case,
the tubercular process lies above and anterior to the capitulum, the two being
separated by a shallow groove. In the ninth rib, the tubercular process is the
larger of the two, but from the twelfth this is altered and the rib articulates
principally by the capitular process.
Ribs nine to fourteen are expanded to form prominent antero-ventral
and postero-dorsal flanges. Because of this, the mid-dorsal ribs particularly
are very flat and broad proximally, and narrow abruptly to a cylindrical
shaft.
The antero-ventral flange rises immediately behind the head of the rib
much as in living crocodiles. In recent genera, the flange is limited to the first
two to four dorsal ribs, the number being greater in older individuals. The
postero-dorsal flange is more gently rounded in shape and is developed slightly
lower down the shaft of the rib. Presumably this flange is homologous with the
cartilaginous ‘uncinate’ process which is developed in this position in living
crocodiles. This process is normally carried on the third to fifth dorsal ribs, and
may occasionally ossify slightly. Both anterior and posterior flanges serve for
muscle attachment. In Orthosuchus the postero-dorsal flange of one rib overlaps
the antero-ventral flange of the succeeding rib.
Posteriorly only the proximal part of ribs in articulation on the left side of
vertebrae eighteen and nineteen are known. These ribs are double-headed, but,
unlike that of the anterior dorsals, the capitular and tubercular processes lie on
the same level. The capitular process is the larger of the two. Each rib is strongly
curved backwards, and is broad and flat immediately behind its head. The rib
rapidly contracts to a cylindrical shaft and there is no evidence of either an
anterior or a posterior flange. Probably at least the last two presacral vertebrae
did not bear ribs.
Overlying vertebrae nineteen and twenty on the right side is an accumula-
tion of the remains of several fine gastralia.
254 ANNALS OF THE SOUTH AFRICAN MUSEUM
THE LIMB GIRDLES AND LIMBS
Pectoral girdle (Fig. 12)
The shoulder girdle shows a remarkable approach towards that seen in
recent crocodiles. It consists of scapula and coracoid only. Both scapulae are
preserved entire, although the right has been somewhat flattened during
preservation. Of the coracoids, the right is fractured and its distal margin
incomplete, while only the proximal end of the left coracoid is present.
The scapula is a tall element, with its superior end considerably expanded
Fig. 11. Orthosuchus stormbergi. Lateral (upper row), and medial
(middle and lower rows) views of ribs (natural size). A & B. Right
fifth cervical. C & D. Right first dorsal. E & F. Right mid-dorsal.
G. Left tenth dorsal. H. Left mid-dorsals showing overlapping of
the rib flanges.
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 255
and drawn out posteriorly. It narrows to a stout shaft which curves sharply
inwards to form a more conservatively expanded inferior region. Charac-
teristically, the posterior margin of this bears a large rough facet which forms
the upper half of the glenoid surface. Another triangular-shaped facet for the
coracoid lies along the lower edge, its broadest part lying posteriorly and below
the glenoid facet. The anterior margin of the lower region of the scapula is
overhung by a ridge, just as in recent forms. Below this ridge, the concave surface
afforded attachment for a large muscle, the scapulo-humeralis posterior.
Fig. 12. Orthosuchus stormbergi. Pectoral girdle
(natural size). A. Lateral view of right side.
B. Anterior view of left scapula.
The coracoid is half as long as the scapula. Its upper region is expanded to
match the lower end of the scapula, while the posterior margin is thickened to
form the lower half of the glenoid. In front of this lies a prominent coracoid
foramen. Distally the coracoid flares out to form a blade-like inferior end. The
long axis of the coracoid, like that of the scapula, lies obliquely orientated so
that in life the bone would have been directed inwards.
The sternum and interclavicle are unknown, and were presumably
cartilaginous in life.
Fore-limb (Figs 13-14)
The fore-limb is completely known from the left side and, in addition, part
of the lower right fore-limb is present.
In general shape the humerus is remarkably similar to that of living
crocodiles. The shaft is well developed and slender. It is twisted along its length,
so that with the proximal expansion lying antero-medially, the distal expansion
faces antero-laterally. In living crocodiles the proximal and distal expansions
lie more or less in the same plane.
256 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 13. Orthosuchus stormbergi. Fore-limb (natural size).
A-C. Left humerus. A. Anterior. B. Posterior. C. Lateral
views. D. Lateral view of the radius. E. Lateral view of the
ulna.
In both Orthosuchus and in living crocodiles, the proximal expansion is of
moderate size, although in recent forms it extends farther medially. The
articular surface lies along the upper posterior edge and is oval in shape. The
deltopectoral crest originates on the lateral edge of the proximal expansion and
curves obliquely down across the anterior face of the shaft. This crest is more
strongly developed in Orthosuchus than it is in living crocodiles, and encloses a
deeper concavity on the anterior surface of the humerus. Distally the bone is
thickened into two condyles. The capitellum is marginally the larger of the two,
and is separated from the trochlea by a shallow groove. There is little projection
of ectepicondyle or entepicondyle, and distal foramina are lacking.
The radius is a slender bone. Proximally it is thickened where it touches and
partly obscures the proximal expansion of the ulna. The ulna extends beyond
a
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 257
Fig. 14. Orthosuchus stormbergi.
Lower fore-limb (natural size).
Anterior view of the left radius,
ulna, carpus and manus.
the limit of the radius, but it does not form an olecranon. The facet for the
humerus is terminal. The articular surface of the ulna faces both forward and
upward.
Distally the radius is expanded medio-laterally and forms an oval articular
surface for the radiale. The ulna is also expanded distally, but in the antero-
posterior direction. The ulna articulates distally with the pisiform and ulnare,
and meets the radiale internally.
As in modern crocodiles, the radius is the shortest of the long bones while
the ulna is the next shortest. However, the proximal part of the fore-limb is only
marginally longer than the distal, the length of the radius being 88 % that of the
humerus. As is typical of archosaurs, the ulna is stronger than the radius and
would have carried the major part of the weight borne by the fore-limb.
The carpus shows the same remarkable degree of specialization found in
modern crocodiles, for the radiale and ulnare are elongated and so add an extra
segment to the fore-foot. Of the two, the radiale is the longer and stouter element.
Proximally the radiale is greatly enlarged to a triangular head which articulates
258 ANNALS OF THE SOUTH AFRICAN MUSEUM
with both radius and ulna. Consequently the weight of the body could be trans-
mitted from both the ulna and radius through the radiale to the middle and
inner digits. The pisiform is a large, flat, kidney-shaped bone which interposes
posteriorly between the ulna and ulnare.
The distal row of carpals consists of two elements and these are not
elongated. The larger of the two is somewhat angular and lies distal to the
radiale. This presumably represents a fusion of distal one and a centrale. The
second is more elliptical in shape, and is formed by fusion of distals three and
four. In modern forms, generally, only one distal carpal is ossified though two
other cartilaginous elements may be present.
Despite the fragile nature of the manus, five digits are known. However,
only the first of these is complete, although very little is missing from the second,
fourth and fifth. About half of the third digit is lacking. Clearly, the first digit is
both the shortest and the stoutest, and the fifth the weakest. Further, the second
and fourth digits are longer than the first, and digit three was probably the
longest.
Each metacarpal is expanded proximally. The first has a shallow articular
surface for the medial distal carpal, which also meets flat articular surfaces on
metacarpals two and three. The proximal articular surfaces on the fourth and
fifth metacarpals are in articulation with the lateral distal carpal. The meta-
carpals overlap one another proximally from medial to lateral sides. Each
metacarpal is also expanded distally and forms a convex articular surface.
Similarly, the phalanges form articular surfaces that are concave proximally,
and convex distally. The first digit bears two phalanges, the terminal phalanx
being a claw. The second digit has two phalanges and the proximal part of a
third. Probably this terminal phalanx is a claw, though it is impossible to be sure.
Of the third digit, only the proximal phalanx and part of the next are known.
The fourth and fifth digits are very weak, and because of this the number of
phalanges present in each case cannot be determined with any certainty.
Probably three phalanges of the fourth, and two of the fifth, are known. The
phalangeal formula was probably 2, 3, 4, 25, 3.
Pelvic girdle (Fig. 15)
The pelvis is complete, although some fragments of bone are missing. The
left pubis was removed to make it possible to clean all the vertebrae. It
originally occupied a position that could have been little changed from the one
held in life. The right pubis is still in situ, but it is displaced from the pelvis. The
ilium on that side is somewhat compressed dorso-ventrally.
The pelvic girdle is remarkably crocodilian in form in that the pubis is
excluded from the acetabulum by the ischium. The acetabulum is large and deep,
and is perforated ventrally. Dorsally a very prominent ridge, the supra-
acetabular buttress, overhangs the acetabulum. In life the femur would have
exerted its thrust against this.
Above the buttress, the blade of the ilium is narrow and is produced
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 259
y) _ NN
Fig. 15 Orthosuchus Stormbergi. Pelvic girdle (natural size).
Lateral view of left side.
anteriorly to form a long, pointed process. This process is reduced in living
crocodiles. Posteriorly the ilium forms a more robust post-acetabular process,
much as in living crocodiles. The outer surface of the blade of the ilium, both
above the supra-acetabular buttress and on the post-acetabular process, is
covered by a series of fine striations. In life, the major extensor of the leg
musculature, the m. ilio-tibialis and the smaller extensor, the m. ilio-fibularis,
would have originated here, as would the powerful abductor, the m. ilio-
femoralis.
Ventrally the ilium forms two stout processes which between them form the
roof and walls of the acetabular opening. The most anterior of these bears two
rounded facets, the upper of which articulates with the pubis; the lower meets
the pubic process of the ischium. The posterior process bordering the acetabulum
also meets the ischium.
Internally, the visible surface of the ilium is smooth, although apart from a
very short median region, the whole of the internal acetabular and post-
acetabular surfaces are supported by a very firm union with the two sacral ribs
(Fig. 10).
The long axis of the ischium passes obliquely back in the dorso-ventral
plane. The main sutural contact with the ilium is posterior, and in this region the
ischium is thickened to form a broad margin to the postero-ventral edge of the
acetabulum. The ischium also sends forward a narrow pubic process which
curves round the ventral and ventro-lateral margin of the acetabulum. It bears a
rounded surface on its leading edge for articulation with the pubis, and above
this for the ilium.
Below this the ischium forms a short, narrow shaft. On both sides the ischia
are fractured at this point, reflecting the distortion of the pelvis. Distally the
ischium is expanded, particularly posteriorly, and its lower margin is striated.
The ischia meet ventrally in the midline.
The pubis is a remarkably rod-like element, flattened on its lateral surface.
It is nearly as long as the ilium and is longer than the ischium. Proximally the
260 ANNALS OF THE SOUTH AFRICAN MUSEUM
head is slightly expanded and bears a large articular surface for the ilium.
Below this lies a smaller surface for articulation with the pubic process of the
ischium. In modern crocodilians contact with the ilium has been lost and the
pubis articulates only with the ischium.
Behind the articular surface there is a shallow depression laterally on the
head of the pubis. This area represents the pubic rim, which is well developed in
early reptiles, and is seen in forms like Howesia. Below this there is torsion in
the pubic shaft so that proximal and distal expansions are at right angles to each
other. The distal expansion is not great, though it is almost twice the size of the
proximal expansion. Clearly the pubes were not fused distally to form a pubic
plate, but it is probable they met distally in cartilage, as they do in modern
crocodiles.
Hind-limb (Figs 16-17)
Unfortunately, both femora are broken. The break in the right femur was
restored in the field, and very little of this bone can be missing. Assuming the
estimated femur length to be correct, then it is still the longest of the long bones,
the humerus being 79% of the length of the femur.
In form the femur is very similar to that of a modern crocodile. It has a long,
curved, somewhat flattened shaft with both proximal and distal ends expanded.
These expansions lie more or less obliquely to one another. There is no constric-
tion between the head and the shaft of the femur. The head has a pronounced .
articular surface which is convexly rounded and oval in shape. As in living
crocodiles, the articular surface is developed on the upper edge of the head, but
in Orthosuchus it extends farther medially.
Fig. 16. Orthosuchus stormbergi. Reconstruction of the
right femur (natural size). A. Anterior. B. Posterior.
C. Medial views.
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 261
Proximally on the posterior face of the femur a shallow depression is
present. This area represents the intertrochanteric fossa. Adjacent to this the
lateral margin of the bone is thickened and possibly represents the remnants of
the internal trochanteric crest. This area carries the insertion of the m. pubo-
ischio-femoralis externus. There is no development of the greater trochanter and
the ilio-femoralis musculature inserted farther down the shaft.
A thickening of bone towards the medial edge on the posterior surface of
the shaft represents the fourth trochanter, to which was attached the coccygeo-
femoral = caudifemoral musculature. The ratio of the distance from the
proximal end of the femur to the centre of the fourth trochanter, over the
distance from the distal end to the centre of the fourth trochanter in Orthosuchus
is 0,46. In a large specimen of Crocodylus acutus it is 0,55, but in two small
specimens of Alligator mississippiensis the ratio is the same as that for Ortho-
suchus. The distal condyles of the femur of Orthosuchus are well developed, the
lateral being the larger of the two.
The left tibia and fibula are entire, and are in articulation with the tarsus
and proximal pes. The proximal part of the right tibia, together with an
impression of the fibula, is in articulation with the femur. The distal parts of
these bones are in articulation with the astragalus and calcaneum.
The tibia is the stouter of the two bones and is particularly well developed
for articulation with the femur. It is expanded posteriorly to form a large
articular surface for the medial condyle of the femur, and also extends laterally
to contribute to the articular surface for the lateral condyle. The fibula is
modestly expanded posteriorly to complete this surface. Both articular surfaces
are concave. The shaft of both the tibia and fibula is circular in cross-section in
the mid-region, but lower down the inner surface becomes gently concave on the
tibia and flat on the fibula.
Distally the tibia is again more developed than the fibula. Laterally the
tibia is about equal in length to the fibula, but it extends farther down medially
to form a large area for articulation with the astragalus. The distal articular
surface of the tibia is rounded medially and has a flat, sloping lateral surface.
This follows a similar contour on the upper face of the astragalus. The fibula
articulates with the astragalus medially in a more or less vertical plane, and with
the calcaneum ventrally where its articular surface is slightly concave.
The tibia and fibula are marginally shorter than the femur, the length of the
tibia being 94% that of the femur. The fore-limb is distinctly shorter than the
hind when only the long bones are considered. However when the length of the
radiale is taken into account, the fore-limb is 91 °%% the length of the hind.
The astragalus and calcaneum of Orthosuchus are identical in shape with
these bones in living crocodiles. The astragalus is a very large element which
occupies both the medial and mid-region of the tarsus. It is somewhat flattened
in the mid-region, although on its medial border it is thickened and forms an
oval convexity anteriorly. The calcaneum is a small, rectangular element
anteriorly, but posteriorly it forms a very prominent tuber. On its posterior face
262
ANNALS OF THE SOUTH AFRICAN MUSEUM
=a
= —
mre LICHT i i]
1
a, ae
j
rs
Fig. 17. Orthosuchus stormbergi. Lower hind-limb (natural size). A-—C. Left
tibia and fibula, tarsus and metatarsus, as preserved. A. Anterior. B. Posterior.
C. Lateral views. D. Reconstruction of left tarsus and pes.
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 263
the tuber is grooved vertically for the passage of the gastrocnemius tendons.
The calcaneum also extends obliquely up over the posterior surface of the
astragalus.
The astragalus articulates on its upper surface both with the expanded
distal end of the tibia and with the medial surface of the fibula, the major
articulation here being between the astragalus and tibia. The astragalus also
articulates with the calcaneum. Superficially this articulation would seem to be
identical with that found in modern crocodiles and probably, as in these forms,
a small ball and socket articulation exists between these two bones. Movement
between them is therefore in the vertical plane and the joint is a crurotarsal one,
with the astragalus functioning with the crus, and the calcaneum with the pes.
In contrast to the articulation between tibia and astragalus, that between the
fibula and calcaneum is a freely movable one.
The astragalus articulates distally with the innermost of two distal tarsals
and metatarsal two. It clearly also met metatarsal one, though this is known only
from an impression. The calcaneum articulates distally with the lateral distal
tarsal. The tarsalia of Orthosuchus occupy the same position in the tarsus as is
found in living crocodiles.
The medial tarsal is the smaller of the two, and meets metatarsals two and
three distally. Presumably it represents tarsal three. The larger lateral tarsal
extends posteriorly and is probably tarsal four. Posteriorly it meets metatarsal
five, anteriorly it meets metatarsal four and just touches metatarsal three.
The metatarsals overlap one another proximally from medial to lateral
surfaces. The third seems to be the stoutest. The fifth is reduced to a hook-like
element which quite clearly lacks phalanges and is held behind the third and
fourth metatarsals. All the elongate metatarsals are incomplete, although the
distal end of one was found in association with a few phalanges of the left pes
overlying the mid-dorsal ribs. This is most likely part of metatarsal two, and the
digits are preserved in inverted sequence in relation to their metatarsals. The
distal end of the metatarsal is grooved on its dorsal surface near to the articular
region. Of this second digit, one phalanx and part of a second are known. A
phalanx and claw of digit one is present, although metatarsal one is known only
from an impression. One phalanx from each of digits three and four are known.
The proximal articular surface of each of these is concave, and its distal end
rounded.
BODY ARMOUR (Fig. 18)
Dorsally the body is covered by a double row of large rectangular scutes.
Neither ventral nor lateral scutes are present, nor is there any evidence which
suggests that the body was protected ventrally in this way. Virtually nothing
remains of the scutes of the caudal region.
The number of scute pairs corresponds to the number of vertebrae present.
In addition, a very small scute is present at the extreme anterior end of the
trunk, and this could correspond to the pro-atlas. The scutes increase in length,
264 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 18. Orthosuchus stormbergi. Dorsal scutes (natural size).
A. Dorsal view of anterior scutes, the most anterior pair are figured
entire. B. Ventral view of cervical scutes.
passing back along the length of the body in an identical manner to that which
occurs in the vertebrae. Hence, the scutes are shortest antero-posteriorly in the
cervical, and longest in the sacral region. A broadening of the scutes also occurs
back to the anterior dorsal series. Behind this they decrease in width. In the
sacral region their exposed dorsal surfaces are rectangular in shape.
The posterior edge of each scute overlies and conceals the anterior edge of
the following scute. Approximately one-third of the length of each scute is
concealed in this way. Where this border is visible it is seen to be smooth, while
the exposed surface of each scute is sculptured by a system of deep pits which
form a pattern similar to that on the skull. The extreme outer edge of each scute
bends sharply down to form a narrow lateral part. In this way, a sharp ridge
borders the lateral margin of each scute dorsally. In the cervical and dorsal
regions the outer anterior edge of each scute extends forwards as a blunt,
peg-like process which lies below the preceding scute.
MYOLOGY OF ORTHOSUCHUS
Although the restoration of musculature in an extinct reptile must always
involve an element of uncertainty, it seems likely that in Orthosuchus the muscle
arrangement could not have been very different from that found in living
crocodiles. The nomenclature used throughout follows that of Romer (1956),
and where this differs from that used by other workers in this field, their
nomenclature has been cited in parentheses.
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 265
Table 2
Measurements of the skull and postcranium in mm
K4639 K409
Length of skull, front of snout to parietal/supraoccipital suture. 54,0 90,0
Preorbital length, front of snout to anterior margin of orbit. : 25,0 46,0
Maximum breadth of skull across external borders of quadratojugals 38,0 70,7
Breadth of cranial table at mid-level of superior temporal fenestrae . 33,0 55,0
Breadth of intertemporal region at mid-level of eae temporal
fenestrae between their inner borders . 6,0 6,0
Breadth of interorbital region at mid-level between i inner borders of
orbits . : , : 11,0 16,0
Breadth of posterior region of the snout immediately anterior to orbits ca 4,5 26,0
Breadth of anterior region of snout across premaxillae . . . 10,0 17,4
Maximum length of superior temporal fenestra . . .. . , 10,0 19,0
Maximum length of inferior temporal fenestra. , 2 : : — 21,0
Maximum height of inferior temporal fenestra . : : ; : — 12,0
Maximum length of orbit : ; : : é ‘ : 13,0 17,0
Maximum length of antorbital fenestra ; : : — 13,0
Distance between tip of snout and anterior border of choana ene — 21,6
Length of lowerjaw. .. oe ts en ae : : : — caii0,0
Length of mandibular symphysis os ie Oa coe A 5 — 9,0
Length of external mandibular fenestra : paar a — 34,0
Maximum height of scapula . : : : ; . ; ; — 46,0
Maximum height of coracoid . ; : E . : — 23,0
Maximum length, antero-posteriorly, of superior ‘scapula : : : — ca 22,0
Maximum length, we of inferior coracoid blade. —_ 152
Length of humerus . : Z : : : : . i : — 45,0
Length of radius : Q | ! ‘ : : : : : : — 39,5
Length of ulna . : . : ; : : : : ; : : — 43,1
Length of radiale : ; : : ' ‘ : ; ' : : — 16,3
Length of ulnare : E : : ’ ‘ ‘ : : : : — 10,0
Length of metacarpal1 . : - : : ; i 2 ; : — 9,0
Length of metacarpal2 . : : : , . : : : : — 9,8
Length of metacarpal3 . c : : : ’ ; : : : — 10,5
Length of metacarpal4 . : E : : q : ’ : : — 9,0
Length of metacarpal5 . : : : 3 ; ; 4 ; : — 13
Length of digit 1 : : : : : : : ; : — 20,5
Maximum length of iliac blade : : ; J ; : : — 36,8
Height of ilium above acetabular fenestra. : ; : : : — 125
Maximum height of acetabulum . . . . . =. . . — 15,3
Length of pubis . : : : : : z d : : : : — 3355
Micadinvor distal pubis . . . . ... ee = 9,0
Height of ischium .. ht Mai nant iat as, — 22,0
Length of ventral ischial margin ial petite oe ae ae — 24,0
Length of femur : : — ca 57,0
Distance between proximal femur and centre of fourth trochanter ‘ — 18,0
Length of tibia . , : : é ; : . : : : : — 53.5
Length of fibula ee gk : Su eee : — 51,0
MenPime@iinciatdisal = A we A ~ 9,5
MUSCULATURE OF THE HEAD (Fig. 19)
A general account of the facial musculature of crocodilians given by
Von Wettstein (1937) was found helpful in this analysis. In addition, Anderson
(1936) based the reconstruction of the jaw musculature of the phytosaur
Machaeroprosopus on that of modern reptiles, particularly Alligator and
266 ANNALS OF THE SOUTH AFRICAN MUSEUM
Spenodon. Colbert (1946) also applied the myology of the head of living
crocodiles to that of the crocodilian Sebecus.
It is generally agreed that in Crocodilia the m. adductor mandibulae
separates into the external, posterior and internal portions as tabulated below,
though Edgeworth (1935) failed to identify a posterior (= medial of Edgeworth)
portion.
Jaw muscles of the Crocodilia based on the nomenclature of Lakjer (1926) and
Anderson (1936)
Innervated by the trigeminal
superficialis
Adductor mandibulae externus medialis
profundus
Adductor mandibulae posterior
pseudotemporalis
pterygoideus dorsalis
= pterygoideus D of Lakjer
Adductor mandibulae internus
pterygoideus ventralis
intramandibularis
Innervated by the facial
Depressor mandibulae
The m. adductor mandibulae externus was probably divisible into three
muscle sheets in Orthosuchus as in living crocodiles. In modern genera the
m. mandibulae externus superficialis arises along the outer edge of the quadrate
between the jaw articulation and the postorbital, and inserts on the dorsal
surface of the surangular. There is no reason to suppose that this muscle had a
different arrangement in Orthosuchus. However, this muscle presumably
inserted mainly on the horizontal flange of the surangular of Orthosuchus. It is
also likely that the m. adductor mandibulae externus medialis occupied a
similar position in Orthosuchus to that seen in Alligator. It probably arose partly
from the ventral surface of the postorbital and partly from the pterygoid face of
the quadrate deep to the superficial sheet of the m. adductor mandibulae
externus. The m. adductor mandibulae externus medialis would also have
inserted on the dorsal surface of the surangular underneath the superficial sheet.
Deeper again and running below the m. adductor mandibulae externus
medialis lies the m. adductor mandibulae externus profundus. This is the only
muscle to penetrate the superior temporal fenestra. Without doubt the major
origin of this muscle in Orthosuchus, as in Alligator, must have been from the
parietal, squamosal and quadrate within the superior temporal fossa, though
other slips may have attached more ventrally to the quadrate and postorbital.
Similarly the major insertion of this muscle in Orthosuchus must have been into
the membrane lying over the adductor fossa.
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 267
In Orthosuchus both the temporal fenestrae and the adductor fossa are very
large relative to the length of the skull. Indeed they are larger in Orthosuchus
than they are-in all the living crocodiles, including Gavialis. The large floored
area of the superior temporal fossa indicates that the m. adductor mandibulae
externus profundus had a much larger insertion area in Orthosuchus than it has
in living crocodiles. Further the large size of the inferior temporal fenestra and
adductor fossa suggest that this muscle required a larger area in which to bulge.
It therefore seems probable that the m. adductor mandibulae externus profundus
was larger and more powerfully developed in Orthosuchus than it is in living
forms.
Fig. 19. Orthosuchus stormbergi. Diagrammatic reconstruction of the inferred general location
of the jaw muscles.
Abbreviations: a: m. adductor mandibulae externus superficialis; b: m. adductor mandibulae
externus medialis; c: m. adductor mandibulae externus profundus; d: m. adductor mandibulae
posterior; e: m. pterygoideus ventralis; f: m. pterygoideus dorsalis; g: m. depressor
mandibulae.
The second part of the adductor, the m. adductor mandibulae posterior, no
doubt also occupied a similar position in Orthosuchus to that seen in living
crocodiles. It must have been a short, stout muscle which attached to the
pterygoid face of the quadrate, and then ran slightly obliquely down to the lower
jaw. In living crocodiles this muscle inserts on the lower surface of the angular
along the posterior part of the external mandibular fenestra, and on the inner
wall of the angular medial to this fenestra. Since the angular does not extend
medially in Orthosuchus as it does in living genera, it seems likely that this
muscle inserted in part on the prearticular.
In Alligator the m. adductor mandibulae posterior is divided at its insertion
by the m. pseudotemporalis. Probably a similar arrangement existed in Ortho-
suchus. This muscle would have originated on the postorbital.
The second part of the m. adductor mandibulae internus, the m. pterygoi-
deus, is extremely large in living crocodiles. There can be little doubt that in
268 ANNALS OF THE SOUTH AFRICAN MUSEUM
Orthosuchus it arose mainly from the posterior edge of the pterygoid flange. It
probably also attached to the ectopterygoid, and to the basisphenoid and
adjacent medial portion of the pterygoid, as it does in living crocodiles.
In eusuchians a ventral slip of the m. pterygoideus wraps round the
lower jaw and inserts on the retroarticular process, below and behind the
glenoid. Since this process is absent in Orthosuchus a comparable muscle must
have had a more anterior insertion principally in front of the glenoid. The
ventro-lateral surface of the posterior region of the angular of Orthosuchus
indicates an area of muscle attachment in life. This is suggested as the insertion
area of the m. pterygoideus ventralis. In this position the muscle would have
acted more or less at right angles to the lower jaw when this was widely open,
and would have been efficient in closing the jaws. The migration of the
m. pterygoideus ventralis on to the retroarticular process in later crocodiles has
reduced its efficiency in closing the jaw. However, as Ewer (1965) has pointed
out, a muscle in this position stabilizes the articulation.
In living crocodiles it is the powerfully developed anterior extension of the
pterygoideus, the m. pterygoideus dorsalis (= the pterygoideus D of Lakjer),
that serves to close the lower jaw. This muscle originates along the length of the
inner surface of the snout. Anteriorly it extends well in front of the orbit, and
attaches to the dorsal surface of the maxilla, palatine and pterygoid. It inserts on
the medial surface of the angular, below and in front of the glenoid.
In view of the large pterygoid flange present in Orthosuchus it is probable
that some development of the m. pterygoideus dorsalis had occurred. However
it could not have been as extensive in Orthosuchus as it is in eusuchians.
Probably it originated from the dorsal surfaces of both the pterygoid and
palatine, but it could not have extended any farther forward than the anterior
margin of the orbit in view of the limited development of the bony secondary
palate. Posteriorly it probably attached to the membrane over the very large
adductor fossa, and may have inserted on the prearticular.
The evidence suggests that the m. pterygoideus ventralis and the
m. pterygoideus dorsalis were not as well developed in Orthosuchus as they are
in living crocodiles. Nevertheless this muscle must have been considerable, as
shown by the large size of the pterygoid flange. Further, the positioning of this
muscle suggests that it was fully effective in closing the jaws. On the other hand,
the large size of the superior temporal fenestra clearly indicates that the
m. adductor mandibulae externus profundus was also extensive, as it is in
Gavialis, so increasing the efficiency of snapping shut the jaws.
Dollo (1884) related the development of this musculature to the size of the
antorbital fenestra. He concluded that where the m. adductor mandibulae
externus was more important than the m. pterygoideus dorsalis, the antorbital
fenestra was reduced. Conversely this fenestra was enlarged where the m. ptery-
goideus dorsalis was well developed. Gregory & Adams (1915) and later workers
supported this idea. Walker (1961) expressed the opinion that in Ornithosuchus
the m. pterygoideus dorsalis originated in large part from a membrane stretched
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 269
across the depressed area on the snout surrounding the antorbital fenestra,
and probably also from the depressed area itself.
Certainly the small size of the antorbital fenestra may be correlated to the
greater importance of the m. adductor mandibulae externus profundus in
Orthosuchus. On the other hand, Ewer (1965) has pointed out that in all later
crocodiles it is the m. pterygoideus dorsalis that is the more extensive of the two
muscles, yet the antorbital fenestra has been lost. This would not have occurred
had the fenestra been the main area of origin of the muscle. There seems little
doubt that in Orthosuchus the antorbital fenestra was unrelated to the jaw
musculature. It probably housed a gland in life.
According to Lakjer (1926) and Anderson (1936), the m. intramandibularis
is differentiated from the m. adductor mandibulae in living crocodiles. This
muscle arises from the membrane lying over the adductor fossa, and then passes
over the coronoid through the Meckelian fossa and along the primordial canal
to insert on Meckel’s cartilage. In Anderson’s view, since the m. adductor
mandibulae externus profundus and the m. pterygoideus dorsalis have slips
which attach to the m. intramandibularis, the anterior insertion of this muscle
increases the efficiency of the other muscles. It is possible that a similar muscle
was present in Orthosuchus, although presumably it passed over the prearticular
before entering the jaw.
In view of the insignificant development of the retroarticular process in
Orthosuchus, it seems probable that the m. depressor mandibulae was not
strongly developed. No doubt it had a similar arrangement to that seen in living
crocodiles, and arose on the occipital surface of the parietal, squamosal and
exoccipital, and inserted on the posterior margin of the articular. In living
crocodiles where the retroarticular process is prominent, the m. depressor
mandibulae is correspondingly strongly developed. Adams (1919) suggests that
since crocodiles generally lie with their heads on the ground, this muscle serves
to lift the cranium rather than lower the bottom jaw. It may be that this muscle
was less well developed in Orthosuchus, and that the skull did not reach the
massive size attained by many of the living species.
MUSCULATURE OF THE SHOULDER AND FORE-LIMB (Figs 20-21)
Fiirbringer (1876, 1900) described in detail the musculature of the
shoulder and upper arm of various reptiles, and this work was followed by
Von Wettstein (1937) in his account of the Crocodilia. Gregory & Camp (1918)
also reviewed the identification of the shoulder muscles. This account follows
the terms used by these workers.
Axial muscles associated with the pectoral girdle
The pectoral girdle of Orthosuchus differs from that of living crocodiles in
that while the scapula is long, relative to the size of the humerus, the coracoid is
shorter. Presumably, therefore, attachment areas of certain muscles would have
differed from those occurring in living crocodiles. On the other hand a clavicle
270 ANNALS OF THE SOUTH AFRICAN MUSEUM
is lacking in Orthosuchus, as it is in living crocodiles, and it seems likely that
specializations in the axial musculature which are associated with this bone’s
absence in modern genera had aiready taken place in Orthosuchus.
In Orthosuchus the m. capito-dorso-clavicularis (= trapezius of Fiirbringer)
probably arose from the parietal and squamosal, and merged posteriorly into
the m. latissimus dorsi, as it does in recent crocodiles. The m. sterno-mastoideus
consists of two parts in crocodiles. The muscle arises on the skull, but is divided
by the atlantal rib before it inserts on the sternum. Presumably this muscle was
similarly modified in Orthosuchus, since a well-developed atlantal rib is present.
Although it is not found in other living reptiles, the m. rhomboideus is
developed in the Crocodilia. It arises from the fascia above the eighth and ninth
cranial nerves, and inserts on the anterior two-thirds of the dorsal inner surface
of the supra-scapula. Since this muscle is also well developed in birds, it seems
probable that it was present in ancestral archosaurs. In Orthosuchus the large
size of the scapula suggests that the rhomboideus muscle may have inserted
principally on this element rather than on the supra-scapula. Similarly, it seems
probable that the deeper m. levator scapulae superficialis inserted on the upper
anterior external margin of the scapula in Orthosuchus. The surface of this
margin shows fine striations which possibly indicate muscle attachment areas.
The muscle presumably took origin on the cervical vertebrae.
In living crocodiles the m. serratus superficialis arises on the last cervical
and first three dorsal ribs, behind the pectoral girdle and below the m. latissimus
dorsi. There is fundamentally no difference between these ribs in Orthosuchus
and those of living crocodiles. However, whereas in living species the ‘uncinate’
processes are normally cartilaginous, in Orthosuchus they are fully ossified. This
muscle most likely inserted on the posterior margin of the scapula. Similarly, the
deep-lying m. serratus profundus presumably arose from the transverse process
of the cervical vertebrae, and inserted on the inner and upper surface of the
scapula. The m. omohyoid must also have inserted on to this surface of the
scapula.
Dorsal muscles of the fore-limb
The m. latissimus dorsi is weakly developed in modern crocodiles and
shows some differentiation into two parts. In Orthosuchus it would have taken
origin from the external surfaces of the dorsal ribs and inserted on the head of
the humerus near its posterior margin. Almost certainly the m. subcoraco-
scapularis (= subscapularis) inserted near to this muscle, and arose from the
rear edge of the scapula. Similarly, it is likely that the m. teres major was present
in Orthosuchus. According to Fiirbringer, although this muscle is absent in
Sphenodon, and occurs in only a few lacertilians (Agamidae), it is developed in
living crocodilians. In Orthosuchus the upper, backwardly projecting margin of
the scapula probably acted as the area of attachment for this muscle, which
extended beneath the m. latissimus dorsi to insert on the lateral surface of the
humerus.
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 271
t
'
4 U /
aw) /
i
Kig!
ToS
aot
1
F
1
=|
ay
~
Fig. 20. Orthosuchus stormbergi. Lateral surface
of the pectoral girdle showing inferred general
areas of muscle attachments.
Abbreviations: a: m. trapezius; b: m. rhom-
boideus; c: m. levator scapulae superficialis;
d: m. latissimus dorsi; e: m. serratus superfi-
cialis; f: m. subscapularis; g: m. teres major;
h: m. scapulo-humeralis posterior; i: m. dorsalis
scapulae and m._ deitoides_ clavicularis;
j: m. triceps; k: m. coracobrachialis brevis;
1: m. supracoracoideus; m: m. biceps brachii.
In living crocodiles the m. scapulo-humeralis anterior is lacking, though the
posterior division is well developed. A similar arrangement almost certainly
existed in Orthosuchus, since there is a well-marked depression ventrally on the
external surface of the scapula, near to its anterior border. A similar area on the
scapula of living crocodiles marks the origin of this muscle. The m. scapulo-
humeralis posterior (= scapulo-humeralis profundus of Fiirbringer) would have
inserted on the lateral surface of the humerus. In Orthosuchus this area of the
humerus is marked by fine striations.
272 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 21. Orthosuchus stormbergi. Diagrammatic reconstruction of the inferred main lines of
action of the muscles of the fore-limb. Abbreviations as in Figure 20.
The deltoid muscle in reptiles is typically a broad but thin sheet which
arises in two parts: one, the m. dorsalis scapulae (= deltoides scapularis
superior of Fiirbringer), arises from the upper part of the scapula, and the
second, the m. deltoides clavicularis (= cleido-humeralis), from the clavicle and
interclavicle. Since the clavicle has been lost in crocodiles, the m. deltoides
clavicularis has shifted its origin to the scapula. A similar arrangement must have
existed in Orthosuchus, and both parts of the deltoid would have inserted on the
outer surface of the deltopectoral crest. Fine striations indicate this area as one
of muscle attachment in life.
Despite the absence of an olecranon process in living crocodilians, the major
dorsal muscle of the arm, the m. triceps, still forms a massive muscle body. In
view of this, and since the proximal end of the ulna is expanded in crocodilian
fashion, there is no reason to suppose that the m. triceps was not well developed in
Orthosuchus. It is also likely that the areas of attachment of this muscle on to the
pectoral girdle and humerus were fragmented into five centres of origin, as they
are in living crocodiles. In these forms the muscle consists of two layers, the
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 273
deeper of which arises from the upper, posterior and medial surfaces of the
humerus, and the outer having its origins on the pectoral girdle. In Orthosuchus
the scapula is grooved above the glenoid facet, and very probably this area
denotes in part the origin of the m. triceps. The insertion of this muscle must have
been by a common tendon on to the head of the ulna.
Ventral muscles of the fore-limb
The m. pectoralis in Orthosuchus no doubt formed a well-developed super-
ficial muscle layer, originating on the interclavicle and ribs and inserting on the
apex of the deltopectoral crest.
In living crocodiles the longus division of the m. coracobrachialis is absent
and a similar arrangement was probably present in Orthosuchus, since distally
the humerus resembles that of extant forms. Probably the m. coracobrachialis
brevis would have arisen on the postero-ventral margin of the coracoid. How-
ever, this area of attachment must have been relatively smaller than that found
in living crocodiles, since distally the coracoid is neither as elongate nor as
expanded in Orthosuchus as it is in the eusuchians. This muscle would have
inserted in the concavity antero-medial to the deltopectoral crest, as it does in
living species. Again, its area of insertion must have been more limited in
Orthosuchus than in modern crocodiles, judging by the more robust nature of
the deltopectoral crest.
According to Von Wettstein, the m. supracoracoideus in living crocodiles
has partially shifted its origin on to the inner face of the coracoid and lower edge
of the scapula because of the considerable development of the m. coraco-
brachialis. It is possible that in Orthosuchus, where the m. coracobrachialis was
more feebly developed, the area of origin of the m. supracoracoideus was
confined to an area above that of the m. coracobrachialis. The m. supra-
coracoideus would then have inserted on the deltopectoral crest lateral to the
point of attachment for the m. pectoralis.
The outer surface of the coracoid of Orthosuchus must also have given
origin to the m. biceps brachii, which would have inserted on to the proximal
part of the radius and ulna. The m. brachialis inferior was probably well
developed, and in life would have attached to the anterior margin of the
humerus, and inserted on the proximal end of the ulna.
Muscles of the lower arm
The following account is based on the work of Von Wettstein, who followed
Ribbing (1907), and of Haines (1939), who made a comparative study of the
extensor muscles of the forearm.
In view of the marked similarity between the lower arm elements of
Orthosuchus and that of living crocodiles, there seems every reason to suppose
that the musculature of this region in Orthosuchus would have resembled that of
eusuchians. In this group in general there is a reduction in the lower arm and
hand musculature.
274 ANNALS OF THE SOUTH AFRICAN MUSEUM
Typically, the long extensors of the forearm attach to the ectepicondyle of
the humerus. This projection is only slightly more developed in Orthosuchus
than it is in living crocodiles, where it is minimal. Very likely, therefore, the
m. extensor digitorum communis (= humerodorsalis of Haines) was as reduced
in Orthosuchus as it is in modern forms. In living crocodiles this muscle no
longer inserts on to all the digits, but chiefly on to metacarpal two, with
extensions to four and the radiale.
There can be little doubt that in Orthosuchus the m. supinator (= extensor
antibrachii radialis of Ribbing) also arose from the ectepicondyle in two parts,
aS in both living crocodiles and lizards, with the m. extensores carpi radiales
attaching to the radius. Similarly, the m. anconeus (= extensor antibrachii
ulnaris of Ribbing) was probably well developed and also took origin on the
ectepicondyle. On the other hand, the m. extensor carpi ulnaris may well have
been somewhat limited in extent. In living crocodiles this muscle is confined to
the ulna, and does not extend to the pisiform.
The m. abductor pollicis longus (= abductor digiti 1 of Ribbing =
supinator manus of Haines) was no doubt extensively developed in Orthosuchus.
It probably originated on the radius as well as on the ulna, and inserted on to
the radiale as it does in living crocodiles. The m. extensores digitorum breves
would have extended on to the digits. In Haines’s view these specializations in
musculature are associated with a great mobility of the wrist joint and the use
of the elongated carpus as an extra limb segment.
Typically, the flexor muscles of the lower arm are more powerful than ‘he
extensors, since the main propulsive effort is a backward push of the distal part
of the limb. The long flexors arise from the entepicondyle of the humerus which,
although of limited development in crocodiles, is a little more pronounced and
rugose in Orthosuchus than it is in later forms.
In living crocodiles, the m. flexor digitorum profundus (= flexor accessorius
of Ribbing) is well developed, while the m. flexor palmaris superficialis (= flexor
primordialis communis of Ribbing) is less extensive. This latter muscle is united
with the m. flexores breves superficiales. Possibly a similar arrangement existed
in Orthosuchus. Certainly in life this medial muscle mass would have inserted on
a palmar aponeurosis and effected flexion of the toes.
In living genera the m. pronator teres (= flexor antibrachii radialis of
Ribbing) is joined together with the m. supinator, but neither the m. flexor carpi
radialis nor the m. epitrochleoanconeus (= antibrachii ulnaris of Ribbing) is
present. Whether or not reduction of musculature had proceeded this far in
Orthosuchus is difficult to determine. These muscles are present in lizards and
Sphenodon, but in these forms the entepicondylar process is well developed. On
balance, it seems likely that in Orthosuchus these muscles were at best only
feebly developed.
Deep in the forearm, the m. pronator profundus, connecting radius and
ulna, probably had a similar arrangement to that seen in living crocodiles,
passing between the m. flexor digitorum profundus and the m. pronator teres.
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 275
As in recent genera, the m. abductor digiti V was most likely well developed,
taking origin on the palmar aponeurosis and inserting on the melararpals and
phalanges of the fifth digit.
MUSCULATURE OF THE PELVIS AND HIND-LIMB (Figs 22-24)
The probable arrangement of the muscles of the pelvis and upper leg of
Orthosuchus is based on that found in Alligator (Romer 1923).
Axial muscles associated with the pelvis
The dorsal axial muscles in Orthosuchus would have run antero-posteriorly
above the lumbar and caudal transverse processes and attached to the dorsal
surfaces of the sacral vertebrae and internal surfaces of the ilium.
Similarly, the ventral axial musculature must have consisted of a series of
muscle sheets covering the flanks of the body. In Alligator the three lateral
members of this series all take origin from the lumbo-dorsal fascia. This fascia
arises from the surface of the dorsal musculature and from the tips of the
transverse processes of the lumbar vertebrae, and attaches posteriorly to the
anterior edge of the blade of the ilium. In Orthosuchus this area of the ilium is
extended forward to form a promjnent anterior process, and may have provided
a stronger base of attachment for the fascia.
Presumably these lateral muscles, the m. obliquus abdominis externus, the
m. obliquus abdominis internus and the m. transversus abdominis, had a similar
arrangement in Orthosuchus. The m. obliquus abdominis externus inserts on the
anterior margin of the acetabulum, the last abdominal rib and on an aponeurosis
over the main part of the m. rectus abdominis. The m. obliquus abdominis
internus inserts on the posterior ribs and gastralia, and the m. transversus
abdominis inserts on the m. rectus abdominis. The m. rectus abdominis no
doubt ran posteriorly in the ventral midline to attach principally to the gastralia
and abdominal ribs, though a part of it may have inserted on the posterior edge
of the pubis and on the m. ilio-ischio-caudalis, as it does in Alligator.
In Orthosuchus the posterior process of the iliac blade extends well back.
Similarly, the postero-ventral edge of the ischium is produced far posteriorly.
This indicates increased attachment areas for the m. ilio-ischio-caudalis. In life
this muscle would have occupied the ventral half of the tail on either side between
the transverse processes and the midline ventrally. It is, therefore, fairly certain
that the tail was a highly muscular organ.
Troxell (1925) has pointed out that the decrease in size of the vertebrae in
both directions from the pelvis in living crocodilians is of advantage to the
animal in swimming, where the tail is the propelling organ. A similar change in
size of the vertebrae occurs in Orthosuchus. Presumably the tail could act as a
propulsive organ in swimming, and no doubt also assisted in movement over
land. However, the nature of the articular surfaces of the centra indicates that
the degree of angular movement between successive vertebrae could not have
been as great as that in living crocodiles, where the vertebrae are procoelous.
276 ANNALS OF THE SOUTH AFRICAN MUSEUM
Dorsal muscles of the hind-limb
Above the acetabulum, and along the upper part of the posterior process,
the ilium bears a number of distinct striae which suggest an area of muscle
attachment. The most dorsal of these would probably have been the m. ilio-
tibialis. This constitutes one part of the major dorsal muscle of the thigh, the
m. quadriceps femoris. In Alligator, the m. ilio-tibialis has three distinct heads,
but it is impossible to determine whether a similar arrangement existed in
Orthosuchus, or whether its origin was still undivided.
ee” =
? os ooo ee Sooo ee Ors
Ld e865 es Se - ® i
o o SS = CY ry Py
of? er aN g a i r) “ib
ty t 2.7 t Soa
Fig. 22. Orthosuchus stormbergi. Lateral surface of the pelvic girdle showing inferred general
areas of muscle attachments.
Abbreviations: a: m. rectus abdominis; b. m. ilio-ischio-caudalis; c: m. ilio-tibialis;
d:m. ambiens; e: m. femoro-tibialis; f: m. ilio-fibularis; g: m. ilio-femoralis; h: m. ischio-
trochantericus; i: m. pubo-ischio-femoralis-internus; j: m. pubo-ischio-tibialis; k: m. flexor,
tibialis internus parts 1 and 2; 1: m. flexor tibialis externus; m: m. flexor tibialis internus
part 3; n: m. adductor femoris; 0: m. pubo-ischio-femoralis externus; p: m. caudi-femoralis
(= coccygeo-femoralis brevis); q: m. caudi-femoralis (= coccygeo-femoralis longus).
The m. ambiens in Alligator has two areas of origin. The smaller part arises
on the proximal medial surface of the pubis, but the major origin is on the
junction of the external surface of the pubis and the cartilage which lies in front
of the acetabulum. The upper edge of the proximal end of the pubis of Ortho-
suchus is marked by a shallow depression. Very probably this denotes the area
of origin of the major part of the m. ambiens. Since the proximal end of the
pubis lies in a more dorsal position, the m. ambiens must have had a higher area
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 277
of origin in Orthosuchus than it does in living crocodilians. In Alligator both
elements of the m. ambiens unite with the m. ilio-tibialis. In addition, the larger
element forms a tendon which passes through the extensor tendon of the
m. ilio-tibialis across the knee to the lateral surface of the leg, where it joins the
external head of the m. gastrocnemius. A similar tendon had probably been
developed in Orthosuchus.
Fig. 23. Orthosuchus stormbergi. Diagrammatic reconstruction of the inferred main lines of
action of the dorsal muscles of the thigh. Abbreviations as in Figure 22.
The anterior (= dorsal of Romer) and medial (= anterior of Romer)
surfaces of the greater part of the shaft of the femur no doubt provided an
attachment area for the origin of the m. femoro-tibialis, the third component of
the m. quadriceps femoris. In Alligator this muscle has a second, more lateral
(= posterior of Romer) origin, but it is impossible to determine whether this
was also the case in Orthosuchus. Its insertion would have been on to the weakly
278 ANNALS OF THE SOUTH AFRICAN MUSEUM
developed cnemial crest of the tibia. Nevertheless it seems that in Orthosuchus,
as in living crocodiles, the m. quadriceps femoris was a powerful extensor of the
thigh. Presumably the m. ilio-fibularis, a smaller extensor, ran parallel to it from
just below the m. ilio-tibialis, and inserted into the head of the fibula. It may also
have connected to the external head of the m. gastrocnemius as it does in
Alligator.
The blade of the ilium above the acetabulum is deeper in Orthosuchus than
it is in Alligator. This expansion no doubt reflects an increased area of origin for
the m. ilio-femoralis, which arises below the m. ilio-tibialis. In life this muscle
would have inserted on the lateral border of the femur for the greater part of the
length of the shaft, and would have been a very powerful abductor.
In Alligator a small muscle, the m. ischio-trochantericus, runs from the
posterior part of the inner surface of the ischium and inserts at the outer anterior
edge of the femur, near to its head. There is no reason to suppose that a similar
muscle did not exist in Orthosuchus.
The m. pubo-ischio-femoralis internus primitively originates on the medial
surface of the pubis. However, Romer (1923) has shown that in Alligator this
muscle is present in two parts and has more dorsal origins. The similarity of the
pelvis in Orthosuchus suggests that this change had already occurred. One part
of the muscle probably originated from the ventral surfaces of the posterior
dorsal vertebrae, and possibly also attached to the inner surface of the anterior
process of the blade of the ilium. This muscle would have inserted on the.
anterior surface of the proximal part of the femur. The evidence of strong muscle
attachment in this area is shown by fine striations marking the bone at this point.
The second part of the m. pubo-ischio-femoralis internus probably originated
from the internal surfaces of the ventral margin of the ilium and dorsal margin of
the ischium. It may also have attached to the ventral portions of the sacral ribs,
as it does in Alligator. This muscle would then have extended anteriorly to insert
on the femur, postero-medially to the insertion of its counterpart. A shallow
depression on the femur of Orthosuchus at this point supports this conclusion.
In life the m. pubo-ischio-femoralis internus would have been a powerful muscle
drawing the femur inward, upward and forward.
Ventral muscles of the hind-limb
In view of the marked development of the dorsal musculature in Ortho-
suchus, it is clear that a corresponding development in ventral musculature must
have existed, as it does in Alligator. In the latter, the superficial layer of muscles
flexing the knee consists of six muscles which unite into two groups at their
insertion on the tibia.
Romer has demonstrated that in Alligator the external group of muscles is
composed of the m. pubo-ischio-tibialis and two parts of the m. flexor tibialis
internus. In lizards the m. pubo-ischio-tibialis arises along the entire ventral
margin of the girdle, but in crocodiles it is confined to a small area on the lower
margin of the anterior process of the proximal ischium. Since this process is more
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 279
Fig. 24. Orthosuchus stormbergi. Diagrammatic reconstruction of the inferred main lines of
action of the ventral muscles of the thigh. Abbreviations as in Figure 22.
strongly developed in Orthosuchus than it is in living crocodiles, it may be that
the muscle too was more extensive. The m. flexor tibialis internus probably
originated both on the medial surface of the ischium at its postero-ventral angle
and from the posterior angle of the iliac blade. As in Alligator, these muscles
would have inserted by a common tendon on to the medial surface of the tibia,
internal to other flexors of the lower leg.
The second, internal group of muscles in Alligator consists of the m. flexor
tibialis externus and two further parts of the m. flexor tibialis internus. In
Orthosuchus the m. flexor tibialis externus almost certainly arose behind the
m. ilio-tibialis on the posterior postacetabular iliac process. The m. flexor
280 ANNALS OF THE SOUTH AFRICAN MUSEUM
tibialis internus commonly arises from the ilio-ischiadic ligament, and pre-
sumably also did so in Orthosuchus. However, there may have been a second
point of origin from the posterior margin of the proximal ischium, as occurs in
Alligator. In both Orthosuchus and Alligator the ischium forms a marked
prominence at this point. The m. flexor tibialis externus, together with the
m. flexor tibialis internus, would have inserted medially on the head of the tibia.
In addition, a second tendon may have passed down to unite with the m. gastro-
cnemius near the foot, as it does in living crocodiles.
Since the m. pubo-tibialis has been lost in both living crocodiles and birds,
it seems reasonable to assume that this muscle was also absent in Orthosuchus.
A m. adductor femoris probably arose both below the m. flexor tibialis
internus on the posterior margin of the ischium, and also from the antero-
ventral angie of the blade of the ischium. This muscle would have inserted on the
posterior (= ventral of Romer) surface of the femur.
The deepest muscle of the ventral musculature, the m. pubo-ischio-
femoralis externus, is a powerful muscle which no doubt had much the same
arrangement in Orthosuchus as it has in Alligator. In the latter, one part arises
from the main area of the blade of the ischium between the two adductors, a
second from the ventro-lateral surface of the pubis, and a third from the dorso-
medial surface of the pubis and adjacent gastralia. These parts unite and insert
on the postero-lateral margin of the femur close to its head. The femur of
Orthosuchus bears a marked rugosity at this point, and there can be little doubt.
that this represents the insertion area of this muscle.
Muscles connecting the femur to the tail, which made possible a backward
and downward pull on the femur, were also well developed in Orthosuchus. The
m. coccygeo-femoralis brevis (= caudifemoralis of Gadow) almost certainly
arose from the last sacral and first caudal vertebrae, and attached to the ventral
surface of the posterior process of the ilium. In Orthosuchus this process is
stouter than it is in Alligator, and possibly the iliac slip of the muscle was more
important than the caudal. This muscle would have inserted on the femur above
the fourth trochanter.
Since the fourth trochanter is relatively less well developed in Orthosuchus
than it is in Alligator, a weaker m. coccygeo-femoralis longus (= caudi
femoralis of Gadow) than that of recent types is indicated. This muscle would
have originated on the ventral surfaces of the caudal vertebrae and inserted both
on the fourth trochanter and on to the fibula.
The muscles of the lower hind-limb
A comprehensive account of the musculature of the lower hind-limb of
living crocodiles was given by Von Wettstein (1937), who followed the work of
Gadow (1882).
Since the form of the lower hind-limb of Orthosuchus is closely comparable
to that of living crocodiles, there seems little doubt that its musculature also
followed a similar pattern. The common extensor of the digits, the m. extensor
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 281
digitorum communis, would have originated on the lateral femoral condyle and
inserted on to the bases of the metatarsals. In Alligator this muscle unites with
the m. tibialis anterior, which arises on the proximal part of the tibia, and there
is a common insertion on to the four long metatarsals. This differs from the
pattern found in lizards, where the two muscles remain separate. Presumably in
Orthosuchus these muscles had an arrangement similar to that found in
Alligator.
In crocodiles, unlike Sphenodon, the m. extensor peroneus brevis (anterior)
and the m. extensor peroneus longus (posterior) are separate. The m. peroneus
brevis arises on the fibula and inserts on the dorsal and lateral surfaces of
metatarsal five, and on the dorso-lateral surface of the calcaneum. The
calcaneum of Orthosuchus shows a shallow depression at this point, and the fifth
metatarsal bears a rugosity. It therefore seems likely that in Orthosuchus the
m. peroneus brevis had a similar relationship and, as in living genera, a compar-
able function of dorsiflexing the foot and elevating its lateral border. In
Alligator the m. peroneus longus arises on the lateral condyle of the femur and
inserts on to the calcaneal tuber, so that it functions to flex the lower limb.
Since the tuber is strongly developed in Orthosuchus, there can be little doubt
that this muscle had an arrangement and function comparable to that in living
crocodiles.
Similarly, the m. abductor and extensor hallucis was probably as well
developed in Orthosuchus as it is in Alligator. It would have originated on the
distal end of the fibula and from the calcaneum, and then crossed the dorsal
surface of the foot to insert on the first metatarsal.
The main flexor of the lower limb, the m. gastrocnemius, is superficial in
position and has two areas of origin. In Orthosuchus one part would have
attached to the lateral condyle of the femur, and the second to the tibia, as in
Alligator. Further, since the calcaneum of Orthosuchus is strikingly similar to
that of living crocodiles, there can be little doubt that this muscle had a similar
insertion. One part of the m. gastrocnemius would have run from the femur to
the calcaneal tuber, partly inserting on its upper surface and forming a stout
tendon which ran down the vertical groove to insert on the plantar aponeurosis.
The second part of the m. gastrocnemius must have inserted entirely on the
tuber. In life this muscle effected flexion of the foot during the propulsive effort.
The arrangement of the m. flexor digitorum longus (= flexor primordialis
communis of Von Wettsiein) in crocodiles is similar to that found in other
reptiles. In Orthosuchus it very probably had several heads of origin, principally
from the lateral condyle of the femur and proximal fibula. These would have
united and run medially to the calcaneal tuber to the plantar aponeurosis,
finally forming tendons inserting on the distal phalanges.
In living crocodiles the m. popliteus (= tibialis posticus of Gadow) is a
powerful muscle which arises from the fibula and tibia, and distally forms a
tendon which passes over a groove on the astragalus to insert on to the bases of
metatarsals one and two. There is no reason to suppose that a similar arrange-
282 ANNALS OF THE SOUTH AFRICAN MUSEUM
ment did not exist in Orthosuchus. Presumably, too, the m. pronator profundus
arose on the proximal part of the tibia and inserted on to the fibula. In living
crocodiles, according to Von Wettstein, the distal part of this muscle, the
m. interosseus, is absent.
AGE VARIATIONS IN THE SKULLS OF RECENT CROCODILIA AND
AN ANALYSIS OF THE DIFFERENCES BETWEEN THE SKULLS
OF THE TYPE, K409, AND K4639
The overall skull plan in K4639 compares well with that seen in the type,
K409. The skull elements of the smaller specimen also have the same relation-
ship to one another that they have in the type. On the other hand, certain
differences are also apparent. Skull dimensions are dissimilar, and in K4639 the
superior temporal fenestra is oval in shape and smaller than the orbit, whereas
in K409 it is more or less circular and approximates the size of the orbit.
In view of the small size and low degree of ossification of the skull elements,
there can be little doubt that K4639 is a juvenile form. The lack of fusion
between the parietals and lack of a squamosal ridge in this specimen could also
be attributed to the juvenile condition. In K409 the parietals are fused, and there
is a ridge marking the position held by ear flaps in life.
The question then is whether the differences in skull proportions can be
attributed to a difference in age between individuals of a single species, or
whether they are specific differences. It cannot, unfortunately, be assumed that
K409 is of breeding size, though clearly it would be helpful if this point could be
established.
With this in mind, some attempt to analyse the differences, and to compare
these with similar dimensions in the protosuchian, Protosuchus, and in
eusuchians, was undertaken. Mook (192la) studied a series of skulls of
Crocodylus acutus (= Crocodilus americanus), Alligator mississippiensis and
Caiman crocodilus (= C. sclerops), and noticed a number of characters which
could be attributed to differences in age. To supplement this a series of nine
skulls of Crocodylus porosus, from the collection at the British Museum (Natural
History), have been examined by the author.
Breadth of skulls compared with length (Fig. 25)
Mook concluded from his data that in Crocodylus acutus and Caiman
crocodilus there is a marked, though irregular, broadening of the skull with age.
In Alligator mississippiensis there is only a slight change from young to old, a
slight narrowing being noticeable in older specimens.
When breadth (ordinates) and length of skulls (abscissae) are plotted
against one another a somewhat curved line is obtained. If both are converted
into log. form, the lines are still curved, but if log. (length) is plotted against
log. (breadth—10) the lines become, for practical purposes, straight.
Regression coefficients (b) for logy )(breadth-10) on log,o(length) were
283
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN
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284 ANNALS OF THE SOUTH AFRICAN MUSEUM
calculated separately for the four living species, and the following values
obtained:
Caiman crocodilus b= 1235
Alligator mississippiensis b = 1,236
Crocodylus acutus == 2a
Crocodylus porosus b == 1,332
The graph shows that the last three of these form a rather close group, with
Caiman standing well outside. In the close group, the regression coefficients of
Crocodylus porosus and of Alligator are the most distinct, but a ¢ test shows that
the difference between these coefficients is not significant (t = 1,950 with 14 d_f.).
While the position of the line for Alligator is perhaps marginally above the
other two, it seems convenient for present purposes to combine the points for
these three species and calculate one regression line. This is given by:
y = 1,236x—0,9207.
From this it can be shown that the ratio of breadth/length of skull has a
minimum value (0,45) for animals of skull length 115 mm. In smaller and in
larger animals the breadth of the skull is relatively greater.
Although the slope of the regression line for Caiman crocodilus (b = 1,235)
is not distinct from the slope of the line (b = 1,236) for the other three combined,
the regression line for Caiman lies clearly separated above that for the other
three groups (t = 11,265 with 34 d.f.). Thus Caiman crocodilus is demonstrated
as a species with a relatively broader skull.
Compared with this, there is no doubt that Orthosuchus (K409 and K 4639)
and Protosuchus have broader skulls than living species. The values for K409 and
K 4639 fall together reasonably well, and the value for Protosuchus is in close
conformity with these.
Relative size of the cranial table (Figs 26-27)
Mook (1921a) established that the cranial table is relatively broader in
young individuals than in older ones. He related the cranial table to the breadth
of the skull across the quadratojugals. The ratios of breadth of cranial table/
breadth of skull in Orthosuchus are:
K409 = 0,78
K4639 = 0,87
These ratios compare favourably with those Mook obtained for Alligator
and suggest that a similar growth pattern occurred in Orthosuchus.
It can also be seen from the data that within a species the relationship
approximates to a direct proportionality between breadth of cranial table and
length of skull.
The estimated ratios (from the graphs) are approximately:
Crocodylus porosus O26
Crocodylus acutus = 027 Gg)
Alligator mississippiensis = 0,28 (Fig. 27)
Caiman crocodilus == 0/35
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 285
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Crocodylus porosus x
©
Orthosuchus stormbergi oO
0
100 200 300 400 500 600 700
Length of skull (mm)
Fig. 26. Graph showing relationship between breadth of cranial table and length of skull in
Crocodylus acutus, C. porosus and Orthosuchus stormbergi.
286 ANNALS OF THE SOUTH AFRICAN MUSEUM
e
co
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0
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Fig. 27. Graph showing relationship between breadth of cranial table and length of skull in
Alligator mississippiensis, Caiman crocodilus and Orthosuchus stormbergi.
Against these Orthosuchus shows a much higher ratio: 0,62 for both K409 and
K4639.
These figures clearly separate Orthosuchus from the eusuchians and also
show a close relationship between K409 and K4639.
Relation of preorbital to postorbital length (Fig. 28)
Mook’s work on modern species showed a progressive increase in the
length of the facial region as compared with the cranial region during growth.
The degree to which this occurs depends on the form of the adult.
The ratios of postorbital over preorbital length for Orthosuchus are:
K409 = 0,96
K4639 = 1,16
These values compare favourably with those Mook obtained for brevirostrate
forms, in particular that of Caiman. On the other hand Protosuchus is shown
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 287
Preorbital length
150
100 Postorbital length
Orthosuchus K409
Protosuchus
oO
Caiman crocodilus
Preorbital length +
Postorbital length ©
Orthosuchus K4639
50
Protosuchia
Preorbital length
Postorbital length
O *
50 100 150 200 250 300
Length of skull (mm)
Fig. 28. Graph showing relationship between preorbital and postorbital length in Caiman
crocodilus, Orthosuchus and Protosuchus.
to have a surprisingly short preorbital region, even assuming that Colbert &
Mook (1951) made no allowance for the damaged tip. Protosuchus could
perhaps be regarded as a juvenile of a species very much larger than Orthosuchus.
However, both types seem to be young animals.
Relative size of the orbits (Fig. 29)
Mook noticed that the orbits of young crocodiiians are relatively much
larger than in full-grown animals, and expressed this as a ratio of length of
orbit/length of skull. Further, he pointed out that there is a progressive increase
in the relative breadth of the interorbital plate.
When the values for Orthosuchus K409 and K4639 are seen against those
for Crocodylus acutus they again fall together reasonably well. In other words,
the relatively longer orbits and the narrower interorbital plate of K4639, as
compared with K409, could be attributed to a size (i.e. age) difference.
Although the values for Orthosuchus correspond closely to those for
Crocodylus acutus, compared to both Alligator and Caiman the orbits of
Orthosuchus are short relative to skull length. On the other hand, the interorbital
region is broader in Orthosuchus than it is in any of these living species of a
comparable size.
Relative proportions and position of the superior temporal fenestrae
Mook suggested that as a general rule, in the very young stages the
superior temporal fenestrae are small and slit-like, that in later stages they
ANNALS OF THE SOUTH AFRICAN MUSEUM
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THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 289
become rounder, and in old animals they usually become small and nearly
circular. That the size and shape of the fenestrae are definitely size (i.e. age)
related characters for the majority of species seems unquestionable. However,
the precise pattern of development varies from species to species.
Mook notes that in young specimens of Caiman crocodilus the fenestrae are
relatively large, and that in older ones they are smaller and may even eventually
close at the surface. Skulls of Crocodylus porosus (British Museum (Natural
History)) indicate that in this species the fenestrae are, in the early stages,
elongate in the longitudinal direction, but when the skull reaches 300 to 400 mm
in length they become round, only to elongate in the same longitudinal direction
in larger specimens (Table 3).
To some extent the very elongate shape of the superior temporal fenestrae
in K4639 may be due to lateral compression of the skull during preservation.
However, even without this consideration it is still possible that this is a juvenile
character in Orthosuchus, the fenestrae becoming broader and assuming a more
or less circular shape in later stages. It is, of course, impossible to say whether
they would have remained large, as they do in Gavialis, or whether they would
eventually have become smaller, relative to skull size, as in other living crocodiles.
Mook also noticed that in crocodiles other than Caiman the superior
temporal fenestrae appear to migrate inwards during growth, due to failure of
growth of the intertemporal region. Hence in young specimens the centres of the
superior temporal fenestrae are immediately posterior to the centres of the
orbits, while in older individuals the centres of the fenestrae are posterior to the
inner part of the orbits. Again, the series of skulls of Crocodylus porosus suggests
that growth between the superior temporal fenestrae is minimal compared with
increase in length and breadth of the skull, in the early stages. However, there is
some evidence of a reversal of this position in skulls over 450 to 500 mm in
length.
In Orthosuchus, despite the fact that K409 is almost twice as long as K4639,
the width of the intertemporal region is the same for the two skulls (i.e. 6 mm).
On balance therefore, there seem to be no strong reasons indicating the
separation of K409 and K4639. In view of the similarity of skull plan and
relationship of individual elements, it seems more likely that the differences are
attributable to size (i.e. age) variation. Whether K409 is juvenile, or of breeding
size, is indeterminable. Much of the evidence seems to indicate that it is a young
animal.
MODE OF LIFE
Orthosuchus was rather lizard-like in body proportions. The type specimen
measures about two-thirds of a metre in length, and although this may well
have been a young animal, other known Triassic protosuchians are all more or
less the same size. The slender jaws and feeble dentition are clearly not those of
a voracious predator, and on land it would have had a number of formidable
competitors.
ANNALS OF THE SOUTH AFRICAN MUSEUM
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THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 291
The Triassic was a time of dramatic faunal changes, with the earlier types,
which were predominantly therapsids, being gradually replaced, mainly by
archosaurs. Unquestionably, the dominant forms of the Upper Triassic lay
among the saurischian dinosaurs. These were prosauropods of generally large
size, and included both herbivorous forms like Thecodontosaurus and Plateo-
saurus, and carnivores like Melanorosaurus. It is now quite clear that the earliest
known ornithischians also occurred at this time, as shown by Fabrosaurus,
Heterodontosaurus and Lycorhinus. Thecodontians also occur in the earlier
Triassic of South Africa, and Sphenosuchus is known from the Red Beds.
Although the mammai-like reptiles were reduced to a few genera, advanced
ictidosaurs, Tritylodon in particular, are characteristic of the Upper Triassic. The
earliest known mammal, Erythrotherium, is also known from these beds of
South Africa. Elsewhere in the late Triassic procolophonids are present,
labyrinthodont amphibians are found in Australia, and rhynchosaurs in Nova
Scotia, Scotland and India.
At the end of the Triassic the large pseudosuchians and many of the
prosauropods disappeared. However, the crocodiles flourished through the
Jurassic, at least in a chiefly aquatic environment, as evidenced by large
teleosaurs like Mystriosaurus and Steneosaurus, and highly specialized marine
forms like Metriorhynchus.
Coupled with the change in fauna there was, at least in South Africa
(Haughton 1924), a progressive increase in aridity from the Lower through to
the Upper Triassic. Yet, however tempting it may be to suppose that Orthosuchus
was essentially terrestrial in habit, there are undeniable indications that it spent
much of its time in water, as do living crocodiles.
This is most clearly shown by the form of the palate. Although Orthosuchus
has only a short secondary palate, the form of the vomers and palatines and
ridging on the pterygoids leave little doubt that a functional secondary palate
extended well back to the base of the skull, as in living crocodiles. The palate of
Orthosuchus differs only in that it was floored by soft tissue rather than by bone.
Hence in Orthosuchus the functional choanae opened well back, and conceivably
a valve apparatus was present which sealed off the glottis when the mouth was
open. Orthosuchus was probably able to remain submerged, leaving only the
external nares open above waiter to act as a snorkling device.
Ear flaps may also have evolved to prevent flooding of the otic region.
Shute & Bellairs (1955) have shown that modern crocodiles close the ear-flaps
when they submerge, though this action does not necessarily prevent water trom
entering the recess. It is difficult to see why ear flaps would have been developed
for a terrestrial habit, when their closure in modern species reduces conduction
of aerial sound to the ear by 10 to 12 db over most of a frequency range of
100 to 6 000 cycles (Wever & Vernon 1957).
Amongst living crocodiles the overall skull shape of Orthosuchus is paralleled
most closely by that of the Indian gharial, Gavialis gangeticus. In both forms the
snout is very slender and set off sharply from the skull table, though in Gavialis
292 ANNALS OF THE SOUTH AFRICAN MUSEUM
the snout is very much more elongate than it is in Orthosuchus. Gavialis feeds
primarily on small fish. An individual will creep towards a shoal of fish, and
then remain motionless whilst snapping sideways at the prey. The skull shape of
Orthosuchus is possibly an adaptation for this kind of aquatic predation.
Most probably Orthosuchus spent much of its time in lakes or swamps, and
this mode of life would have afforded certain advantages. Food in the form of
small fishes and aquatic invertebrates would have been pientiful and more
easily obtainable than a comparable diet on land, while the water would have
provided an admirable escape route from terrestrial predators.
Ewer (1965) came to the conclusion that the antorbitai fenestra in
Euparkeria housed a gland, suggesting that this might have been a salt gland.
The nature of the fenestra in Orthosuchus also strongly indicates that in life it
housed a gland, whose secretion would have passed through a duct occupying
the groove leading to the lower orbital margin.
However, whereas thecodontians like Euparkeria typically have a large
antorbital fenestra, it decreases in relative size through Orthosuchus to early
aquatic mesosuchians like Teleosaurus, is nearly always absent in highly
specialized marine forms like Metriorhynchus and Geosaurus and is lost in
living crocodiles. Presumably therefore the gland was more important in
terrestrial than in marine forms.
In contrast, a salt-secreting gland is predominantly important in marine
animals. Salt secretion by marine turtles is particularly well known, and occurs’
from one of the orbital glands. Among other reptiles salt secretion is known to
occur from the nasal gland in lizards. Although some terrestrial forms, such as
the tropical lizard, Iguana iguana, and the American desert lizard, Dipsosaurus
dorsalis, secrete significant quantities of salt, they do not do so in the large
amounts found in their marine relatives (Schmidt-Nielsen 1963). Birds also
possess a salt-secreting nasal gland, which is normally located in the orbit.
Again, in terrestrial forms it is very small, but is large in marine species. Indeed,
to some extent its size varies with exposure to salt loads.
Unfortunately, almost no work has been carried out in this field on
crocodiles. Schmidt-Nielsen (1960) found that a single specimen of the estuarine
crocodile, Crocodylus porosus, did not respond to osmotic loads. This is the
most thoroughly aquatic and frequently marine species of all living crocodiles.
In living crocodiles a well-developed nasal gland lies anterior to the
preconcha, with its duct running anteriorly. Both a lachrymal and a large
Harderian gland lie within the orbit. The antorbital gland of Orthosuchus was
possibly an additional orbital gland, and the patches of glandular tissue which
occur on the conjunctiva of the lower lid in Crocodylus porosus may represent
the vestige of this gland.
Taking fore- and hind-limb lengths as humerus + radius + radiale, and
femur + tibia respectively, the length of the fore-limb approximates that of the
hind-limb in both Orthosuchus and contemporary crocodilians (table 4). Two
large specimens of Crocodylus niloticus showed a fore-limb 83% and 88% the
293
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN
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294 ANNALS OF THE SOUTH AFRICAN MUSEUM
length of the hind. A smaller individual of C. acutus had a ratio of 98%, while
in a juvenile of Alligator mississippiensis the ratio was 91%. Although both
femora of Orthosuchus are now broken, the suggested measurement of 57 mm
can only be a little short, if at all, of the true length. Using this measurement,
the fore-limb is then 91% the length of the hind. Orthosuchus falls well within
the range for living crocodilians and was quadrupedal (Fig. 30).
Ewer (1965) noted the significance of trunk length in determining the type
of locomotion. Trunk length is taken as the distance along the vertebral column
from a point directly over the pectoral glenoid to one over the acetabulum.
The longer the trunk, relative to the length of the hind-limb, the more
quadrupedal the type.
Of four large specimens of Crocodylus studied, it was found that in
C. niloticus the hind-limb was 44% and 52% the length of the trunk, in C. acutus
it was 49%, and in C. porosus 50%. In a juvenile Alligator the trunk is shorter
relative to the length of the hind-limb, as is also the case in Orthosuchus, a value
of 61% being obtained in both types. Ewer found a value of 40% and 50% in
Varanus and Sphenodon respectively, and 70° in the fast-running quadrupedal
Agama.
There can be little doubt that Orthosuchus could rest on its belly with the
femur held horizontally as is the practice of modern reptiles. On the other hand,
the structural features of the femur and pelvis indicate that the femur was
pulled into a more nearly vertical position when the animal was moving.
Living crocodiles when walking unhurriedly over land, and when hauling out of
water, do so with the limbs held vertically ; a movement described by Cott (1961)
as the high walk. A comparison of the femur of Orthosuchus with that of living
crocodiles shows remarkable similarity in form between the two types. However,
in Orthosuchus the articular surface of the femoral head is more medially
directed, suggesting that the femur was usually held in a vertical position. The
deep acetabulum and pronounced supra-acetabular buttress would have been
advantageous with the femur held in this position. The musculature essential for
this type of movement was well developed.
The crurotarsal ankle joint of Orthosuchus is admirably suited to the action
of a vertically orientated femur. By virtue of the longer length of the tibia
posteriorly, it is possible for the pes to be forwardly directed while the crus
passes backwards. The presence of a calcaneal tuber clearly gives greater
leverage to movement of the foot. Orthosuchus would therefore have moved in a
plantigrade manner, much as living crocodiles do when walking over land.
Although the pectoral musculature was apparently well developed, in
certain respects the humerus of Orthosuchus is more primitive in form than that
of living crocodiles. The pronounced medially directed deltopectoral crest seems
to obviate any possibility of the humerus being held vertically. In this position
it would foul the coracoid, on protraction at least. The humerus also shows
greater twisting through its axis, so that the distal articular surface, and the
lower fore-limb, must have been more laterally directed than it is in living
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 295
crocodiles. On the other hand, the elongation of the radiale and ulnare appears
to be a means of both lengthening the fore-limb, and providing a more flexible
wrist movement, so making quadrupedal locomotion more efficient.
On rare occasions juvenile crocodiles move by galloping. Cott (1961) likens
this movement to the bounding of a squirrel. In view of the more primitive
posture held by the fore-limb of Orthosuchus it seems unlikely that the animal
could have moved in this way. The lengthening of the coracoid in living croco-
diles has presumably increased the length of the adductor muscles of the fore-
limb, and also their efficiency.
Similarly, the tail in living crocodiles is probably used more efficiently as a
propulsive organ when swimming than it was in Orthosuchus. The vertebrae of
Orthosuchus are amphicoelous, whereas in eusuchians they are procoelous
except for the centrum of the first caudal, which is biconvex. Further, the
caudifemoral musculature was probably less well developed, as seen by the small
size of the fourth trochanter. Presumably therefore the tail was less mobile than
it is in living crocodiles, where it can be swung through a full circle of 360 degrees.
RELATIONSHIP OF ORTHOSUCHUS TO THECODONTIANS
Orthosuchus demonstrates many characters which are indicative of a
thecodontian heritage. The snout is slender, as is typical of thecodontians like
Cerritosaurus (Price 1946), and the maxilla is excluded from the boundary of
paired, laterally orientated external nares. The teeth are thecodont, conical and
undifferentiated.
An antorbital fenestra, bounded by the maxilla antero-ventraily and by the
lachrymal postero-dorsally, is present in Orthosuchus as in thecodontians.
Similarly, both temporal openings are well developed. The posterior margin of
the inferior temporal fenestra is angular.
There is no tabular or supratemporal in Orthosuchus, and a postparietal is
also absent. This element is known in a few thecodontians such as Euparkeria
(Ewer 1965), but is lost in later forms. A parietal foramen is absent, as it is in the
majority of thecodontians. A very tiny parietal foramen is present in certain early
types such as Chasmatosaurus (Brink 1955) and Erythrosuchus (Broom 1905).
An otic notch is present in Orthosuchus. Although this is not developed in
early thecodontians like Chasmatosaurus (Broili & Schréder 1934), it is generally
present in later forms. In these types, as in Orthosuchus, the notch is open
posteriorly.
In Orthosuchus the head of the quadrate rests in a socket of the squamosal
adjacent to the paroccipital process. This is characteristic of thecodontians such
as Stagonolepis. Similarly, a small basipterygoid process of the basisphenoid is
present in Orthosuchus. In thecodontians such as Chasmatosaurus and
Euparkeria these joints are highly developed and mobile. Case (1922) describes
prominent basipterygoid processes in Desmatosuchus spurensis and Walker
(1961) notes their presence in Stagonolepis, and (1964) in Ornithosuchus. Short
296 ANNALS OF THE SOUTH AFRICAN MUSEUM
basipterygoid processes allowing for some possibility of movement are also seen
in phytosaurs such as Machaeroprosopus (Camp 1930).
In that the pterygoid is firmly applied both to the quadrate and to the
basipterygoid process, it seems unlikely that the basal articulation of Orthosuchus
was movable. Similarly, movement between the squamosal and the superior
edge of the quadrate and quadratojugal seems unlikely, and the supraoccipital
and parietal are firmly united. The skull of Orthosuchus should not therefore be
regarded as kinetic, though it cannot be far removed from an ancestor with this
type of skull.
In both Desmatosuchus and Stagonolepis a simple pit lies between prominent
basipterygoid processes in exactly the same position as in Orthosuchus. It
therefore seems probable that the median eustachian system of the Crocodilia
does not correspond to the spiracular gill slit, but is a secondary formation
developed as an intucking of the basisphenoid.
The lower jaw of Orthosuchus is slender with a very large external
mandibular fenestra. This is characteristic of thecodontians, with the exception
of primitive forms such as Chasmatosaurus and Erythrosuchus. The internal
adductor fossa is also large.
Thecodontians show a range in the count of presacral vertebrae. Euparkeria
has 22, while both Vjuskovia (Von Huene 1960) and Stagonolepis have 25. In
Orthosuchus the number is probably 24. There are seven cervicals in Euparkeria,
while the probable number in Orthosuchus is eight. The centra are amphicoelous .
in each case.
Behind the axis, the cervical vertebrae of both Orthosuchus and thecodon-
tians like Euparkeria and Ornithosuchus show strong ventral keeling, and
hypapophyses are not developed. The dorsal vertebrae, however, are not
keeled.
The areas of attachment for the cervical ribs of Orthosuchus are distinctive
and show the same progressive change in rib articulations that occurs in
archosaurs generally. The parapophysis of the anterior cervical vertebra lies
antero-ventrally on the centrum. Above this the diapophysis lies anteriorly, low
down on the neural arch. Back through the cervicals the parapophysis moves up
the centrum, while the diapophysis rises and moves posteriorly so that at the end
of the series it is borne on a transverse process.
The migration of the parapophysis continues through to the dorsals, where
early on it comes to lie wholly above the neurocentral suture. In Euparkeria it
reaches this position on vertebra fourteen (i.e. dorsal six), in the phytosaur
Machaeroprosopus on vertebra twelve (i.e. dorsal five) and in Stagonolepis on
vertebra ten (i.e. dorsal one). In Orthosuchus this occurs on vertebra twelve
(i.e. dorsal four). The diapophysis does not change its position further back in
the series, though the parapophysis continues to move upwards until the two
facets come to lie at the same level on the posterior dorsals. They become
confluent on the last presacral. This can be seen in Chasmatosaurus and
Ornithosuchus, and also occurs in Orthosuchus. In many thecodontians, such as
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 297
Euparkeria and Chasmatosaurus, the transverse processes become shorter from
the mid-dorsal vertebrae back to the sacrals, as they do in Orthosuchus. In
others, such as Stagonolepis and Parringtonia (Von Huene 1939), they remain
the same size.
The sacral vertebrae of Orthosuchus are very similar to those of many
thecodontians. Orthosuchus has retained the primitive count of two sacrals, as
found in various thecodontians, including Erythrosuchus, Aétosaurus (Walker
1961) and Ticinosuchus (Krebs 1963a). In other thecodontians, however, the
number of sacrals has been increased. In Ornithosuchus there are three, and in
Schleromochlus (Woodward 1907) there are four.
Atlantal and axial ribs are poorly known in thecodontians, but in some
types at least, as in Euparkeria and Chasmatosaurus, they are single-headed, as
in Orthosuchus. Other cervical ribs of Orthosuchus have a shape which is typical
of both thecodontians and crocodilians. That is, the blade of the rib is directed
antero-posteriorly with two heads rising vertically from a point along the
length of the shaft. The length of the shaft is variable. It is long and slender
relative to the length of the cervical vertebra in Chasmatosaurus, but it is short
in Orthosuchus, as it is in the unrelated phytosaurs.
In both thecodontians and crocodilians the cervical ribs grade into the
quite different form of dorsal rib. Coupled with the change which occurs in the
position of the rib facets, the capitulum becomes terminal on the rib shaft, and
the tubercle gradually approaches this. In Orthosuchus the two processes remain
distinct throughout the major part of the dorsal series. A similar condition
occurs in Euparkeria.
The scapula of Orthosuchus is similar to that of Stagonolepis and Euparkeria
in that it is a tall, narrow element expanded at both its upper and lower ends. In
many of the less specialized lepidosaurs like Sphenodon the scapula is relatively
short and broad.
The humerus is long and slender in Orthosuchus as in thecodontians
generally, as shown by Hesperosuchus (Colbert 1952) and Ornithosuchus. It
further resembles that of thecodontians like Euparkeria and differs from
lepidosaurs in that distally the two condyles are well apart, and the ect-
epicondylar and entepicondylar expansions are slight.
In Orthosuchus as in all archosaurs, the ulna is stouter than the radius.
Further, there is no development of an olecranon process, and the head of the
ulna is expanded for weight support. In thecodontians generally there is little or
no development of an olecranon, though Walker (1961) notes its presence in
Stagonolepis.
The ilium of Orthosuchus is, in general form, persistently primitive. As in
Erythrosuchus and Machaeroprosopus it has a low iliac blade with a well-
developed posterior extension. There is also a pronounced supra-acetabular
buttress overhanging the acetabulum.
In the presence of a fourth trochanter and absence of a greater trochanter,
the femur of Orthosuchus closely resembles that of thecodontians. Ewer descrites
298 ANNALS OF THE SOUTH AFRICAN MUSEUM
the fourth trochanter of Euparkeria as lying well down the shaft, but in both
Orthosuchus and Stagonolepis it arises about one-third of the way down.
The ankle joint of Orthosuchus is of crurotarsal type, a feature which has
been considered of diagnostic significance for the Crocodilia. Yet, as Krebs
(19635) has shown, the crurotarsal ankle joint is common to all but a few
pseudosuchians of Lower and Middle Triassic age. Euparkeria has the crurotarsal
type of ankle joint, though it lacks the elaborate articulation between the
astragalus and calcaneum that is found in crocodiles. The ankle joints of
Aétosaurus and Typothorax are also crurotarsal (Walker 1961). In these forms,
as in Orthosuchus, two distal tarsals are present. A pronounced tuber is present
on the posterior face of the calcaneum of Orthosuchus, and this too occurs in
some thecodontians, such as Saltoposuchus (Von Huene 1921) and Aétosaurus.
Thecodontians in general show some reduction of the fifth digit. In
Euparkeria the fifth metatarsal is somewhat hooked in shape, and bears three
shortened phalanges. In Aétosaurus the fifth digit is more or less similar to this
but bears four phalanges. In Orthosuchus this metatarsal is further reduced and
phalanges are lacking. The pseudosuchian Schleromochlus apparently shows a
condition identical to that seen in Orthosuchus.
Orthosuchus was protected dorsally throughout its length by a paired row
of bony plates. Many thecodontians show a similar development of armour.
Ornithosuchus has, as far as is known, only dorsal armour, the scutes being
arranged in a paired, longitudinal series. Ticinosuchus and Euparkeria have both
dorsal and lateral scutes, while Stagonolepis was protected at least in part on its
ventral surface as well. Ornithosuchus and Stagonolepis also show the same
bending down of the outer portions of the scutes at right angles to the dorsal
portions, as seen in Orthosuchus. Furthermore, the characteristic peg and socket
articulations between adjacent scutes in Orthosuchus also occurs in both
Stagonolepis and Ornithosuchus.
Gastralia are present in Orthosuchus, as they are in thecodontians such as
Euparkeria, Ticinosuchus, Stagonolepis, Schleromochlus and Rutiodon (McGregor
1906).
RELATIONSHIP OF ORTHOSUCHUS TO CROCODILIANS
There are few unquestionable crocodilians of Upper Triassic age. Proto-
suchus richardsoni from Arizona is the best known of these. The type described
by Colbert & Mook (1951) is a nearly complete, articulated skeleton. Seven
other partial skeletons were also collected from the same locality and
stratigraphic level.
Two partial skeletons of Notochampsa istedana are known. The type
described by Broom (1904) from the Cape Province is an impression of the
nasal and temporal regions of the skull, a good pectoral girdle and fore-limb,
part of the hind-limb and dorsal armour. The other specimen, British Museum
(Natural History) No. R8503, as yet undescribed, is from Lesotho and consists
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 299
es
\
ics
Fig. 31. Dorsal views of the skulls of A. Orthosuchus stormbergi (2 natural size). B. Steneo-
saurus durobrivensis, after Andrews (1913a) (5 natural size). C. Gavialis gangeticus (juvenile).
of the left temporal region of the skuil and part of a jaw ramus, a few vertebrae,
some isolated limb elements and armour.
Erythrochampsa longipes is rather poorly known only from part of the
postcranium and armour. The type specimen is from the Cape Province, and was
described by Broom (1904). Further material from the Orange Free State was
assigned to this genus by Broili & Schréder (1936).
In general form the skull of Orthosuchus is very similar to those of Proto-
suchus and Notochampsa. In all these forms the cranial table is broad and flat,
and is of characteristic crocodilian shape. The snout is slender and clearly
marked off from the rest of the skull, a pattern which is seen in many meso-
suchians, notably teleosaurs, metriorhynchids and pholidosaurs, and one which
is retained in certain eusuchians, principally Gavialis (Fig. 31). Further, the
bones are sculptured in true crocodilian fashion in all these forms.
It seems likely that the external nares of Orthosuchus are similar to those of
Notochampsa. Haughton (1924) felt that there was sufficient evidence in the
nasal region to prove that they were paired and somewhat lateral. The tip of the
snout of Protosuchus is not known, but in Steneosaurus (Andrews 1913a) the
external nares are confluent. Ventrally the premaxilla of Steneosaurus is not
extensive and differs little from that of Orthosuchus.
300 ANNALS OF THE SOUTH AFRICAN MUSEUM
In both Orthosuchus and Protosuchus there is a short region of the snout
which does not bear teeth. In Protosuchus this is described as being comparable
to the notch region in later crocodiles, though in Orthosuchus a prominent
notch also occurs. Behind this there are in Orthosuchus three or four maxillary
teeth, in Notochampsa between six and nine, and in Protosuchus about eleven.
Von Huene (1925) described the teeth of Notochampsa as not compressed and
without edges, and this description is equally applicable to Orthosuchus and
Protosuchus.
The orbits of Orthosuchus and Protosuchus are directed outward and
forward rather than predominantly upward as in more advanced forms. There
is now no doubt that Colbert & Mook (1951) interpreted the orbital region of
Protosuchus incorrectly. They considered the orbital region to be wide, but
clearly in both Orthosuchus and Protosuchus this region is relatively narrow and
is bordered on either side by two supraorbital elements. Unfortunately, the
orbital region of Notochampsa is not preserved, though in the more recent
material, British Museum (Natural History) No. R8503, the anterior margin of
the left postorbital bears a facet for a supraorbital element. This surely indicates
that the orbit was outwardly directed as in Orthosuchus.
The postorbital-jugal bar in Orthosuchus and Protosuchus is superficial in
position, as it is in teleosaurs. The superior part becomes sunken in
metriorhynchids, pholidosaurs and notosuchids, and it is wholly ae
internally in goniopholids and atoposaurs, and in all eusuchians.
Much has been written regarding the position of the orbits of crocodiles in
relation to an aquatic mode of life. It seems that the change in orientation from
the more lateral position found in early crocodiles to the dorsal inclination of
living forms should be attributed to the movement of the jugal upwards from an
essentially horizontal position in Orthosuchus into a nearly vertical one. This is
coupled with the inward movement of the postorbital bar. Supraorbital elements
lie over the roof of the orbit in the eyelid in both modern crocodiles and
Orthosuchus.
Despite the position of the orbits in Orthosuchus, there are clear indications
in the form of the palate and overall shape of the skull, that the trend towards a
semi-aquatic mode of life was already well established. In view of the remarkable
similarity in form between the North American and southern African proto-
suchians, it seems that Protosuchus too was rather more aquatic in habit than
has previously been supposed.
Differences between the protosuchians lie in the shape of the superior
temporal fenestrae. Notochampsa is characterized by the moderately sized,
almost pear-shaped superior temporal fenestra, which lies with its broadest end
anteriorly. In Protosuchus this fenestra is circular in shape, and is smaller than
the orbit, and in this respect shows an inherent thecodontian condition. In
Orthosuchus the superior temporal fenestra is also circular, but equals the size
of the orbit. This indicates a progression toward the condition shown by early
mesosuchians such as Teleosaurus (Andrews 1913a) where the fenestra is greatly
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 301
enlarged at the expense of the parietal and squamosal.
Kalin (1955) suggests that the enlargement of the superior temporal
fenestra seen especially in long snouted groups, such as the gharial, is correlated
with the form of the musculature. In those longirostrate forms adapted for fish
eating there is a need for a well-developed m. adductor mandibulae externus,
which arises within the superior temporal fossa. On the other hand, in breviro-
strate crocodiles, where prey is held in the water to drown it, or where large
animals are torn apart, it is the m. pterygoideus which is strongly developed.
Since Orthosuchus would appear to be more progressive than Protosuchus
with regard to the size of the superior temporal fenestra, it is surprising that an
antorbital fenestra is present in Orthosuchus and lacking in Protosuchus.
Unfortunately this question remains unresolved for Notochampsa. Haughton
(1924) omits to mention this point at all. According to Von Huene (1925) an
antorbital fenestra is present in Notochampsa, though Broom (1927) considered
it absent. In fact, the material is too incomplete for a proper determination to
be made.
Among mesosuchians an antorbital fenestra is seen in teleosaurs like
Steneosaurus and Pelagosaurus (Eudes-Deslongchamps 1864) and also occurs in
notosuchids such as Notosuchus and Araripesuchus (De Gasparini 1971). It has
previously been supposed that this represents a secondary formation, but the
presence of an antorbital fenestra in Orthosuchus demonstrates that this is not
the case. In all types the opening lies between the maxilla and lachrymal.
In Orthosuchus the frontal contributes to the anterior border of the superior
temporal fenestra, as it does in ail mesosuchians with the exception of a few
notosuchids like Notosuchus. In view of this it seems most unlikely that the
frontal is excluded from the fenestra in Pretosuchus, as Colbert & Mook (1951)
suggest. They indicate a short suture running from the medial border of the
superior temporal fenestra on each side, and it seems likely that this represents
the fronto-parietal suture. In eusuchians this suture is located further
forward, so that the parietal articulates with the postorbital.
In both Orthosuchus and Notochampsa British Museum (Natural History)
No. R8503 a deep otic recess is made between the overhanging squamosal and
sloping quadrate. In each case the squamosal bears a longitudinal groove,
indicating that in life the recess was concealed by ear-flaps.
As is also characteristic for the Crocodilia, in both these forms the quadrate
contributes to the floored-in area of the superior temporal fossa. The relation-
ships of the quadrates of Notochampsa and Orthosuchus are, in fact identical.
In both the otic notch is shaliow and the quadrate has a posterior articulation
with the squamosal. Mesosuchians differ in that the notch is deepened by the
loss of the posterior articulation, as seen in Pelagosaurus. In eusuchians the
squamosal extends down to close the notch posteriorly (Fig. 32).
Notochampsa also has a highly fenestrated quadrate, though it differs from
that of Orthosuchus in its overall pattern. A well-developed rhomboidal sinus
lying behind the prootic-opisthotic suture, between the braincase and
302 ANNALS OF THE SOUTH AFRICAN MUSEUM
A\
eT
ee
N
Fig. 32. Lateral views of the skulls of A. Orthosuchus stormbergi (natural size). B. Pelagosaurus
typus, after Eudes-Deslongchamps (1864) (4 natural size). C. Crocodylus, after Romer (1956).
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 303
exoccipital, is present in Notochampsa. This clearly indicates an early stage in
the enlargement of the tympanic cavity in crocodiles. The rhomboidal sinus
probably opened into the throat through lateral eustachian foramina. These can
be seen in Orthosuchus. |
In living crocodiles the tympanic cavities are further enlarged and linked
together by large air spaces within the supraoccipital. Each cavity is also
connected to air passages within the quadrate and articular as well as to a
rhomboidal sinus. This is linked to the throat by a lateral eustachian tube which
passes up between the basisphenoid-basioccipital suture. In addition, a third
eustachian tube passes up in the midline and divides above into an anterior and
a posterior canal. Each of these forks into horizontally running canals which
also terminate in the rhomboidal sinus.
This complex system of eustachian tubes is partially developed in Pelago-
saurus and Steneosaurus, where the anterior branch of the median eustachian
tube, with its horizontal branches, is found. Unfortunately, in Orthosuchus it is
impossible to determine whether the comparable anterior canal also leads to a
rhomboidal sinus on either side.
A very large horizontal canal connects the median and lateral eustachian
foramina on each side in Pelagosaurus and Steneosaurus. These surely represent
the canals of the posterior branch of the median eustachian system. These
canals are not developed in Orthosuchus. On the other hand, in both Orthosuchus
and Steneosaurus an additional canal leads backwards from the median opening
and, in Steneosaurus, runs into the body of the basioccipital and Andrews
(1913a) was of the opinion that this canal was vascular in function.
In mesosuchians the lateral eustachian tubes are membranous, as is the
median tube, and each lateral foramen opens above directly to a rhomboidal
sinus. Presumably the enclosure of these tubes by bone occurred as a result of
growth of the basioccipital and basisphenoid. This, coupled with the more
limited extension of the pterygoid posteriorly, must account for the apparent
change in position of the eustachian foramina from the pterygoid-basisphenoid
suture, as in Orthosuchus, to that between the basisphenoid and basioccipital, as
in later crocodiles. |
This complex eustachian system in crocodiles probably functions as a
series of resonance chambers to improve auditory capacity. Wever & Vernon
(1957) have demonstrated that because the two middle ear cavities are inter-
connected, the application of sounds to one ear stimulates the other ear almost
equally well.
The pterygoid of Steneosaurus is essentially similar in form to that of
Orthosuchus (Fig. 33). In both types it has a posterior process which extends
back along the side of the basis cranii as far as the basisphenoid-basioccipital
suture and makes contact with the exoccipital so that the quadrate does not
meet the basisphenoid as it does in modern forms. In Pelagosaurus this process
of the pterygoid does not extend as far back, and in modern forms only a
vestige remains.
304 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 33. Basicranial region of the skulls of A. Orthosuchus
stormbergi (natural size). B. Steneosaurus durobrivensis,
after Andrews (1913a) (4 natural size).
In Orthosuchus, as in Steneosaurus and Pelagosaurus, the flange of the
pterygoid does not project below the level of the alveolar border to the same
extent as seen in recent crocodiles. This downward movement of the flange in
living crocodiles has involved a change in orientation of the ectopterygoid,
so that it lies essentially in a vertical position. In Orthosuchus and Steneosaurus
it is more or less horizontal.
Mesosuchians, again, differ from Orthosuchus in that the palatine forms
part of the wall and floor of the secondary palate. This is brought about by
fusion of the palatines ventrally in the midline, and has also resulted in the
enlargement of the postpalatine fenestra on each side. In forms like Pelagosaurus
and Pholidosaurus (Andrews 1913b) the choanae are large and confluent, and
the anterior border is formed by the palatine and the pterygoid forms the roof.
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 305
In recent crocodiles the pterygoids also contribute to the secondary palate
and are hollowed out behind the palatines and between the pterygoid flanges so
that the narial tubes are continued posteriorly. The ridges on the body of the
pterygoids of Orthosuchus mark the position of the choanae in eusuchians
(Fig. 34). A transitional stage between these is seen in the mesosuchian
Theriosuchus (Joffe 1967) where the internal nares lie in a depression which is
walled laterally by the pterygoids and roofed by the palatines.
The prefrontal of Orthosuchus forms an internal process which extends
down towards the palate. In living crocodiles this process unites with that from
the other side in the midline, and also with the palatine and pterygoid. A similar
process occurs in Pelagosaurus British Museum (Natural History) No. 32599
(oe)
Fig. 34. The palate of the skulls of A. Orthosuchus stormbergi (3 natural size). B. Pelagosaurus
typus, after Eudes-Deslongchamps (1864) (¢ natural size). C. Alligator mississippiensis, after
Kalin (1955) (? natural size).
and also in Steneosaurus and Metriorhynchus, though it is incomplete in each
case. Andrews (1913a) was of the opinion that in the latter types these processes
did meet in the midline and also met the palatine ventrally.
The symphysial region of the lower jaw of Protosuchus is unknown, but in
Notochampsa (Haughton 1924) and Orthosuchus it is short. In Protosuchus and
many thecodontians the rear end of the lower jaw lacks the marked prolongation
of the angular and articular bones that characterizes mesosuchians and
eusuchians, and this is probably also the case in Orthosuchus.
Orthosuchus is the first Triassic crocodile known to possess a prearticular
(the internal surface of the lower jaw is unknown in Protosuchus) and, indeed,
306 ANNALS OF THE SOUTH AFRICAN MUSEUM
this element is known in only two other crocodilians. It is well developed in
Pelagosaurus, of Lower Jurassic age, but in the Upper Jurassic form Metrio-
rhynchus it is quite short. It is absent as a separate element in Crocodilia from
more recent strata. In contemporary crocodilians, where the prearticular fuses
with the articular during development (De Beer 1937) the region normally
occupied by the prearticular is filled by a medial process of the angular.
The gradual elimination of the prearticular through the Crocodilia is shown
in Figure 35.
Fig. 35. Medial view of the lower jaws of A. Orthosuchus stormbergi (natural size). B. Metrio-
rhynchus cultridens, after Andrews (1913a) (4 natural size). C. Crocodylus vulgaris, after Kalin
(1955) ( natural size).
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 307
Although the entire vertebral column of Protosuchus is known, it is
obscured dorsally, and to a lesser extent ventrally, by scutes. Colbert & Mook
(1951) estimated a presacral count of 24 vertebrae, as is suggested for Ortho-
suchus. All early crocodilians possess thecodontian spool-shaped, amphicoelous
vertebrae, and this condition was retained generally among mesosuchians.
The anterior vertebrae are keeled in Orthosuchus, but in the marine Jurassic
crocodiles this keeling is lost. In living crocodiles keeling has been extended to
include the anterior dorsals.
A similar change in position of the rib facets through the cervical series
back down the column occurs in both Protosuchus and Orthosuchus. In Proto-
suchus the parapophysis lies at the base of the transverse process above the
neurocentral suture on vertebra twelve, and in this respect shows an identical
condition to that seen in Orthosuchus. Further, in both types the two rib facets
remain distinct up to the last presacral vertebra, a condition retained in forms
like Steneosaurus.
Protosuchus and Orthosuchus also share the primitive count of two sacral
vertebrae. In both forms, and in later crocodiles, the extremities of the sacrum
are large because each sacral rib articulates both with the centrum and with the
transverse process.
The cervical ribs of Protosuchus and Erythrochampsa (Broili & Schréder
1936) closely resemble those of Orthosuchus, but are also indistinguishable from
those of many thecodontians as seen in Ticinosuchus.
The dorsal ribs of Protosuchus are of particular interest, and are identical
to those of Orthosuchus. In both types the shaft is flanked by an antero-ventral
and a postero-dorsal flange. In the thecodontians Euparkeria and Ticinosuchus,
they are not flanged in this way, while in teleosaurs only anterior flanges are
developed in the first two or three dorsal ribs.
The scapulae of Orthosuchus, Protosuchus and Notochampsa are strikingly
similar in shape. They are crocodilian in form in that the upper end is expanded,
particularly posteriorly, and is very much larger in size than the lower end. This
expansion is not found in teleosaurs (Fig. 36), but this no doubt is an
aquatic adaptation. However, the coracoids of these forms are closely similar in
shape, and are elongate elements expanded both proximally and distally. This
represents an advanced stage over that seen in typical thecodontians where the
coracoid is rather an insignificant element, subcircular in shape.
In addition, these early crocodilians differ from, and are more specialized
than, thecodontians in that they lack a clavicle. Von Huene (1925) did
suggest that a clavicle is present in Notochampsa, though Broom (1927) expressed
the opposite view and believed the element in question to be a scute. Broom
appears to have been correct in this.
In 1924 Haughton expressed doubts concerning the crocodilian form of the
humerus of Notochampsa. He was of the opinion that it is more thecodontian in
form, much like that of Stagonolepis. However, the description does not support
this view, and the recent find of Notochampsa includes two humeri which are
308 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 36. Lateral views of the pectoral girdles of A. Pretosuchus richardsoni, after Colbert &
Mook (1951) (natural size). B. Notochampsa istedana, after Broom (1927) (natural size).
C. Orthosuchus stormbergi (natural size). D. Steneosaurus durobrivensis, after Andrews (1913a)
(3 natural size). E. Crocodylus acutus (= C. americanus) (reversed), after Mook (19215)
(4 natural size).
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 309
very similar to those of Orthosuchus and Protosuchus. In these forms, as in
living crocodiles, the humerus bears a deltopectoral crest which rises to an apex
about one-third of the way down a long slender shaft.
A very striking crocodilian character can be seen in the form of the carpus.
Broom (1927) commented on the metacarpal-like nature of the radiale and
ulnare in Notochampsa, and an identical condition is present in both Protosuchus
and Orthosuchus. In the latter form the ulnare is only just shorter than the
longest metacarpal (the third), and the radiale is more than one and a half times
its length. Hence, as in modern crocodiles, the carpus is elongated to add an
extra segment to the fore-limb.
Protosuchus further resembles Orthosuchus in that a large pisiform element
is preserved in the carpus. However, in Orthosuchus two distal carpals are
present, whereas in Protosuchus and Notochampsa there is only one (Fig. 37).
In these forms the distal carpal lies in articulation with the ulnare.
Fig. 37. Carpus and manus of A. Protosuchus richardsoni (reversed), after Colbert & Mook
(1951) (4 natural size). B. Orthosuchus stormbergi (3 natural size). C. Crocodylus, after Romer
(1956).
The pelvis in Orthosuchus is closely similar to that of Protosuchus. In both
the acetabulum is perforated ventrally, a feature separating these forms quite
clearly from all thecodontians where the acetabulum is imperforate. The ischia
are also very alike and resemble that of Erythrochampsa. The most significant
feature here is the development of the anterior process to exclude the pubis from
the acetabuium. Unfortunately, on neither side of the body is the pubis of
Erythrochampsa complete, although Broom was of the opinion that the pubis
does not enter the acetabulum.
In all three forms the iliac blade is produced anteriorly to form a bluntly-
pointed process. This was reduced in teleosaurs, and only a vestige remains
in eusuchians (Fig. 38). In Orthosuchus, anterior to the acetabulum, the
ilium is notched. The inner of the two processes meets the ischium, and the
outer, together with the ischium, articulates with the pubis. A similar condition
probably exists in Protosuchus. In Steneosaurus the head of the pubis occupies a
more ventral position, and in living crocodiles the pubis articulates only with
the ischium.
310 ANNALS OF THE SOUTH AFRICAN MUSEUM
A its SX
SV SSS
Fig. 38. Lateral views of the pelvic girdles of A. Protosuchus richardsoni (reversed), after
Colbert & Mook (1951) (natural size). B. Orthosuchus stormbergi (natural size). C. Steneo-
saurus leedsi, after Andrews (1913a) (4 natural size). D. Alligator, after Romer (1956).
The femur of Orthosuchus is remarkably similar to that of Protosuchus, and
in fact differs little from that of living crocodiles. The tibia lacks a distinct
cnemial crest in each case. Similarly, there is little difference between the tarsus
of Orthosuchus and that of Protosuchus. In both types the calcaneum bears a
tuber which is more prominently developed than it is in living crocodiles and
which is grooved on its posterior surface. However, whereas in Orthosuchus
there are two distal tarsals, in Protosuchus only one is known.
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 311
The structure of the tarsus of Erythrochampsa is difficuit to determine.
According to Haughton (1924) it consists of two proximal and two distal
elements. The calcaneum bears a smail posterior tuber. Broom (1927) described
the same tarsus as similiar to that of the Upper Jurassic mesosuchian Alligatorel-
lus. Lortet (1892) had incorrectly described three distal tarsals in this type, but
Broom figured the tarsus of Erythrochampsa as consisting of astragalus and
calcaneum only. At most, only two elements can be identified in the type, as
shown by Broom, though it is questionable as to what these are.
The metatarsus of Orthosuchus is composed of four elongated metatarsals
with the fifth reduced to a hook and heid behind the other four. Protosuchus
shows a similar condition. Broom (1904) described four digits in Erythrochampsa,
but in 1927 figured four elongated elements and a short, hook-shaped fifth.
Von Huene (1925) figured the same specimen showing four elongated and a
slightly shortened fifth metatarsal in articulation with two phalanges.
At my request, Dr M. A. Cluver (South African Museum) examined the
type specimen and came to the conclusion that both Broom and Von Huene
had misinterpreted the specimen. Broom gave the correct number of unreduced
metatarsals (four), but did not indicate that the fourth is incomplete. Von Huene
miscounted and regarded the broken fourth metatarsal as a fifth member and
figured it in articulation with phalanges. Although the material is difficult to
interpret, there is a piece of bone underlying the proximal end of the fourth
metatarsal which does not seem to be part of the distal end of the tibia or
fibula. This could well be a reduced fifth metatarsal. On balance it seems likely
that the metatarsus of Erythrochampsa is identical to that of both Orthosuchus
and Protosuchus (Fig. 39). In view of this, it is unfortunate that Kalin (1955)
chose to reproduce Von Huene’s incorrect figure.
Protosuchian armour may be distinguished from thecodontian armour by a
combination of characters. The dorsal scutes are arranged in two rows, with the
anterior margin of each overlapped by the preceding scute. The articular surface
of each dorsal scute is smooth, and behind this the exposed surface is strongly
pitted. Further, the lateral portion of each scute is strongly bent downward and
the bend is strengthened by a dorsai ridge.
Many mesosuchians such as Alligatorellus show an identical development
of dorsal armour. In others like Crocodileimus (Lortet 1892), although the scutes
covering the neck and lumbar region are paired, the mid-dorsals are arranged in
four rows. In living genera as many as ten scutes may be present in a row.
The most anterior scutes of the dorsal armour of Orthosuchus and Noto-
champsa carry a peg-like process which extends forward from the anterior
margin of the dorsal ridge, and fits into a groove on the ventral surface of the
preceding scute. This feature is inherited from thecodontians, and is one which
was passed on to at least some of the mesosuchians, like Steneosaurus,
Pholidosaurus and Goniopholis.
Although ventral armour of the type specimen of Notochampsa is not
known, five ventral scutes form part of the material of specimen British Museum
312 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 39. Tarsus and pes of A. Protosuchus richardsoni (reversed), after Colbert & Mook (1951)
(¢ natural size). B. Erythrochampsa longipes, after Broom (1927). C. Erythrochampsa longipes
(reversed), after Von Huene (1925). D. Orthosuchus stormbergi (2 natural size). E. Crocodylus,
after Romer (1956).
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 313
(Natural History) No. R8503. Protosuchus and Erythrochampsa were also
protected by dorsal and ventral, but not lateral scutes, except over the tail.
Probably Orthosuchus, too, was covered in this way, though its ventral armour
is as yet unknown. The teleosaur Mystriosaurus has a similar covering of
ventral armour, and it is also found in Crocodileimus and Pholidosaurus,
although here the scutes are polygonal in shape. Ventral armour is not known
in atoposaurs, and is generally absent in living crocodiles.
Ventrally and at the level of the distal portion of the pubis, both Orthosuchus
and Protosuchus show an accumulation of gastralia. In life these were probably
more numerous and extended farther forwards. Gastralia are known in some
mesosuchians, like Atoposaurus and Alligatorellus, and are present in eusuchians.
LIMB AND LIMB GIRDLE PROPORTIONS IN
CROCODILIA AND THECODONTIA
Relationship between lengths of scapula and humerus (Fig. 40)
The relationship between length of scapula and length of humerus for the
seven available pairs of measurements on contemporary animals may con-
veniently be shown by plotting log,, humerus length (x axis) against logy) of
scapula (y axis). They may be represented by the regression line y = 1,167x—
0,522. If Y and X are the actual lengths of scapula and humerus respectively, the
equation becomes Y = 0,301X?1”.
The regression coefficient 1,167 is significantly different from 1,0 (t = 4,674
with 5 d.f.). This shows that the scapulae of these contemporary crocodilians
increases in length at a proportionally greater rate than the humerus, i.e. the
ratio between the two is greater in the larger animals.
In Figure 40 are included points for selected fossil crocodilians and
thecodontians, though of these the thecodontians Vjushkovia and Euparkeria,
and the protosuchians Orthosuchus and Protosuchus, form a group of very
ancient animals which may reasonably be separated from the mesosuchians,
whether land or marine forms.
The regression coefficient for the above four is given by b = 1,170. This
is patently not significantly different from the above value 1,167. The separation
of these two regression lines is however significant at the 1% level (t = 3,780
with 7 d.f.). That is, these four animals form a group separate from the con-
temporary crocodilians. The equation for the above four is: y; = 1,170x,;—
0,3107 so that the initial dimensions are related by: Y, = 0,489X,?”.
Comparing this with the previous equation it is seen that in these forms the
ratio of scapula length to humerus length is about 1,6 times as great, for any
particular value of scapula length. These higher ratios are in fact very apparent
in Table 4, and clearly apply to Orthosuchus. The very high value of the ratio for
Vjushkovia is not out of place, and can be attributed to the fact that it is a
member of this ancient group, and is a very large animal.
ANNALS OF THE SOUTH AFRICAN MUSEUM
314
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THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN
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ANNALS OF THE SOUTH AFRICAN MUSEUM
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THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 317
The mesosuchians on this graph show rather more scatter than the
eusuchians, but no different trends. The regression slope, and the relationship
with size which it shows, appear to be applicable to all of these forms.
Relationship between lengths of scapuia and coracoid (Fig. 41)
It can be seen from Table 4 that in the thecodontian Vjushkovia, and the
protosuchians Orthosuchus and Protosuchus, the coracoid is approximately half
as long as the scapula, whereas in the modern crocodilians these elements are
more or less equal in iength. The relationships may be shown when log,) scapula
is plotted against log,, coracoid. This has been done for all contemporary and
fossil forms.
Two regression lines are of interest. The regression of log coracoid (y) on
log scapula (x) for the seven contemporary forms is given by: y = 0,9815x +
0,0150. The regression for Vjushkovia, Orthosuchus and Protosuchus however is
given by: y; = 1,1007x, — 0,4605.
The difference between the coefficients 0,9815 and 1,1007 in the above
equations is significant at the 5% level.
It now becomes interesting to note that Steneosaurus, Alligatorium and
Crocodileimus lie closely on the first line, while Alligatorellus lies virtually on the
second. Mycterosuchus is remarkabie in that the scapula is actually shorter than
the coracoid, but this is sureiy due to aquatic adaptation.
It is also surprising to find that Vjushkovia falls into a group with the
protosuchians, since in thecodontians the coracoid is normally less elongate
relative to the scapula than it is in crocodilians. In forms like Euparkeria and
Stagonolepis, for example, the coracoid is considerably wider than it is high.
Relationship between lengths of humerus and radius (Fig. 42)
The relationship between lengths of radius (Y) and of humerus (X) in fossil
and contemporary forms is again best seen in a plot of y = log Y against
x = log X. All available vaiues have been plotted.
Two regression lines are given, one for the thecodontians Vjushkovia and
Euparkeria, together with the protosuchians Orthosuchus and Protosuchus
(upper line, b = 1,007), and the other for the eight contemporary crocodilians
(lower line, b = 0,9028).
The lower line has a regression coefficient significantly smaller than 1,0
(t = 3,721, significant at the 1% level). The regression coefficients of the two
lines, however, are not significantly different (t = 1,811). The spatial separation
of the two lines though is significant at the 1 % level. Thus the-difference between
these two groups of animals is well established on this evidence.
The radius/humerus ratios in Orthosuchus and Protosuchus and in the
thecodontians Vjushkovia and Euparkeria are all very similar and very much
higher than in the living genera. This high ratio was retained, or very nearly, by
certain of the small mesosuchians Alligatorellus, Alligatorium and Theriosuchus.
However, in Crocodileimus, a mesosuchian of similar age and size, the radius is
ANNALS OF THE SOUTH AFRICAN MUSEUM
318
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THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 319
shorter relative to the length of the humerus. This animal agrees well, in respect
of radius/humerus ratio, with living genera. In these the ratio, in each case, is
less than 0,70, the ratio becoming smaller in older (i.e. larger) animals. In the
more fully aquatic mesosuchians like Steneosaurus, the ratio is rather similar to
that in contemporary crocodiles.
Relationship between lengths of radius and radiale
Data is available here for the thecodontian Chasmatosaurus, the proto-
suchians Orthosuchus and Protosuchus, the mesosuchians Alligatorellus,
Alligatorium and Crocodileimus, and for five contemporary crocodiles. In each
of these the radiale is an elongate bone, averaging in the eleven examples about
29,5% of the length of the radius. In Orthosuchus it is 41,3 % of the length of the
radius. This is significantly longer, relative to the radius, than in the other ten
animals above (significant at the 5% level).
Relationship between lengths of femur and tibia (Fig. 43)
This relationship is again brought out in a plot of log,, tibial length against
logi) length of femur. On this evidence there is no reason for separating the
mesosuchians from the contemporary crocodiles, though the thecodontians and
protosuchians could at first sight be supposed somewhat different, as could the
more fully aquatic forms.
The regression coefficient for the mesosuchians and modern forms taken
together is b = 0,8822. The difference between this value and 1,0 is significant
at the 0,1 % level. The relationship between length of femur (X) and length of
tibia (Y) for these specimens is represented by: Y = 1,31X°®°.
This equation shows that the relative length of tibia to length of femur
decreases significantly as the size of the animal increases. In other words, the
tibia elongates more slowly than the femur.
The thecodontians Vjushkovia and Euparkeria, and the protosuchians
Orthosuchus and Protosuchus, lie rather above this line, and Steneosaurus and
Mycterosuchus below it.
The regression coefficient for the thecodontians and protosuchians is
b = 1,0125 and is not significantly different from b = 0,8822 above (t = 2,038
with 12 d.f.). A test for the separation of the two lines, however, gives t = 3,768
with 12 d.f., significant at the 1% level, hence showing the thecodontians and
protosuchians to have significantly higher tibia/femur ratios at corresponding
animal sizes than the mesosuchians and eusuchians.
The tibia is therefore long relative to the length of the femur in Orthosuchus
and Protosuchus, as in the thecodontians Euparkeria and Vjushkovia. In
eusuchians, the femur grows at a faster rate than the tibia during development
from juvenile to mature stage, so that the tibia/femur ratio decreases with age
(i.e. size). The more fully aquatic Steneosaurus and Mycterosuchus have very low
tibia/femur ratios, ca 0,50, values which are low even though the large sizes of
these forms are taken into account. In Metriorhynchus this value is further
reduced to about 0,40, and this again is clearly a result of aquatic adaptation.
ANNALS OF THE SOUTH AFRICAN MUSEUM
320
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THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 321
Fig. 44. Dorsal views of the skulls of A. Orthosuchus
stormbergi (% natural size). B. Protosuchus richardsoni, after
Walker (1970) (% natural size).
RELATIONSHIP OF THE PROTOSUCHIANS TO FORMS OF
QUESTIONABLE CROCODILIAN AFFINITY
In 1967 Sill suggested Proterochampsa barrionuevoi, of late Middle Triassic
age from South America, as the most ancient of known crocodilians. Sill
believed that Proterochampsa is closely related to Notochampsa, and that
Protosuchus is representative of a more aberrant line of crocodilian heritage.
According to Sill, in both Notochampsa and Proterochampsa the skull is
relatively long compared to width, is flat, and the orbits lie in the horizontal
plane. There is, however, no reason at all for supposing that the orbits of
Notochampsa are dorsally orientated as Sill claimed. On the contrary, it seems
much more likely that they face laterally as in Orthosuchus. Again, Sill relied
heavily on Haughton’s description (1924) of the auditory region of Notochampsa,
and claimed that a further resemblance to Proterochampsa could be found in the
322 ANNALS OF THE SOUTH AFRICAN MUSEUM
og Vd MON
Fig 44 continued C. Notochampsa istedana, after Broom
(1927) (¢ natural size). D. Proterochampsa barrionuevoi,
after Sill (1967) ( natural size).
presence of an auditory canal on the posterior face of the skull. However, this
description of the type specimen of Notochampsa cannot be correct; the skull of
specimen British Museum (Natural History) No. R8503 shows a deep otic
recess, and pronounced otic notch.
The resemblances between Notochampsa and Proterochampsa are clearly
not impressive, whereas the similarity between the skulls of Orthosuchus,
Protosuchus and Notochampsa is striking (Fig. 44). Sill (1968 personal com-
munication) now agrees that he cannot substantiate his earlier claims, though
he doubts the phytosaurian affinity of Proterochampsa postulated by Walker
(1968). In Walker’s view (1970) the position of the choanae alone is sufficient to
exclude Proterochampsa as a crocodilian ancestor.
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 323
The vertebral column of Proterochampsa is known only from the thirteen
most anterior vertebrae. Rather surprisingly, the parapophysis retains a low
position on the centrum throughout this series. Presumably it changes position
farther back along the column, and so may parallel the condition found in some
primitive thecodontians like Chasmatosaurus and Erythrosuchus where the
parapophysis does not begin to rise until the eleventh vertebra (Hughes 1963).
Indeed, Romer (1971, 1972a) has now demonstrated a close relationship between
Proterochampsa, Cerritosaurus, Gualosuchus and Chanaresuchus. He has argued
convincingly that they are a sterile group, representing a modest advance over
the Proterosuchia, which in Middle Triassic times occupied a position in the
ecology similar to that of the phytosaurs of the late Triassic and the crocodilians
of the later Mesozoic.
Stegomosuchus longipes, from the Upper Triassic of the Connecticut Valley,
is poorly known because of the form of its preservation. As Walker (1970) has
commented, resemblances to protosuchians are seen in the broad, flattened
cranial table, in the longitudinal groove which runs along its margin, and in the
presence of two supraorbital elements on each side. The skull roof is pitted as
in protosuchians. Postcranially, crocodilian affinity is shown by the broad,
triangular apex of the scapula, the presence of a calcaneal tuber and four
elongated metatarsals of the hind-foot, and the similarity of the dorsal armour.
This evidence, though slender, suggests that Stegomosuchus may reasonably be
regarded as a protosuchian.
Microchampsa scutata from the Upper Triassic of the Yunnan, China, is
known only from part of the postcranium. In view of this, Simmons (1965) was
hesitant to establish its systematic position but suggested that Microchampsa
might represent a new type of crocodilian. He described the vertebral structure
as most crocodilian, but it is difficult to see what is meant by this, since the
pattern found in protosuchians is also common to thecodontians. Further,
the neural spines of the posterior dorsal vertebrae of Microchampsa are
terminally expanded to accommodate the median row of dorsal scutes, and this
is certainly not the case in any of the protosuchians. Spine tables are developed
on the posterior cervical and anterior dorsal vertebrae of Euparkeria, and on the
posterior dorsal vertebrae of Stagonolepis.
The ribs of Microchampsa are also quite different from those of Protosuchus
and Orthosuchus. In Microchampsa they are short and entirely devoid of the
double flanges so characteristic of protosuchians. Furthermore, in Micro-
champsa the lumbar ribs are fused to lateral scutes; an impossible condition
in protosuchians which lack scutes in this region. The armour, moreover, is
not of the type found in protosuchians. In Microchampsa there is but a single
median row of dorsal scutes, not pitted or ornamented in any way. This row is
flanked by a row of dorso-lateral scutes on each side. These are simply pitted
and have rugosities. These features, taken together, make a good case for
excluding Microchampsa from the Crocodilia, and regarding it, for the present,
as a pseudosuchian.
324 ANNALS OF THE SOUTH AFRICAN MUSEUM
The archosaur material from the Upper Triassic of South Wales, at present
being studied by Kermack, includes several partly associated skeletons. The
skull has an antorbital fenestra, and the carpus is crocodilian in type in that the
radiale and ulnare are elongated. With regard to the number of distal carpals
present, the specimens from Wales resemble Protosuchus and differ from
Orthosuchus. It is however surprising to find the pelvis and hind-limb more
primitive in type than that of both Orthosuchus and Protosuchus. In Kermack’s
opinion (1956), the pubis contributes to the formation of the acetabulum, and is
perforated by an obturator foramen. It is possible that in Protosuchus the pubis
contributes marginally to the acetabulum, but in neither Protosuchus nor
Orthosuchus is it perforated. On the other hand the acetabulum is open, and the
ischium similar in each case.
Kermack described the metatarsus of his material as having a fifth digit
with two reduced phalanges, and likened it to the metatarsus of Erythrochampsa.
However he relied upon Von Huene’s interpretation of Erythrochampsa which is
now shown to be incorrect. In view of the crocodilian nature of the pelvis, and
also of the carpus, there is little doubt that the Welsh form is a primitive
crocodilian, differing in a number of respects from the South African and
North American forms. ;
Hemiprotosuchus leali, from the Upper Triassic of Argentina, is represented
by a skull and jaws, and several postcranial elements. Bonaparte (1971) believes
it to be a protosuchian, and has closely compared Hemiprotosuchus with
Protosuchus.
Hemiprotosuchus is of moderate size, much the same as other known
protosuchians. The cranial table has a characteristic crocodilian configuration,
though the sculpturing is in the form of rugosities rather than pitting. The
interorbital area is slender, as it is in Orthosuchus, but the orbit and preorbital
region is higher than in other protosuchians. The nares are terminal, as in
Orthosuchus, and the superior temporal fenestra is elongate, as in Notochampsa.
An antorbital fenestra is present, as in Orthosuchus, and leads to the ventral
edge of the orbit.
As is characteristic of crocodilians, the quadratojugal and quadrate are
sutured to the postorbital and squamosal in the anterior dorsal region of the
inferior temporal fenestra. Further, the squamosal overhangs the otic region
which is open posteriorly as in protosuchians. A short, bony secondary palate is
developed, and the pterygoid and basisphenoid show the same relationship as
found in Orthosuchus. The lower jaw is persistently primitive, as in Orthosuchus
in that a prearticular is present, but a retroarticular process is lacking. However,
whereas the jaw in Hemiprotosuchus is deep, in Orthosuchus it is quite slender.
The characters of the skull indicate that Hemiprotosuchus should be regarded
as a protosuchian, and the associated postcranial elements confirm this diagnosis.
The coracoid is elongated postero-ventrally, the astragalus and calcaneum are
crocodilian in type, and the scutes covering the trunk are paired and imbricated.
Unfortunately it is not possible to determine whether the scutes are sculptured.
THE MORPHOLOGY AND RELATIONSHIPS OF A CROCODILIAN 325
Walker (1970) has suggested that a number of other Upper Triassic genera
of disputed affinities are also closely related to, though distinct from, the ‘true’
crocodiles. These are Pedeticosaurus from the Cave Sandstone of South Africa,
Sphenosuchus from the underlying Red Beds, Hesperosuchus from the Chinle
Formation of Arizona, Saltoposuchus from the Stubensandstein of Germany and
Platyognathus from the Dark Red Beds of the Lower Lufeng Series, Yunnan,
China.
Bonaparte (1971) has since demonstrated that the skull of Pseudhespero-
suchus jachaleri is closely similar to that of Sphenosuchus acutus. He is of the
opinion that rather more significance should be placed on differences between
the skulls of sphenosuchians and protosuchians, notably the absence in
sphenosuchians of the typical crocodilian cranial table, and the pseudosuchian
nature of the basicranium, suspensory region and palate. Hence, although
sphenosuchians are transitional between thecodontians and crocodilians, in
Bonaparte’s view they show a series of characters which relate them most
closely to thecodontians.
Bonaparte has further suggested that Pseudhesperosuchus is closely linked
to Hesperosuchus agilis. Evidence for this is based mainly on the form of the
coracoid and humerus. Bonaparte and Walker are both of the opinion that the
“problematical’ element figured by Colbert (1952) is a coracoid, and there can be
little dispute about this identification. However, according to Colbert, this
element consists of two bones sutured together, the suture being interrupted by
a large foramen. In my opinion this element resembles a therapsid coracoid
rather than that of an archosaur. Colbert had, in addition, identified the proxi-
mal portion of the left coracoid of Hesperosuchus.
Should the ‘problematical’ element prove to be the coracoid of a different
animal, and not that of Hesperosuchus, then the latter shows no crocodilian
characteristics. In Sphenosuchus the coracoid is elongate, and in Pseudhespero-
suchus the coracoid, radiale and ulnare are all elongated, but the presence of
certain diverse crocodilian features in these types does not in itself confirm
crocodilian affinity.
Pedeticosaurus leviseuri (Van Hoepen 1915) is preserved mainly as an
impression of the right side of the skull and limbs. The skull, as far as can be
seen, differs little from that of known protosuchians. The scapula, humerus,
radius and ulna appear to be similar in general outline to these elements in
Orthosuchus, though the coracoid is unknown. Walker has suggested that the
element previously identified by Van Hoepen as a metacarpal is an elongated
radiale associated with the first metacarpal and two phalanges. This could
indicate crocodilian affinity, but since no other bones of the carpus and manus
are present, this cannot be confirmed. Although the evidence is slight, on
balance it seems that Pedeticosaurus lies closer to protosuchians than to
forms like Sphenosuchus.
Saltoposuchus is at present too inadequately known for a meaningful
comparison to be made.
326 ANNALS OF THE SOUTH AFRICAN MUSEUM
Platyognathus hsui represents an unusual type in that the teeth are irregularly
polygonal, and finely denticulate. The jaw also differs from that of proto-
suchians in that it is short and terminally expansive. The development of a
rudimentary secondary palate in this type was regarded by Simmons (1965) as a
crocodilian characteristic, but this type of development has also been noted in
such diverse forms as Sphenosuchus, Erpetosuchus and phytosaurs. It does
not compare with the form of the palate in Orthosuchus.
Simmons (1965) described the vertebral column of Platyognathus as being
protosuchian, but with procoelous centra. The latter is clearly a very remarkable
character in a form of Upper Triassic age, and is found in neither thecodontians
nor protosuchians. The vertebrae also differ from those of protosuchians in the
development of spine tables. Further, the dorsal ribs show the unusual condition
of being flanged on the sternal segment. The dorsal scutes are paired and over-
lapping, but bear rugosities as well as pits. Indeed, the only crocodilian character
present is seen in the elongate form of the coracoid. This is clearly insufficient
reason to link Platyognathus with the Protosuchia, and its affinities more
probably lie with the Pseudosuchia.
ACKNOWLEDGEMENTS
I should like to express my sincere appreciation to Mr J. Attridge of the
University of London, Birkbeck College, for arranging the loan of the specimens
of Orthosuchus from the South African Museum, and for his continued
encouragement and interest. I am also indebted to Professor A. W. Crompton,
Director of the Museum of Comparative Zoology at Harvard University, for
allowing me to work on this material.
I owe special thanks to Dr A. D. Walker for his hospitality and for the
interesting discussions we had during my visit to Newcastle upon Tyne; for
sending me casts of Stegomosuchus and Stagonolepis and for sending me
drawings and photographs of Sphenosuchus prior to publication. I should like
to thank Dr A. J. Charig and Miss A. G. C. Grandison for their assistance in
allowing me to examine the material in the fossil and living Reptilia sections
respectively of the British Museum (Natural History). I am also indebted to
Dr E. P. Jeffree for his invaluable advice and assistance in carrying out the
statistical analysis of data.
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ABBREVIATIONS
a angular m maxilla
af _— antorbital fenestra n nasal
ar articular p parietal
bo __ basioccipital pa __ prearticular
bpt __ basipterygoid process pl palatine
bs basisphenoid pm __— premaxilla
Cc coronoid pmf premaxillary foramen
ch choana po __ postorbital
con condyle popr paroccipital process
d dentary prf _ prefrontal
ec ectopterygoid (transverse) pt pterygoid
en external naris q quadrate
€0 exoccipital qj quadratojugal
eu median opening of eustachian tubes s stapes
eul lateral eustachian pit sa surangular
f frontal sO supraorbital
fch position of functional choana soc supraoccipital
ic foramen for internal carotid sp splenial
gl jaw articulation (glenoid fossa) sq squamosal
itf inferior temporal fenestra stf superior temporal fenestra
j jugal tm tympanic membrane
] lachrymal Vv vomer
Is laterosphenoid
A.M.N.H. American Museum of Natural History, New York
B.M.N.H. British Museum (Natural History), London
S.A.M. South African Museum, Cape Town
Zool. Mus. Zoologisches Museum, Berlin
Foramina for cranial nerves in roman numerals
.
CA ge ta
ae het
ig Sais
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Figs 14-15A
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DIANE S. NASH
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BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
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FIscHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines.—Archs
Zool. exp. gén. 74: 627-634.
Konan, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.—
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Koun, A. J. 1960b. 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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ARCHAEOCYATHA PROVENANT DE BLOCS ERRATIQUES DES
TILLITES DE DWYKA (AFRIQUE DU SUD)
Par
FRANCOISE DEBRENNE
Institut de Paléontologie
Museum National d’ Histoire Naturelle, Paris
(Avec 11 figures et 1 tableau)
[MS accepté 20 Janvier 1975]
RESUME
L’étude paléontologique des Archéocyathes contenus dans les blocs glaciaires des tillites
de Dwyka montre une grande similitude avec la faune d’Antarctique. Il se confirme que
l’origine des blocs est a rechercher dans cette région. L’affinité de la faune avec |’Australie,
d’autre part, renforce I’hypothése de l’unité d’un continent austral dés le Cambrien inférieur.
Sont décrits: Pseudosyringocnemididae fam. nov., Flexanulus oosthuizeni gen. et sp. nov.,
Statanulocyathus oosthuizeni gen. et sp. nov., Andalusicyathus cooperi sp. nov.
ABSTRACT
ARCHAEOCYATHA FROM GLACIAL ERRATICS FROM THE DWYKA TILLITES (SOUTH AFRICA)
Palaeontological studies of the Archaeocyatha embedded in the erratic cobbles of the tillites
from the Dwyka sub-group give evidence of great similarity with the Antarctic faunas. They
show that the origin of these cobbles has to be looked for in Antarctica. On the other hand,
the affinities with the Australian faunas give support to the hypothesis of the unity of the
southern continents since the Lower Cambrian. Pseudosyringocnemididae fam. nov., Flexanulus
oosthuizeni gen. and sp. nov., Statanulocyathus oosthuizeni gen. and sp. nov. and Andalusicyathus
cooperi sp. nov. are described.
TABLE DES MATIERES
PAGE
Introduction ; : : . s . od2
Description paléontologique du matériel : 5 “aoe
Stapicyathus incisus (Hill) . : . : : : . “232
Thalamocyathus tectus Debrenne. . ; . 334
Flexanulus oosthuizeni gen. et sp. nov. oo» « & 330
?Ladaecyathus sp. q . , ; . 336
Thalamopectinus arterialis Debrenne : : : . 2.339
Erugatocyatus scutatus (Hill) : : : : : . 340
?Erugatocyathus sp... . 2 «342
Statanulocyathus oosthuizeni gen. ‘et sp. nov. . . . 344
?Didymocyathus sp... . 346
Archaeopharetra cf. typica R. & W. R. Bedford 5 . 347
Chouberticyathus cf. fragilis R. & W.R. Bedford . . 347
Protopharetra densa Bornemann. . a ee
Protopharetra grandicaveata Vologdin ae ae ee
?Protopharetra pauciseptata (Gordon) : : : y. «ook
Andalusicyathus cooperi sp. nov. : ; so “352
Pseudosyringocnema cf. gracilis (Gordon) . ‘ : e-, “350
Pseudosyringocnema cf. uniserialis (Hill) . ; : 5. "336
Conclusions ; : : ; ; ; ; : 2358
Remerciements . ; ; : : ; : ; . 360
Reterencest =) ; : ae eee : . 360
Sail
Ann. S. Afr. Mus. 67 (8), 1975: 331-361, 11 figs, 1 table
332 ANNALS OF THE SOUTH AFRICAN MUSEUM
INTRODUCTION
L’ensemble du matériel, comportant les Archéocyathes provenant de blocs
erratiques des tillites de Dwyka, 4 Zwartskraal a été découvert et récolté par
M. R. Oosthuizen entre 1966 et 1968. Bien que ces enclaves contenant des
Archéocyathes soient extrémement rares, on peut les suivre a travers la tillite
depuis sa base meuble, a travers la partie centrale plus dure jusqu’au sommet
plus meuble de nouveau. R. Oosthuizen a découvert quelques exemplaires in
situ dont quelques-uns sont inclus dans la gangue. M. M. Cooper a pris
l’initiative de me confier l’étude du matériel confié par M. Oosthuizen au South
African Museum. Je les remercie trés vivement et souligne le grand intérét
scientifique de cette découverte, la premiére en Afrique du Sud.
DESCRIPTION PALEONTOLOGIQUE DU MATERIEL
Famille Robustocyathidae Debrenne, 1964
Genre Stapicyathus Debrenne, 1964
1964 — Archaeocyathellus (Stapicyathus) Ford.—Debrenne: 127.
1970— Stapicyathus Debrenne.—Debrenne: 43.
1972—Stapicyathus Debrenne.— Hill: E 66.
Espéce-type: Archaeocyathus stapipora Taylor, 1910: 118.
Diagnose
Calices 4 2 murailles poreuses et cloisons radiales imperforées, sauf une
rangée de pores qui se combine avec l’unique rangée verticale de pores que la
muraille interne posséde en face de chaque cloison. La muraille externe est
simple et réguliérement poreuse.
Discussion
In Debrenne 19746.
Répartition
URSS, Extréme-Orient, Australie, Antarctique.
Stapicyathus incisus (Hill), 1965
Fig. la
1920—Thalamocyathus tubavallum Taylor.—Gordon: 168, pl. 1 (fig. 15).
1965 — Robustocyathus incisus Hill: 68-69, pl. 3 (figs 2-5).
1974b— Stapicyathus incisus (Hill).—Debrenne: 121, pl. 25 (fig. 3).
Holotype: British Museum (Natural History) S 8398.
Matériel étudié: 4 exemplaires, SAM-K4495 B.
Description
Espéce conique, atteignant de 5 4 6 mm de diamétre, avec un intervallum
de 1,2 mm et des cloisons distantes de 0,6 4 0,7 mm. Le coefficient pariétal
‘OL X ‘opessoAsuen odnod “qLI- S6rpVI-IN VS ‘ouueIqed snj2aj snyjoAooWDjoYT, “S{ “317A
"OZ X ‘apesioAsues) odnoo oun,p [1e19q ‘OI-V S6PYN-INVS ‘ouusIged sj2aj snyjodoowDjoY] “qt “SIy
‘OI X ‘anbijgo ayesiaasued adnod “q¢sI- S6PVS-WVS “CIIIH) Sms1ou1 snyjodoidojgs “ey “314
333
ARCHAEOCYATHA PROVENANT DES TILLITES DE DWYKA
334 ANNALS OF THE SOUTH AFRICAN MUSEUM
est de 3, les chambres interseptales de 1/2. La muraille externe, mal conservée,
a 4a 5 rangeées de pores par intersept (diametre environ 0,05 mm-—linteaux et
épaisseur 0,10 mm). Le pore de la muraille interne est large (0,27 mm) séparé
par des linteaux importants (0,20-0,25 mm) d’épaisseur 0,20 mm.
Affinités
Correspond en tous points aux mesures et coefficients des formes d’Ajax
Mine, de Weddel Sea et de Wichaway Nunatak.
Famille Cyclocyathellidae Zhuravleva, 1959
Thalamocyathus tectus Debrenne, 1973
Fig. 1b-c
1920—Thalamocyathus trachealis (Taylor) pars.—Gordon: pl 2 (figs 24-25).
1936—Bronchocyathus trachealis (Taylor) pars.—R. & J. Bedford: 25, pl. 26 (figs 100-102).
1955—Thalamocyathus trachealis (Taylor) pars. — Hill: 94-96, pl. 7 (fig. 2a-e).
1955 — ?Thalamocyathus sp.— Hill: 94-96, pl. 7 (figs 7a—b—8a-b).
1973 —Thalamocyathus tectus Debrenne: 11, figs 5B, 6-8, pl. 1 (figs 5-7), pl. 4 (fig. 2).
Holotype: Princeton University 86722 (165).
Matériel étudié: 6 calices, SAM—-K4495.
Description
Calices coniques a cloisons droites qui perdent leur porosité quand les
individus atteignent 7 mm de diamétre. Le coefficient pariétal est élevé (7), les
chambres interseptales ont un rapport des cétés de 1/4,8. La muraille externe
porte des tumuli peu nombreux (2 par intersept dans les calices jeunes, souvent
un seul ensuite). Leur paroi est mince (0,05 mm); ils forment des monticules
assez élevés (0,06 mm au-dessus de la muraille) qui couvrent une ouverture de
0,10 mm. Ces monticules peuvent étre interprétés comme pores-diaphragmes
turgescents (Debrenne 1973: 6, 11, fig. SA—B). La muraille interne est annulaire
avec une forme en auge, concave vers le haut; au milieu et sous cette gouttiere
se développe une caréne plus ou moins bien marquee. La largeur de l’anneau
est de 0,27 mm, la distance entre 2 anneaux, 0,16 mm. L’épaisseur moyenne de
la paroi est 0,06 mm. A la base, on peut observer des anneaux incomplets, ce
qui permet de rapprocher les stades jeunes du genre Sanarkocyathus Zhuravleva.
Il n’y a pas de planchers pectinés visibles.
Affinités
La distinction entre les tumuli (type Kijacyathus Zhuravleva ou Annulo-
cyathella V ologdin) et les pores diaphragmes turgescents a été exposée antérieure-
ment (Debrenne 1973: 6, 11). Il est actuellement difficile de se prononcer sur la
valeur systématique de ces saillies hémisphériques (vacuoles—bubbles dans
Hill 1955: 95). Actuellement une espéce, tectus, rattachée au moins provisoire-
ment au gentre Thalamocyathus a été établie (Debrenne 1973: 11). Elle présente
ARCHAEOCYATHA PROVENANT DES TILLITES DE DWYKA 335
une muraille externe avec capsules hémisphériques. Son coefficient pariétal est
plus faible que celui de trachealis (Debrenne 1973: 11).
Certaines formes décrites en Antarctique par Gordon et Hill et dans la
composition de l’espéce trachealis avaient déja été incluses dans tectus (Debrenne
1973). Les échantillons d’Afrique du Sud, par leurs caractéristiques anatomiques
et leurs coefficients font également partie de l’espéce.
?Famille Kijacyathidae Zhuravleva, 1964
Genre Flexanulus gen. nov.
Diagnose
Calices a 2 murailles et cloisons radiales peu perforées. La muraille externe
est formée d’un ensemble de tubes allongés, courbés avec ouvertures dans le
plan vertical. Au point de rebroussement un crochet recourbé vers le bas se
développe, fermant partiellement le tube immédiatement inférieur (Fig. 2).
Les cloisons s’appuient sur les parois latérales des tubes et forment avec elles
des parties squelettiques épaissies. La muraille interne est annulaire; les anneaux
sont en S trés allongés se recouvrant |’un |’autre.
Espéce-type: Flexanulus oosthuizeni sp. nov.
Affinités
Genre proche d’Annulocyathus Vologdin et de Kijacyathus Zhuravleva, il
en différe par la forme particuliére des tubes externes et des anneaux internes.
Flexanulus oosthuizeni sp. nov.
Figs 2-3a—b
Holotype: SAM-K4495 B-12a.
Matériel étudié: 4 calices.
Description
Calices coniques de petite taille (5,4 mm) traversés de cloisons peu perforées,
délimitant des chambres interseptales de rapport 1/5. Le coefficient pariétal est
de 5,7. Les canaux de la muraille externe, 2 par intersept, se développent en
s’appuyant sur les cloisons a l’intérieur de l’intervallum; leur diamétre est de
0,20 mm, I’€paisseur de leur paroi 0,01 mm et leur longueur 0,21 mm. Ils sont
formés d’une partie intervallaire en tube oblique vers |’extérieur et le haut,
puis se redressent brusquement presqu’a angle droit au niveau des ouvertures
externes. A ce point se développe un crochet recourbé vers le bas, symétrique
de la partie redressée par rapport a4 la paroi du tube (Fig. 3a—b). La muraille
interne est formée d’anneaux en S trés allongés et imbriqués: |’ouverture entre
les anneaux est de 0,16 mm, la hauteur des anneaux 1,15 mm, l’épaisseur de
leur paroi 0,04 mm. Les cloisons sont droites, peu poreuses, épaisses de 0,04 mm.
A ffinités
Une seule espéce connue actuellement.
336 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 2. Reconstitution des tubes de la
muraille externe de Flexanulus gen. nov.
Famille Erbocyathidae Vologdin & Zhuravleva, 1956
?Ladaecyathus sp.
Fig. 4a
1 exemplaire, SAM-K4495 A-9.
Description
Fragment d’un petit calice dont les murailles externe et interne paraissent
recouvertes d’une seconde enveloppe microporeuse. Les cloisons sont €paisses
onduleuses et parfois bifurquées.
Dimensions
Diamétre: 6,5 mm; intervallum: 1,65 mm; distance entre les cloisons
variable: en moyenne 0,6 mm; muraille externe principale, diamétre des pores:
32)
ARCHAEOCYATHA PROVENANT DES TILLITES DE DWYKA
cI x
“ST X
‘g7eulpnyisuo] odnoD “eZ]
-I S6PYS-WVS ““Aou ‘ds 39 “Wad IuaziInyjsoO SnjnuDXxeaLT “QE ‘SIq
‘gresioasues} odnoZD “e7[-d S6PVA-NVS “aou ‘ds 39 ‘UdS MazINyIsooO Snjnuvxa,y “eg “BIy
Be
ANNALS OF THE SOUTH AFRICAN MUSEUM
338
0Z X ‘(eploe,] & gnbeye o11es[e9) geurpnysuoy odnoD ‘¢[-€ S6PYM-INVS “ouUusIqed $7/0149,1D snuljzadoupjoy [ “Gp “314
‘Oz X ‘anbijqo ajesioasuen odnoD *6-Y S6hYM-INVS “ds snysodoavpvTj, “ep ‘SIq
ARCHAEOCYATHA PROVENANT DES TILLITES DE DWYKA 339
0,16 mm, linteaux: 0,05 mm, épaisseur: 0,10 mm; muraille interne, diamétre:
0,16 mm, linteaux: 0,08 mm, épaisseur: 0,21 mm, épaisseur des cloisons:
0,09 mm, pores: 0,10 mm.
Discussion
Les espéces rapportées par Hill (1955) a Ladaecyathus Zhuravleva, 1960,
sont représentées par des fragments d’exemplaires de grande taille (60 mm pour
L. pratti Hill, 35 mm pour fortiseptatus Hill). Il est difficile de rattacher le
fragment découvert ici a l’une ou l’autre espéce. D’autre part l’attribution au
genre Ladaecyathus de ces fragments aussi bien ceux décrits par Hill, que celui
dont il est question ici, est douteuse. La fagon dont se construisent les doubles
murailles et la relative irrégularité des cloisons font penser plutét 4 Metaldetes
ramulosus (R. & J. Bedford)—Debrenne 1974a: 226-227, fig. 24.
Famille Bronchocyathidae R. & J. Bedford, 1936
Thalamopectinus arterialis Debrenne, 1972
Fig. 4b
1936—Bronchocyathus trachealis (Taylor).—R. & J. Bedford: 25, pl. 25 (fig. 99C-D).
1955 —Thalamocyathus trachealis (Taylor).— Hill: 96, pl. 7 (fig. 3).
1973 — Thalamopectinus arterialis Debrenne: 8.
Materiel étudié: SAM-K4495 B-13.
Description
Calice cylindrique a cloisons radiales peu ou pas perforées et planchers
pectinés assez réguliérement répartis, distants d’un millimétre en moyenne. Le
diameétre atteint 5 mm avec un intervallum de 0,75 mm; l’écartement des cloisons
est de 0,15 mm, soit un coefficient intervallaire de 0,15 et un rapport de 1/5 pour
les chambres interseptales. La muraille externe a une porosité simple, elle est
formée d’une lame perforée réguliérement de pores arrondis disposés en quin-
conce, 3 rangées par intersept (diamétre: 0,06 mm, linteaux: 0,02 mm). La
muraille interne est annulaire avec une section en auge soulignée vers le bas
par une petite caréne; les anneaux sont distants de 0,1 mm, |’épaisseur de la
paroi est variable (de 0,1 4 0,2 mm) suivant l’endroit de la mesure. La largeur
totale de l’anneau est de 0,25 mm.
Affinités
Une étude récente sur la valeur systématique qui devait étre accordée aux
planchers pectinés (Debrenne et al. 1973) a établi qu’il était nécessaire de classer
en des genres distincts les formes avec et les formes sans planchers pectinés.
Le genre Thalamocyathus Gordon est réservé aux formes a cloisons peu poreuses,
le genre Thalamopectinus Debrenne aux formes a cloisons peu poreuses et
planchers pectinés.
340 ANNALS OF THE SOUTH AFRICAN MUSEUM
L’espéce découverte en Afrique du Sud est en tout point comparable a
celle figurée par Hill (1955, pl 7 fig. 3) (British Museum (Natural History)
58416); bien qu’elle ne signale pas de planchers pectinés cette structure a
été observee sur la lame et apparait méme sur la figuration citée.
Un exemplaire de la collection Bedford (Princeton University 248) est
trés voisin de celui étudié ici; il semble donc que les formes d’Antarctique et
d’Afrique du Sud entrent dans les limites de l’espéce arterialis Debrenne qui
aurait donc existé dans les trois régions.
Famille Anaptyctocyathidae Debrenne, 1969
Gentre Erugatocyathus Debrenne, 1969
Espéce-type: Coscinocyathus papillatus R. & W. R. Bedford, 1934: 3, fig. 12.
Diagnose
Coscinocyathe avec muraille externe 4 double porosité et muraille interne
protégée par des crochets trés recourbés recouvrant le pore de base.
Discussion
La seconde enveloppe microporeuse n’est pas indépendante de la muraille
principale. Elle est de type Erbocyathus (Debrenne 1973: 18) et non Tomocyathus.
Erugatocyathus scutatus (Hill), 1965
Fig. 5a—b
1965 — Torgaschinocyathus scutatus Hill: 104-105, pl. 8 (figs 1-4).
Holotype: British Museum (Natural History) S 8434.
Matériel étudié: 12 calices.
Description
Calices coniques ne dépassant pas 12 mm de diamétre. Un exemplaire
montre que |’animal débute par un stade a 2 murailles et intervallum vide sauf
quelques lames de tissu vésiculeux. A 2 mm de diamétre se forment les premiers
planchers poreux qui précédent l’apparition des cloisons. La muraille externe
est d’abord compacte et épaissie par du stéréoplasme. La double porosité est
acquise trés t6t, au niveau des premiers planchers, lorsque le stéréoplasme
disparait. Les individus les plus nombreux ont un diamétre de 5 a 7 mm avec
un intervallum de 1 a 1,5 mm. Les cloisons radiales sont assez espacées, formant
des chambres interseptales dont le rapport des c6tés varie de 1/1,25 a 1/2, tandis
que le coefficient pariétal s’établit entre 3 et 3,7.
La porosité de la muraille externe est souvent dificile 4 observer, la fossili-
sation étant souvent grossiére, et les structures souvent masquées par des algues
qui s’accolent aux calices. Cependant certains endroits plus favorables montrent
la double enveloppe. Les pores principaux ont un diamétre de 0,15 mm, des
linteaux de 0,10 mm; l’épaisseur totale de la muraille est de 0,10 mm. La muraille
341
ARCHAEOCYATHA PROVENANT DES TILLITES DE DWYKA
0
"CL
Tx *
x ‘ayeuIpnyisuo] sdnoD “e¢-q S6rpyaI-NVS
gjesioasued) odnoD */-€ S6pVI-NVS “(I[MH) Sniwinas snyyodoojos8naq °q¢ “314
“(I[NH) Snivinos snyjodoojosnAq “eG “317
342 ANNALS OF THE SOUTH AFRICAN MUSEUM
interne comporte 2, plus rarement 3 rangées de pores en quinconce par intersept.
Le développement des crochets courbés vers le haut et complétés par des épines
vers le bas est variable: complets et bien marqués dans les faibles diamétres,
ils diminuent d’importance et ont presque disparu dans le calice de 12 mm.
Les cloisons sont trés poreuses: 5 a 8 pores en rangées réguliéres (diamétre
0,05 mm, linteaux: 0,16 mm, épaisseur: 0,04 mm). Les planchers ont des pores
de méme diametre, mais plus denses (8 par intersept, diamétre: 0,05 mm,
linteaux: 0,04 mm).
Affinités
L’observation personnelle des collections du British Museum (Natural
History) S 8433-S 8434 (lames) nous a permis d’observer la présence d’une
double muraille externe, souvent masquée par des algues et décrite par Hill
(1965) comme ‘thin projecting collar’. Les plaques recourbées au-dessus des
pores de la muraille interne sont tout a fait semblables a celles du type de
Erugatocyathus Debrenne. Les mesures et coefficients de E. scutatus (Hill)
correspondent a ceux des échantillons étudiés ici qui doivent donc étre rapportés
a cette espéce.
E. scutatus (Hill) différe de E£. papillatus (R. & W. R. Bedford) par une
plus grande minceur des éléments squelettiques, une plus faible porosité des
cloisons et des chambres interseptales différentes (1/2,5 contre 1/5).
?Erugatocyathus sp.
Fig. 6a
Un exemplaire: SAM-K4495 B-17a.
Description
Un calice de 5 mm de diamétre, ayant un coefficient pariétal de 3,6 et des
chambres interseptales de 1/2,7 de cdté présente un seul pore recouvert d’une
papille par intersept a la muraille interne.
Discussion
La présence d’un seul pore, caractére généralement considéré comme
important pour la systématique, éloigne cet exemplaire des E. scutatus types.
Cependant en l’absence de sections supplémentaires dans cet exemplaire, il
n’est pas possible de savoir si la figure observée traduit un trait constant ou
seulement un accident sporadique.
Famille Porocoscinidae Debrenne 1964
Genre Statanulocyathus gen. nov.
Espéce-type du genre: Statanulocyathus oosthuizeni sp. nov.
Diagnose
Calices 4 2 murailles, cloisons et planchers poreux. La muraille externe a
des pores recouverts de bractées; la muraille interne est principalement cons-
ARCHAEOCYATHA PROVENANT DES TILLITES DE DWYKA
Oo
O
343
Fig. 6a. Erugatocyathus sp., SAM-K4495 B-17a. Coupe transversale, x 20.
Fig. 6b. ?Didymocyathus sp., SAM-K4495 A-13a.
344 ANNALS OF THE SOUTH AFRICAN MUSEUM
tituée par des pores recouverts par des bractées, un par intersept; ces bractées
se soudent périodiquement pour former un anneau complet, au niveau de
chaque plancher.
Les cloisons et les planchers sont normalement poreux.
Discussion
Il est probable que nous ayions affaire a des faux anneaux formés par la
fusion et le développement plus important des bractées qui soulignent les pores
internes. Ce caractére a précédemment été décrit dans différents genres
(Cadniacyathus R. & J. Bedford, Tennericyathus Rozanov, Denaecyathus
Zhuravleva). L’auteur (Debrenne, 19745) ne considére pas que ces formations
soient de vrais anneaux. La forme découverte et décrite ici montre que ces
fusions en anneaux peuvent étre périodiques et que la muraille principale reste
une muraille 4 canaux et bractées. C’est pourquoi ce nouveau genre est inclus
dans la famille Porocoscinidae Debrenne et non pas Sigmocoscinidae R. & J.
Bedford.
Statanulocyathus oosthuizeni sp. nov.
Fig. 7a—b
Holotype: SAM-K4495 A-7.
Matériel étudié: 1 exemplaire.
Description
Petits calices de 5 mm de diameétre, avec un intervallum de 1 mm de large,
traversé de cloisons distantes entre elles de 0,21 mm et de planchers plus irrégu-
liers répartis (de 0,54 mm a 1 mm). Le coefficient pariétal est de 12. La muraille
externe a 2 rangées de pores par intersept, d’ouverture 0,12 mm, recouverts de
bractées recourbées formant des cupules de 0,04 mm de hauteur au-dessus de
la muraille dont l’épaisseur totale atteint 0,08 mm. La muraille interne comporte
une rangée de pores par intersept, aplatis dans le sens de la hauteur et alignés hori-
zontalement, ils sont soulignés de bractées. L’ensemble se présente comme un
court tuyau en V, d’ouverture 0,12 mm et de largeur 0,16 mm; l’épaisseur du
squelette de ces éléments ne dépasse pas 0,03 mm. Au niveau de chaque plancher
se construit une formation annulaire large et épaisse (0,26 mm) recourbée vers
le haut.
Il semble que les systeémes poreux des cloisons et des planchers soient les
mémes (pores: 0,03 mm, linteaux: 0,07 mm).
Famille Sigmocyathidae Krasnopeeva, 1955
Genre Didymocyathus Debrenne & Rozanov, 1972
Espéce-type: Didymocyathus hillae Debrenne & Rozanov, 1972.
Diagnose
Calices a 2 murailles annulaires, cloisons et planchers poreux.
345
ARCHAEOCYATHA PROVENANT DES TILLITES DE DWYKA
Ol x
‘so[[rernu sop [eo "L-V S6bY I-INVS “AOU °
‘gnbijgo oyesioAsued} odnoD °*L-V S6vbyA-WVS
ds 30 ‘uo8 muazinyjsoo snyjodoojnuDjVIS “QL ‘34
“Au ‘ds Jo “Uod muaziInyjsoo snyjvADO[NUDIDIF “VL “BI
346 ANNALS OF THE SOUTH AFRICAN MUSEUM
Affinités
Différe de Sigmocyathus par la porosité des murailles.
Répartition
Jusqu’a maintenant seulement Ajax Mine (Australie).
Afrique du Sud?
?Didymocyathus sp.
Fig. 6b
Un exemplaire: SAM-K4495 A-13a.
Description
Fragment d’une coupe transversale oblique permettant d’observer l’amorce
des anneaux des murailles, les cloisons et les planchers poreux.
Diamétre: 10 mm environ, intervallum: 1,5 mm, coefficient intervallaire:
0,15, interseptum: 0,65 mm, chambres interseptales: 1/2,3.
La largeur des murailles est de 0,4 mm, louverture entre les anneaux de
0,01 mm, |’épaisseur de l’anneau varie, elle s’amincit vers la terminaison libre.
Les cloisons sont perforées de 6 rangées de pores, 4 pores des planchers apparais-
sent dans la largeur de l’intersept.
Discussion
La petitesse et la rareté du matériel ne nous a pas permis de mettre indubi-
tablement en évidence la présence d’anneaux externes, tels qu’ils sont suggérés
par la coupe observée. Cependant les structures visibles sont trés nettement en
faveur de cette hypothése. Les formes de Didymocyathus connues a ce jour sont
de grande taille (jusqu’a 62 mm); les calices jeunes ou plus petits ne sont pas
connus ce qui ne permet pas d’identifier la forme d’Afrique du Sud aux espéces
d’Australie.
Famille Archaeopharetridae Debrenne, 1970
Genre Archaeopharetra R. & W. R. Bedford, 1936
Espéce-type: Archaeopharetra typica R. & W. R. Bedford, 1936.
Diagnose
Petits calices 4 muraille externe imperforée, muraille interne assez tardive-
ment différenciée. L’intervallum comporte des éléments squelettiques irrégu-
liérement disposés et du tissu vésiculeux en lames horizontales.
Affinités
Proche de Bicyathus Vologdin qui pourrait étre un synonyme récent
d’Archaeopharetra Bedford.
ARCHAEOCYATHA PROVENANT DES TILLITES DE DWYKA 347
Archaeopharetra cf. typica R. & W. R. Bedford, 1936
Fig. 8a
1936—Archaeopharetra typica R. & W. R. Bedford: 17, pl. 17 (fig. 75).
1937—Archaeopharetra typica R. & W. R. Bedford.—R. & J. Bedford: 30-31, pl. 29
(fig. 120A-—B).
1970— Archaeopharetra typica R. & W. R. Bedford.—Debrenne: 29.
1974a— Archaeopharetra typica R. & W. R. Bedford. —Debrenne: 195-196, fig. 3a—b.
Holotype: South Australian Museum P 969.
Matériel étudié: un exemplaire.
Description
Petit calice (4 mm) a cavité centrale étroite (0,5 mm). L’épaisseur de la
muraille externe (0,15 mm) et celle des barres intervallaires (0,10 mm) sont
inférieures a celle de l’espéce australienne.
Famille Chouberticyathidae Debrenne, 1974
Genre Chouberticyathus Debrenne, 1964
Espeéce-type: Chouberticyathus clatratus Debrenne, 1964: 208, pl. 32 (figs 1-7).
Diagnose
Calices a muraille externe compacte, striée horizontalement, muraille
interne simplement poreuse. L’intervallum est traversé de barres cylindriques
horizontales disposées le plus souvent radialement et reli¢es par des barres
obliques horizontales ou verticales. Tissu vésiculeux parfois présent.
Répartition
Maroc, Espagne, Australie.
Chouberticyathus cf. fragilis (R. & W. R. Bedford), 1936
Fig. 8b
1936 — Dictyocyathus fragilis R. & W. R. Bedford: 13, pl. 11 (fig. 57).
1964— Chouberticyathus fragilis (R. & W. R. Bedford). —Debrenne: 208.
1974a— ?Chouberticyathus fragilis (R. & W. R. Bedford).—Debrenne: 192-193, fig. 2.
Holotype: South Australian Museum P. 945 (59).
Matériel étudié: 4 calices.
Description
Petits calices de 4 mm de diamétre, avec intervallum de 0,7-0,8 mm traversé
de barres cylindriques horizontales radiales. La muraille externe compacte est
épaisse et présente une structure feuilletée (0,20 mm). La muraille interne est
grossi¢rement poreuse (diamétre 0,3 mm).
ANNALS OF THE SOUTH AFRICAN MUSEUM
348
‘OL x “onbiqo efeurpnysuo] ednoD “T9-Vv S6h/A-INVS ‘P1OJPOM “A “MA 79”
OZ x “onbi[qgo afeurpnyisuo] adnoD “es |-g S6ryM-WVS ‘PIOJPOM “AM
U SUISdAf “JO snywdo1saqnoy7D
“YU Va1dd1 “Jo vasvydoavyoap
q8 “314
eg “S14
ARCHAEOCYATHA PROVENANT DES TILLITES DE DWYKA 349
Discussion
Par la taille et la répartition des éléments squelettiques cette forme se
rapproche de Il’espéce australienne C. fragilis (R. & W. R. Bedford). L’attribu-
tion générique dans les 2 cas est faite avec réserve, la rareté et la petitesse du
matériel ne permettant pas de mettre en évidence les relations des structures
intervallaires.
Répartition
Australie du Sud, Afrique du Sud.
Famille Protopharetridae Vologdin, 1957
Genre Protopharetra Bornemann, 1884
Espéce-type: (Simon 1939) Protopharetra polymorpha Bornemann, 1884 (1887,
pl. 5 (fig. 4)).
Diagnose
Calices souvent ramifiés, parfois solitaires. Muraille externe souvent
imperforée ou, dans les grandes formes, constituée par l’ouverture du réseau
intervallaire recouvert d’un pellis sans pores visibles. La muraille interne est
une simple ouverture du réseau intertaenial. L’intervallum est rempli de plaquettes
branchues relic¢es entre elles par leurs branches ou par des tigelles en position
de synapticules. Elles peuvent s’orienter d’une fagon privilégiée en pseudo-
cloisons, surtout vers la muraille interne. Le tissu vésiculeux est abondant.
Répartition
Mondiale.
Protopharetra densa Bornemann, 1887
Fig. 9a
1887—Protopharetra densa Bornemann: 48, pl. 8 (figs 6c—7b-8).
1887 —Protopharetra polymorpha Bornemann: pl. 5 (fig. 5).
1940a—Protopharetra laqueata Vologdin: 40—42, fig. 7.
21959 — Syringocnema colevilensis Greggs: 72-73, pl. 13 (figs 5-6).
1967—Protopharetra densa Bornemann.— Zhuravleva et al.: 90-91, pl. 39 (figs 5-6).
Holotype: \lectotype Bornemann, 1887, pl. 8 (fig. 8).
Matériel étudié: 1 échantillon, SAM—K4495 A-3.
Description
Petits calices de 4 mm de diamétre a cavité centrale étroite (1 mm) mais libre
d’éléments squelettiques et de tissu vésiculeux.
La muraille externe est compacte et porte de courts bourgeonnements.
L’intervallum est rempli de plaquettes fortement liées en taeniae denses (distantes
radialement de 0,3 a 0,4 mm) épaisses de 0,15 mm. La muraille interne a une
ouverture par espace intertaenial de 0,2 mm.
ANNALS OF THE SOUTH AFRICAN MUSEUM
350
“gyes1oAsuvs} odnos op yuowselyz “7-G C6PVYH-NVS “(UOpIOD) vn dasianod vAjasDYdOJOAd{, °96 “Bt
‘Ol x ‘anbifqo adnod *,eZI-V S6bbA-WYVS “UIPsOjOA DIDAADIIPUDAS DAJIADYAOJOAT *Q6 “SIA
"OL X ‘ayessoasues3 9dNOD “LE-V S6PPA-INVS “UURlUsUIOg DsSuap DAJIADYAOJOAT *C6 “314
O
ARCHAEOCYATHA PROVENANT DES TILLITES DE DWYKA 351
Affinités |
Trés proche par la taille et les coefficients de la forme type.
Répartition
Sardaigne; URSS: Mongolie, Tuva, Altai Sajan; Maroc; ?USA.
Protopharetra grandicaveata Vologdin, 1940
Fig. 9b
1940b—Protopharetra grandicaveata Vologdin: 42, pl. 3 (figs 1-2).
1960—Protopharetra grandicaveata Vologdin. — Zhuravleva et al.: 139, pl. 10 (figs 7-8).
1964—Protopharetra grandicaveata Vologdin.—Repina et al.: 239, pl. 26 (figs 5-6).
1964—Protopharetra aff. grandicaveata Vologdin.—Debrenne: 216, pl. 32 (figs 8-9).
Holotype non précisé.
Matériel étudié: 5 échantillons, SAM—-K4495 A-12a-a!, B-SaT, A-1, B-15a.
Description
Calices solitaires de forme conique irrégulicre de 6 4 8 mm de diametre;
la cavité centrale est peu importante (1,5 mm maximum) souvent traversée de
lames de tissu vésiculeux. La muraille externe est compacte, épaisse de 0,1 mm,
sans pores visibles. L’intervallum, assez large, est rempli de plaquettes rares,
grossiérement orientées verticalement et radialement avec quelques liaisons
tangentielles. Leur épaisseur est du méme ordre que celle de la muraille externe
(0,08 mm—0,1 mm). La distance moyenne entre les plaquettes est de 0,4-0,5 mm.
La muraille interne est simple, avec des ouvertures de 0,25 mm de diamétre.
Le tissu vésiculeux est abondant et traverse l’intervallum et la cavité centrale.
Affinités
Tout a fait semblable aux formes de Protopharetra a rares éléments inter-
vallaires groupées dans l’espéce grandicaveata Vologdin.
Répartition
URSS: Salair, Sajan oriental, Tuva, Kuzneck-Alatau; Maroc; Afrique
du Sud.
?Protopharetra pauciseptata (Gordon), 1920
Fig. 9c
1920 — Archaeocyathus pauciseptatus Gordon: 687, pl. 6 (figs 63-64).
Holotype: King’s College S 148.
Materiel étudié: 1 exemplaire.
Description
Petits calices 4 taeniae presque droits vers la muraille interne et dissociés
vers la muraille externe. Le diamétre atteint 5 mm avec un intervallum de 1,2 mm.
La muraille externe, mal conservée par places, parait compacte. La distance
352 ANNALS OF THE SOUTH AFRICAN MUSEUM
entre les pseudo-cloisons est de 0,4 mm. La muraille interne est simple, a un
seul pore par espace intertaenial (diametre 0,4 mm).
Le tissu vésiculeux est peu abondant ici.
Affinités
Il est difficile de déterminer cet unique calice. La présence de taeniae radiales
presque completes éloigne cette forme des Protopharetra typiques; le faible
développement du tissu vésiculeux est la seule différence marquante avec
l’exemplaire décrit par Gordon, que |’on doit également rapporter avec doute
au genre Protopharetra. Sur des formes de si petite taille il est difficile d’étre
parfaitement affirmatif quant a leur position systématique.
Répartition
Antarctique (Weddel Sea); Afrique du Sud.
?Famille Flindersicyathidae R. & J. Bedford, 1939
Genre Andalusicyathus Perejon MS
1939 — Archaeocyathellus (Archaeofungia) Simon: 76.
1964 — Spirocyathella Vologdin.—Debrenne: 137.
Espéce-type: Archaeocyathellus (Archaeofungia) andalusicus Simon, 1939,
Cordoue, Las Ermitas (Espagne.)
Diagnose (d’aprés Perejon, Manuscrit)
‘Calices solitaires ou coloniaux de forme cylindrique ou légérement conique, —
avec expansion basale pour la fixation.
Muraille externe simple avec 3 rangées longitudinales de pores par intersept,
pouvant présenter de légeres ondulations longitudinales. Intervallum avec
septes (ou pseudo-septes) a larges pores généralement allongés dans le sens
longitudinal et réunis par des synapticules qui se situent au voisinage de la
muraille externe qui semblerait avoir un pore par intersept.
Il existe du tissu vésiculeux localisé dans des zones plus restreintes et qui
détermine une modification de la fossilisation des éléments squelettiques et
Vapparition d’un pellis.’
L’auteur consideére en outre que les espéces (Spirocyathus) latus Vologdin et
(Spirocyathus) extremus Vologdin tombent en synonymie avec andalusicus Simon.
Répartition
URSS: Sanachtygkol; Espagne: Las Ermitas; Afrique du Sud.
Andalusicyathus cooperi sp. nov.
Fig. 10
Holotype: SAM-K4495 A-S.
Description
Calices a large cavité centrale et intervallum construit par des pseudo-
cloisons largement perforées reliées par des synapticules et quelques lames de
393
ARCHAEOCYATHA PROVENANT DES TILLITES DE DWYKA
0
IT x
G
anbiyqo syesiaAsuesy ednoD
S
-“V S6bbx
“WVS
“AOU
ds t4adoo2 snyiodoisnjppuy
Ol
rd |
KY
bea
354 ANNALS OF THE SOUTH AFRICAN MUSEUM
tissu vésiculeux. Le diametre est de 10 a 11 mm, l’intervallum de 2,5 mm. La
muraille externe est perforée de 2 pores par espace intertaenial (diamétre;
0,10 mm, linteaux: 0,10 mm) souvent recouverte vers l’extérieur d’une membrane
de méme nature que le tissu vésiculeux. La muraille interne est indépendante
du réseau intervallaire et comporte 2 pores en quinconce entre 2 pseudo-septes
(diamétre; 0,21 mm, linteaux: 0,05 mm) pouvant porter de courtes épines.
L’espace entre 2 pseudo-septes est de 0,5 mm., I’épaisseur des plaquettes de
0,08 mm.
Affinités
Cloisons moins rapprochées et cavité centrale plus large que dans andulusi-
cus et tissu vésiculeux plus abondant.
Ordre des SYRINGOCNEMIDIDA
Définition: Intervallum a loculi prismatiques radiaux poreux.
Discussion
La définition de tubes poreux intervallaires, de section polygonale et de
disposition radiaire, n’a jusqu’ici pas été donnée avec une precision suffisante
et bien souvent, on a confondu sous ce terme des constructions de types
différents.
En effet, l’interprétation de cloisons ondulées reliées par des synapticules
aplaties (type Pycnoidocyathus decipiens) comme des tubes quadratiques a
souvent été proposée. Ces ‘tubes’ different essentiellement de ceux des Syringoc-
nemidida par leur forme (section quadratique), leur porosité (un pore par coté),
leur orientation (oblique vers l’extérieur et le haut, trés rarement subhorizontale).
Enfin, et c’est l’argument majeur, ces tubes évoluent plus ou moins rapidement
au fur et a mesure de la croissance de l’individu en pseudo-cloisons radiales
planes (type Pycnoidocyathus synapticulosus). L’évolution des tubes prismatiques
des Syringocnemidida en pseudo-cloisons radiales n’est jamais observeée.
C’est pourquoi il faut reprendre avec soin l’étude de la composition de cet
ordre:
Le genre Syringsella Krasnopeeva, 1961 est problement un synonyme récent
d’ Archaeofungia Taylor, 1910, sous réserve de l’examen de la porosité des cloisons.
Le genre Batenevia Krasnopeeva, 1961 est proche de Flindersicoscinus Debrenne,
1970 dont il différe par l’acquisition d’une muraille externe 4 double porosite.
Le genre Tubocyathus Vologdin, 1940 est 4 rapprocher des Pycnoidocyathidae.
Malheureusement sa position systématique a l’intérieur du groupe ne peut
étre précisée actuellement, la constitution des murailles n’étant pas connue en
détail. D’aprés la figuration il y aurait peut-étre une enveloppe microporeuse
externe et interne.
Beticocyathus Simon, 1939, genre par ailleurs caduc car les murailles ne sont
ARCHAEOCYATHA PROVENANT DES TILLITES DE DWYKA 355
pas conservées, n’est pas non plus une forme a loculi prismatiques, mais a
taenia, pseudo-cloisons ondulées et synapticules.
Syringocyathus Vologdin, 1940 a fait l’objet d’une étude particuliére par I. T.
Zhuravleva (1960: 42-46) qui considére ce genre comme appartenant aux
Regulares; l’intervallum comporte des cloisons radiales réguli¢rement perforées
reliées non par des synapticules simples, mais par des plaquettes de liaison
perforées, délimitant des loges verticales incompletes.
Par contre doivent étre maintenus dans l’ordre des Syringocnemidida les
familles et genres suivants:
Famille des Syringocoscinidae Vologdin & Jasmir, 1969
muraille externe simple
“ooops qhiesre Sttee Syringocoscinus V ologdin-Jasmir
Famille des Pseudosyringocnemididae fam. nov.
muraille externe ? simple ou 1 tumulus | Pseudosyringocnema
muraille interne: ? un canal oblique h Handfield, 1971
1 pore par cété du tube intervallaire
Famille des Fragilicyathidae fam. nov.
ill 2
murai e externe a canaux horizontaux \_ Frazilicyathus Beljacva, 1969
muraille interne 4 canaux coudés o6
tubes a plusieurs pores par cdété
Famille des Syringocnemididae Taylor (Debrenne emend.)
muraille externe a crible multiperforé
ae a : iat
muraille interne: un canal coudé hy POOR NOE Say OE
Famille Pseudosyringocnemididae fam. nov.
Genre Pseudosyringocnema Handfield, 1971
Espéce-type: Pseudosyringocnema uniporus Handfield, 1971: 76, pl. 15 (figs 3a—c,
4-5).
Diagnose originale
‘The cups may be solitary or colonial with the intervallum containing
hexagonal tubules. The tubules sides have only a single row of pores. The inner
wall has oblique ‘S’ shaped pores tubes, 2 or3 per tubule in vertical arrangement.’
Discussion
L’examen des figurations plate 15, figures 3a, 3c suggére le fait que les
tubules intervallaires se recourbent vers le bas, comme c’est le cas dans toutes
les formes 4 tubes hexagonaux, et se rétrécissent jusqu’a se raccorder a un seul
canal interne et non 2 ou 3; cette hypothése est confirmée par la section oblique
figure 3a ot le raccord de chaque tube intervallaire se fait avec un canal corres-
pondant de la muraille interne.
La diagnose proposée doit étre ainsi modifiée: calices dont |’intervallum
356 ANNALS OF THE SOUTH AFRICAN MUSEUM
est constitué de tubules dont les cdtés sont perforés d’une seule rangée de pores.
Muraille externe mal définie, probablement un pore au sommet d’un tumulus.
Muraille interne un canal par tubule, dirigé vers le centre et le haut de la cavité
centrale.
Affinités
Differe de Syringocnema par la porosité des tubes intervallaires. Se rapproche
de certaines especes de Pycnoidocyathus mais ne forme jamais de pseudo-cloisons
radiales.
Pseudosyringocnema cf. gracilis Gordon, 1920
Fig. lla
1920—Syringocnema gracilis Gordon: 699, pl. 4 (fig. 43).
1965 — Syringocnema gracilis Gordon. Hill: 136, pl. 11 (figs 17-18).
Holotype: King’s College, Londres; syntypes S 108—S 112—S 115—S 118—
S 120—S 121.
Matériel étudié: 4 calices.
Description
Petits calices de 6 4 7 mm de diamétre avec un intervallum de 2 mm de
large. Les tubes hexagonaux sont perforés d’une rangée de pores de 0,2 mm
de diamétre. Les alvéoles ont une ouverture de 0,5 mm dans leur partie médiane.
Les tubes se rétrécissent et se raccordent aux pores de la muraille interne dont
le diamétre est de 0,2 mm. Les tubes sont fermés extérieurement par une muraille
bombée au sommet de laquelle se trouve probablement un pore, mais cette
ouverture n’a pas été observée.
Discussion
La forme décrite ici différe de l’espéce-type et des échantillons rapportés
a l’espéce par Hill (1965) par un intervallum plus large pour un méme diameétre
(2 mm contre 1,5 mm) mais les autres éléments squelettiques ont des mesures
comparables, notamment les tubes intervallaires.
?Pseudosyringocnema cf. uniserialis (Hill), 1965
Fig. 11b-c
1965 — Flindersicyathus uniserialis Hill: 123, pl. 11 (figs 1-2).
Holotype: British Museum (Natural History) S 8441.
Matériel étudié:; 3 fragments.
Description
Fragment d’un calice conique assez haut (12 mm) avec un intervallum
relativement étroit (2,5 mm pour un diamétre de 5 mm). La muraille externe
n’est pas conservée. Les alvéoles de l’intervallum ont un diamétre de 0,4 mm
357
ARCHAEOCYATHA PROVENANT DES TILLITES DE DWYKA
Ol xX “wnyyeArejzur | sIaAvI} & OdNOD ‘eE[-V S6PVH-INVS “(I[NH) Soldasiun “Jo pluausosulddsopnasd
‘Or x
‘Ol
¢
Sv
.
gyeurpnyisuoy odnoD *;e¢I-V S6rhpA-WVS
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358 ANNALS OF THE SOUTH AFRICAN MUSEUM
L’épaisseur des parois est de 0,07 mm percée de pores d’un diamétre de 0,15 mm.
La muraille interne a un court tube d’ouverture 0,20 mm. |
Discussion
L’appartenance de l’espéce wniserialis au genre Flindersicyathus comme le
démontre Hill (1965: 123-124) repose sur la forme quadratique et non hexa-
gonale des tubes intervallaires. Structure difficile 4 mettre en évidence. La forme
étudiée actuellement bien que légérement plus petite, peut étre comparée a
uniserialis Hill; elle parait avoir des loculi hexagonaux et non quadratiques.
Mais le matériel est trop rare et trop fragmentaire pour que l’on puisse décider
de trancher ici le délicat probleme des formes 4 loculi prismatiques dans
V’intervallum.
CONCLUSIONS
Les résultats de l’étude paléontologique sont reportés dans le Tableau 1.
Sur les 18 espéces décrites 10 sont communes avec |’Antarctique, 8 avec
Australie et 5 avec les deux.
Ce tableau appelle un certain nombre de précisions:
(1) les espéces les plus fréquentes sont Erugatocyathus scutatus (Hill), Pseudo-
syringocnema cf. gracilis (Gordon), Pseudosyringocnema cf. uniserialis (Hill),
espéces connues dans |’Antarctique—et Thalamocyathus tectus, connue en
Antarctique et en Australie.
(2) Les espéces des genres Protopharetra et Archaeopharetra sont cosmopolites
et ne peuvent apporter d’arguments décisifs dans ce cas.
(3) La présence de formes a planchers pectinés (Thalamopectinus Debrenne)
est trés importante; si l’on suit l’hypothése émise récemment (Debrenne et al
1973: 33, fig. 4) selon laquelle les planchers pectinés seraient liés aux climats
équatoriaux, les blocs erratiques de Dwyka viendraient de la zone équatoriale
Australo-Antarctique.
(4) La découverte d’espéces et méme de deux genres nouveaux peut s’expliquer,
non par l’endémicité de ces taxons, mais par le caractére sporadique et incomplet
des récoltes faites en Antarctique (dragage, moraines, prises en place limitées
en nombre) et par la rareté actuelle des localités étudiées en Australie, dont la
faune d’Archéocyathes connue est, en fait, restreinte 4 la région d’Ajax Mine.
Il est probable que ces nouvelles formes, rares (1 ou 2 exemplaires découverts
ici) pourront étre mises en évidence plus tard dans les régions antarctiques et
australiennes. On peut affirmer, grace a la similitude des espéces que les blocs
erratiques des tillites de Dwyka proviennent de la région antarctique.
L’hypothése d’un continent austral, futur Gondwanaland, en bordure
duquel se développaient les Archéocyathes, peut trouver ici une justification
supplémentaire.
ARCHAEOCYATHA PROVENANT DES TILLITES DE DWYKA 359
Stapicyathus incisus (Hill)
Thalamocyathus tectus Debrenne
Flexanulus oosthuizeni sp. nov. .
?Ladaecyathus .
Thalamopectinus arterialis
Debrenne
Erugatocyathus scutatus (Hill)
?Erugatocyathus sp.
Statanulocyathus oosthuizeni sp.
nov. .
? Didymocyathus sp.
Archaeopharetra cf. typica (Born.)
Chouberticyathus cf. fragilis (Bed.)
Protopharetra polymorpha
Bornemann . :
Protopharetra densa Bornemann .
Protopharetra grandicaveata
Vologdin.
?Protopharetra pauciseptata
Gordon .
Andalusicyathus cooperi sp. nov. .
Pseudosyringocnema cf. gracilis
(Gordon)
Pseudosyringocnema cf. uniserialis
(Hill)
Tableau J
Australie | Antarctique | Afrique du Sud | Autres localités
—$_—— EE
= == =
aie ae ==
O O ==
+ += 5
= == 2?URSS
== ==
ee oe | i
O O =F
== O ==
a O + 2URSS
a= O ==
aie ai mondiale
? ~ Sardaigne,
Maroc, URSS,
2?USA
@) O == URSS, Maroc
O a= ==
O © ==
O =e a=
O = ==
360 ANNALS OF THE SOUTH AFRICAN MUSEUM
REMERCIEMENTS
Nous exprimons notre gratitude 4 M. R. Oosthuizen qui a découvert ce
matériel, a M. M. Cooper et au South African Museum qui a bien voulu nous
en confier |’étude, et au Dr W. H. Ball, British Museum (Natural History) tant
pour son hospitalité dans ses locaux ol nous avons eu accés aux collections
d’Australie et d Antarctique, que par les nombreux préts temporaires qu'il a
consentis pour les comparaisons et révision des faunes.
REFERENCES
BEDFORD, R. & BEDFORD, J. 1936. Further notes on Cyathospongia (Archaeocyathi) and other
organisms from the Lower Cambrian of Beltana, South Australia.—Mem. Kyancutta
Mus. 3: 21-26.
BEDFORD, R. & BEDFORD, J. 1937. Further notes on Archaeos (Pleospongia) from the Lower
Cambrian of South Australia.— Mem. Kyancutta Mus. 4: 27-38.
BEDFORD, R. & BEDFORD, W. R. 1934. New species of Archaeocyathinae and other organisms
from the Lower Cambrian of Beliana, South Australia.— Mem. Kyancutta Mus. 1: 1-7.
BORNEMANN, J. G. 1887. Die Versteinerungen des Cambrischen Schichten-systems der Insel
Sardinien nebst vergleichenden Untersuchungen tiber analoge Vorkommisse aus andern
Landern.— Nova Acta Acad. Caesar. Leop. Carol. 51: 1-148.
DEBRENNE, F. 1964. Archaeocyatha. Contribution a l’étude des faunes cambriennes du Maroc,
de Sardaigne et de France.— Notes Mém Serv. Mine. Carte géol. Maroc. 179: 1-265,
266-371.
DEBRENNE, F. 1970. A revision of Australian genera of Archaeocyatha.—Trans. R. Soc. S.
Aust. 94: 21-48.
DEBRENNE, F. 1973. Modifications de la porosité premiére de la muraille externe chez les
Archéocyathes Réguliers.— Annls Paléont. (Invert.) 59: 1-24.
DEBRENNE, F. 1974a. Les Archéocyathes Irréguliers d’Ajax Mine (Cambrien inférieur, Australie
du Sud).— Bull. Mus. nat. Hist. nat., Paris (3) 195 (Sci. Terre 33): 185-258.
DEBRENNE, F. 19746. Anatomie et systématique des Archéocyathes Réguliers sans plancher
d’Ajax Mine (Cambrien inférieur, Australie du Sud).—Géobios 7: 91-138.
DEBRENNE, F. & ROZANOV, A. Ju. 1972. O Kol’cevykh structurakh naruzhnykh stenok
Arkheociat. [Sur les structures annulaires de la muraille externe des Archéocyathes.]
In: ZHURAVLEVA, I. T., ed. Problemy paleontologii i biostratigraphii nizhnego Kembrija
Sibiri i Dalgeno Vostoka: 235-237, pls. 42-44. Moskva: Nauka, Akademiya Nauk SSSR,
Sibirskoe Otdelenie.
DEBRENNE, F., ZHURAVLEVA, I. T. & ROZANOV, A. Ju. 1973. Grebentchatije dnishcha u
Arkheociat i ikh sistematitcheskoe znatchenie. [Planchers pectinés chez les Archéocyathes
et leur signification systématique.] In: ZHURAVLEVA, I. T., ed. Problemy paleontologii i
biostratigraphii nizhnego Kembrija Sibiri i Dalnego Vostoka: 33, 38, 6 figs. Novosibirsk:
Nauka, Akademiya Nauk SSSR, Sibirskoe Otdelenie.
Gorpon, W. T. 1920. Scottish National Antarctic Expedition, 1902-1904: Cambrian organic
remains from a dredging in the Weddell Sea.— Trans. R. Soc. Edinb. 52: 681-714.
Grecos, R. C. 1959. Archaeocyatha from the Colville and Salmo areas of Washington and
British Columbia. —J. Paleont. 33: 63-75.
HANDFIELD, R. C. 1971. Archaeocyatha from the Mackenzie and Cassiar Mountains, North-
west Territories, Yukon Territory and British Columbia.—Bull. geol. Surv. Can, 201:
i-v, 1-119.
Hit, D. 1955. Contribution to the correlation and fauna of the Permian in Australia and
New Zealand.—J. geol. Soc. Aust. 2: 83-107.
Hit, D. 1965. Archaeocyatha from Antarctica and a review of the phylum.—Scient. Rep.
transantarct. Exped. 10: 1-151.
Hit, D. 1972. Archaeocyatha. In: TEICHERT, C., ed. Treatise on invertebrate paleontology.
Part E. 2nd ed. 1. Boulder: Geological Society of America; Lawrence: University of
Kansas.
ARCHAEOCYATHA PROVENANT DES TILLITES DE DWYKA 36]
PEREJON-RINCON, A. Estudio paleontologico y bioestratigrafico de los Arqueociatides de
Sierra Morena (S.W. de Espana).— Monografias Consejo Investnes cient. Madrid. (Sous
presse.)
REPINA, J. N., KHOMENTOVSKII, V. V., ZHURAVLEVA, I. T. & ROZANOv, A. Ju. 1964. Biostrati-
grafiya nizhnego Kembriya Sayano-Altaiskoi skladchatoi oblasti. [Biostratigraphy of the
Lower Cambrian in the Sayan—Altai folded region.] Moskva: Izdatel’stvo ‘Nauka’.
Simon, W. 1939. Archaeocyathacea. I. Kritische Sichtung der Superfamilie. II. Die Fauna im
Kambrium der Sierra Morena (Spanien). — Abh. senckenb. naturforsch. Ges. 448: 1-87.
VoLoGDIN, A. G. 1940a. Arkheotsiaty i vodorosli kembriyskikh izvestnyakov Mongolii i
Tuvy Chast’l.— Trudy mongol’. Kom. 34: 1-268.
VoLocpINn, A. G. 1940b. Atlas rukovodyashchikh form iskopaemykh faun SSSR. (1) Kembriy.
Moskva, Leningrad: Gosgeolizdat.
ZHURAVLEVA, I. T. 1960. Novye dannye ob arkheotsiatakh sanashtykgol’skogo gorizonta. —
Geologiya Geofiz. Novosibirsk 1960 (2): 42-46.
ZHURAVLEVA, I. T., KRASNOPEEVA, S. V. & CHERNYSHEVA, S. V. 1960. Tip Archaeocyathi.
Arkheotsiati. In: KHALFINA, L. L., ed. Biostratigrafiya paleozoya Sayano-Altayskoy
gornoy oblast.— Trudy sib. nauchno-issled. Inst. Geol. Geofiz. miner. Syr’ya (Sniggims) 19:
97-140.
ZHURAVLEVA, I. T., ZADOROZHNAYA, N. M., OSADCHAYA, D. V., POKROVSKAYA, N. V.,
Ropionova, N. M. & Fonin, V. D. 1967. Fauna nizhnego kembriya Tuvy (opornyy raznez
r.Shivelig-Khem). Moskva: Izdatel’stvo ‘Nauka’.
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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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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
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In describing new species, one specimen must be designated as the holotype; other speci-
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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.
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FRANCOISE DEBRENNE
ARCHAEOCYATHA PROVENANT
DE BLOCS ERRATIQUES
DES TILLITES DE DWYKA
(AFRIQUE DU SUD)
VOLUME 67 PART 9 OCTOBER 1975 | ISSN 0303-2515 |
_ ANNALS
OF THE SOUTH AFRICAN
he : ? ah a ape a at 1 eg
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.
Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.—
Ann. Aa. nat. Hist. (13) 2: 309-320.
Konn, A. 1960b. 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-A (frika 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 67 ~~ Band
October 1975 Oktober
Part 9 Deel
A NEW SPECIES OF MEIOSQUILLA
(CRUSTACEA, STOMATOPODA)
FROM SOUTH AFRICA
By
RAYMOND B. MANNING
Cape Town Kaapstad
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A NEW SPECIES OF MEIOSQUILLA (CRUSTACEA, STOMATOPODA)
FROM SOUTH AFRICA
By
RAYMOND B. MANNING
Smithsonian Institution, Washington, D.C., U.S.A.
(With | figure)
[MS accepted 4 March 1975]
ABSTRACT
Meiosquilla barnardi sp. nov. from Natal, South Africa, is described and a key to
Mediterranean and African species of Meiosquilla is provided.
CONTENTS
PAGE
Introduction . Haas : A : ‘ Nie : : . 363
Description . f ' 3 é : : : : : . 363
Key to Mediterranean and African species of Meiosquilla . . 366
Acknowledgements . : : A ; : : ; : . 366
References 5. |) : Aete : : : Soe . 366
INTRODUCTION
The new species of Meiosquilla reported below was first recorded from
South Africa by Barnard (1950) who identified the single specimen available to
him with the Mediterranean Squilla desmarestii Risso. I followed Barnard in
identifying additional material from off Durban with Risso’s species (Manning
1969), then known as Meiosquilla desmarestii. Subsequently, in studying material
of other species of Meiosquilla from West Africa, which resulted in the descrip-
tion of two new species (Manning 1974), I came to the conclusion that the
South African Meiosquilla represented a new species which is described below.
DESCRIPTION
Meiosquilla barnardi sp. nov.
Fig. 1
Squilla desmarestii Barnard, 1950: 842, fig. 1a.
Meiosquilla desmarestii Manning, 1969: 13.
Holotype
1 3g, total length 30 mm; Cape Natal, South Africa; SAM-A1328.
Paratypes
1 3, total length 26 mm; | juvenile, total length 15 mm; 1 fragment,
carapace length 6,4 mm; off Durban, Natal; 29.37,5S, 31.33E; 175-200 metres,
sand, mud; 8 September 1964; University of Cape Town.
1 9, total length 27 mm; data same; USNM 125361.
363
Ann. S. Afr. Mus. 67 (9), 1975: 363-366, 1 fig.
364 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 1. Meiosquilla barnardi sp. nov. Female paratype, TL 27 mm: a, anterior portion of body;
b, eye. Male holotype, TL 30 mm: c, outline of raptorial claw; d, lateral processes of fifth,
sixth, and seventh thoracic somites; e, sixth abdominal somite, telson, and uropod.
(Setae omitted in all figures.)
Description
Eye large, triangular. cornea bilobed, set almost transversely on stalk.
Ocular scales subtruncate. Anterior margin of ophthalmic somite rounded.
Antennal protopod subequal to or slightly longer than carapace. Rostral plate
cordiform, length and width subequal, apex rounded. Carapace smooth, lacking
spines or carinae except for short reflected marginals and laterals on posterior
fourth. Dactylus of raptorial claw with 5 teeth. Propodus of claw shorter than
A NEW SPECIES OF MEIOSQUILLA FROM SOUTH AFRICA 365
carapace, greatest depth near midlength, not markedly tapering distally. Dorsal
ridge of carpus of claw undivided, terminating in rounded lobe. Mandibular
palp absent; 4 epipods present. Exposed thoracic somites lacking submedian
carinae, intermediates present on sixth to eighth somites. Lateral process of
fifth thoracic somite a slender, oblique, laterally-projecting lobe, rounded
laterally ; sharp ventral spine present under each lateral process. Lateral processes
of sixth and seventh thoracic somites broadly rounded. Ventral keel of eighth
thoracic somite slender, erect, apex rounded. Abdomen lacking submedian
carinae on anterior 5 somites, abdominal carinae spined as follows: submedian 6,
intermediate 5—6, lateral 5-6, marginal 4-5. Telson with 3 pairs of marginal
teeth, bases inflated in male holotype, prelateral lobes absent; submedian teeth
with movable apices; carinae of submedian teeth short; denticles sharp,
6-8, 10-11, 1; ventral surface of telson with short postanal keel. Uropodal
exopod broad, distal segment longer than proximal; proximal segment with
5 movable spines on outer margin, distalmost not extending past midlength of
distal segment. Basal prolongation of uropod with inner margin crenulate, not
armed with spinules, inner spine with low, rounded lobe on outer margin.
Colour
Completely faded in all specimens examined.
Size
Males, total length 26-30 mm; only known female, total length 27 mm;
juvenile, total length 15 mm. Other measurements of male holotype, total
length 30 mm; in mm: carapace length ca 7 (damaged); cornea width (damaged) ;
length antennular peduncle 7,6; length, width rostral plate, 2,4; 2,4; raptorial
claw propodus length, depth 6,6; 2,0.
Discussion
Meiosquilla barnardi is the fifth species of the genus to be recorded from
localities outside of the Americas. It resembles the Mediterranean and West
African species and differs from the American species of the genus in having
5 rather than 4 teeth on the dactylus of the raptorial claw (the number 4 used by
me (1969: 4) for M. barnardi in a key to South African species was a /Japsus).
It differs from M. pallida (Giesbrecht) and M. calypso Manning and resembles
M. africana Manning and M. desmarestii (Risso) in having a short antennular
peduncle; in M. calypso and M. pallida the antennular peduncle is longer than
the carapace and rostral plate combined. It differs from M. africana in lacking
erect spinules on the inner margin of the basal prolongation of the uropod. It
agrees with M. africana and differs from M. desmarestii in having a broad
propodus on the raptorial claw. The propodus in both M. africana and
M. barnardi does not markedly taper distally whereas the propodus in M. calypso
M. desmarestii, and M. pallida is broadest near its midlength and markedly
tapers distally.
Meiosquilla barnardi appears to mature at a smaller size than M. desmarestil.
366 ANNALS OF THE SOUTH AFRICAN MUSEUM
The holotype of the new species shows marked swellings at the bases of the teeth
of the telson, a secondary sexual characteristic that does not become apparent in
specimens of M. desmarestii smaller than 50 mm in total length.
It is a pleasure to dedicate this species to Dr K. H. Barnard whose work on
South African decapod and stomatopod crustaceans contributed so much to our
knowledge of these animals.
The 5 species of Meiosquilla occurring in African waters are very similar
morphologically and very difficult to distinguish without comparative material.
The following key may help to separate them.
KEY TO MEDITERRANEAN AND AFRICAN
SPECIES OF MEIOSQUILLA
1. Antennular peduncle as long as carapace and rostral plate combined . . . . 2
Antennular peduncle shorter than or subequal to carapace in length . . . 3
2. Lateral process of fifth thoracic somite parallel to body line, flattened dorsoventrally.
Uropod with 5-6 movable spines on outer margin. (Propodus of claw with greatest
depth at midlength) . : . M. pallida (Giesbrecht)
Lateral process of fifth thoracic somite ‘obliquely flattened, appearing as slender lobe in
dorsal view. Uropod with 8 spines on outer margin. Cee of claw with greatest
depth at midlength) .__. SV hreuellvae calypso Manning
3. Basal prolongation of uropod with 9-17 erect spinules on inner margin. (Propodus of
claw with greatest depth distally) . : ; j : M. africana Manning
Basal prolongation of uropod at most crenulate, unarmed oninner margin. . . 4
4. Propodus of claw slender, tapering distally, greatest depth at midlength
M. desmarestii (Risso)
Propodus of claw broad, not markedly tapering beyond midlength . M. barnardi sp. nov.
ACKNOWLEDGEMENTS
I thank Dr B. F. Kensley, South African Museum, for allowing me to study
the specimen originally reported by Barnard (1950). The illustrations were
prepared by my wife Lilly. My studies on stomatopods from the Indian Ocean
have been supported by the Smithsonian Institution sO its Research
Awards Program.
REFERENCES
BARNARD, K. H. 1950. Descriptive list of South African stomatopod Crustacea (mantis
shrimps).— Ann. S. Afr. Mus. 38: 838-864.
MANNING, R. B. 1969. Notes on some stomatopod Crustacea from southern Africa.—
Smithson. Contr. Zool. 1: 1-17.
MANNING, R. B. 1974. Stomatopod Crustacea. Campagne de la Calypso dans le Golfe de
Guinée et aux Iles Principe, Sao Tome et Annobon (1956), et Campagne aux Iles du Cap
Vert (1959) (suite). —Annls Inst. océanogr., Monaco 50 (1): 1-22.
pied
ae
i v
-
ft
-
i
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 correnes 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...
...1n C. namacolus (Fig. 10)...’
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RAYMOND B. MANNING
A NEW SPECIES OF MEIJOSQUILLA
(CRUSTACEA, STOMATOPODA)
FROM SOUTH AFRICA
VOLUME 67 PART 10 ~—- OCTOBER 1975 ISSN 0303-2515
OF THE SOUTH AFRICAN |
CAPE TOWN
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BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
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FISCHER, P.-H., DuUvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines.—Archs
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Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.—
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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 67 Band
October 1975 Oktober
Part 10 #£Deel
PIVE SPECIES-OF JMEROPSIS
FROM THE SOUTHERN INDIAN OCEAN
(CRUSTACEA, ISOPODA, ASELLOTA)
By
BRIAN KENSLEY
Cape Town Kaapstad
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FIVE SPECIES OF JAEROPSIS FROM THE
SOUTHERN INDIAN OCEAN (CRUSTACEA, ISOPODA, ASELLOTA)
By
BRIAN KENSLEY
South African Museum, Cape Town
(With 10 figures)
[MS accepted 10 March 1975]
ABSTRACT
Five species of Jaeropsis are figured and diagnosed. These include J. waltervadi, a new
species from Walter’s Shoal, and J. beuroisi, a new species from St Paul and Amsterdam
Islands, as well as J. paulensis Vanhoffen from St Paul and Amsterdam Islands, and J. curvicornis
(Nicolet) from Marion Island. The common intertidal South African species, which was
previously misidentified, is given a new name, i.e. J. stebbingi.
CONTENTS
PAGE
Introduction ; : : . . ; 307
Description of material . . . . 367
References . : : Ee I: ; . 380
INTRODUCTION
Material of the genus Jaeropsis from the following sources necessitated this
short report:
Two specimens from Walter’s Shoal (33.138, 43.51E) collected by the
R/V Anton Bruun during the International Indian Ocean Expedition in 1964.
One specimen from Marion Island (46.53S, 37.52E) collected by the third
South African Biological Expedition to the island in 1972-3.
Numerous specimens of two species from St Paul Island (38.448, 77.30E)
and Amsterdam Island (37.55S, 77.40E) collected by the French Expedition to
these islands in 1970-2.
In addition, the species commonly found intertidally around both the west
and east coasts of South Africa was found to be misidentified, and has been
included in this report.
In the accompanying figures, dimensions are in millimetres.
DESCRIPTION OF MATERIAL
Family Jaeropsidae
Genus Jaeropsis Koehler
Jaeropsis waltervadi sp. n.
| Figs 1-2
Diagnosis
Cephalon dorsally broadly convex, with frontal plate anteriorly slightly
concave, lateral margins smooth. Lateral margins of pleotelson bearing 7 small
367
Ann. S. Afr. Mus. 67 (10), 1975: 367-380, 10 figs.
368
sitll pee
u
ern CL So
ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 1. Jaeropsis waltervadi sp. n.
A. Holotype in dorsal view. B. Antenna. C. Antennule. D. 2nd maxilla. E. 1st maxilla.
F. Maxilliped. G. Mandible. H. Uropod.
FIVE SPECIES OF JAEROPSIS FROM THE SOUTHERN INDIAN OCEAN 369
Fig. 2. Jaeropsis waltervadi sp. n.
A. VIIth peraeopod. B. Ist pleopod, g. C. 2nd pleopod, 3. D. 3rd pleopod, ¢.
denticulations. Uropodal basis about as long as broad, inner distal angle
rounded, unarmed, hardly extending beyond narrowly rounded pleotelsonic
apex. Body with broadly rounded median raised area on pleotelson and Vth to
VIlth pereional segments, becoming a double, more defined ridge on perieonal
segments I to IV.
Material
Holotype SAM-A13646 ¢ 2,9 mm Walter’s Shoal, 38-46 metres depth.
Paratype SAM-A13646 3 2,4 mm Walter’s Shoal, 38-46 metres depth.
Remarks
The median dorsal ridge, unarmed uropodal basis, and shape of the
frontal plate of this species make it distinct from any previously described
species of Jaeropsis.
Jaeropsis curvicornis (Nicolet)
Figs 3-4
Jaeropsis curvicornis (Nicolet), Menzies & Schultz, 1967: 174, figs 27-28 (complete synonymy).
non J. curvicornis: Barnard, 1914: 224; 1940: 494.
Diagnosis
Cephalon with frontal plate bearing small median point, lateral margins
smooth. Lateral margins of pleotelson with single incision and stout seta.
Uropodal basis longer than wide, with small hook on inner distal angle,
370 ANNALS OF THE SOUTH AFRICAN MUSEUM
ASAIO
SSS
SSSMh>E¥O™
SS
<n
ew
Fig. 3. Jaeropsis curvicornis (Nicolet)
C. Antennule. D. Mandible. E. 2nd maxilla.
F. Maxilliped. G. Uropod. H. 1st maxilla.
A. ¢ in dorsal view. B. Antenna.
FIVE SPECIES OF JAEROPSIS FROM THE SOUTHERN INDIAN OCEAN Sat
Fig. 4. Jaeropsis curvicornis (Nicolet)
A. VIIth peraeopod. B. Ist pleopod, g. C. 2nd pleopod, 3. D. 3rd pleopod, ¢.
extending well beyond pleotelsonic apex. Body bearing broad band of red-brown
pigment dorsally, expanded on cephalon to include eyes, also somewhat
expanded on pleotelson.
Material
SAM-A13647 ¢ 5,0 mm Marion Island.
Distribution
Patagonia, Falkland Islands, Macquarie Island, Chile, Magellan Straits,
Fuegian Archipelago.
Remarks
The colour pattern of the present specimen agrees well with that given by
Richardson (1909) for J. patagoniensis (= J. curvicornis). Slight differences in
the mouthparts are apparent between the present specimen and the description
given by Menzies & Schultz (1967). The Marion Island specimen has nine
serrated spines on the mandible (13 in the Antarctic specimens) and five fringed
setae on the middle segment of the mandibular palp (as opposed to eight).
For the rest, the specimen agrees well with the figures in the abovementioned
descriptions.
Jaeropsis paulensis Vanhoffen
Figs 5-6
Jaeropsis paulensis Vanh6ffen, 1914: 531, fig. 59a-1. Barnard, 1965: 201, fig. 2b.
Diagnosis
Cephalon with frontal plate rounded, medially entire, lateral margins entire.
Lateral margins of pleotelson entire, fringed with alternating long and short
372 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 5. Jaeropsis paulensis Vanhoffen
A. 3 in dorsal view. B. Antennule. C. Antenna. D. Ist maxilla. E. Mandible.
F. Mandible. G. Maxilliped. H. 2nd maxilla.
FIVE SPECIES OF JAEROPSIS FROM THE SOUTHERN INDIAN OCEAN 373
Fig. 6. Jaeropsis paulensis Vanh6ffen
A. Ist peraeopod. B. VIIth peraeopod. C. Uropod. D. Operculum, 2. E. Ist pleopod, ¢.
F. 2nd pleopod, g. G. 3rd pleopod, g. H. 4th pleopod, g. I. 5th pleopod, ¢.
374 ANNALS OF THE SOUTH AFRICAN MUSEUM
setae. Uropodal basis with broadly rounded lobe medially, pa nile medio-
distal margin serrulate.
Material
SAM-A12285, Gough Island, 2 99 1 ¢.
St Paul Island Amsterdam Island
Station No. Station No.
8a. 1 ovig. 9, 5 99, 14 dS a4. 8 ovig. 99,9 gg
8b. 2 ovig. 99, 1 9,3 dd a8. 2 9°
Sea 2-ovie. 9073-22 9 SS a9.1
b3. ovig. 99, 69S 36
14. 1 ovig. 3, 2 29, 2 dd
27/3/1970/b. 1 3
93.4 ovig. 99, 2 99.766
Remarks
Barnard (1965) remarks that Vanhoffen figures the maxillipedal palp
displaced, and with the second segment unlobed. Figure 5G above shows that
in fact this segment is lobed on the inner margin.
Jaeropsis beuroisi sp. n.
Figs 7-8
Diagnosis
Cephalon with frontal plate obtusely angled, lateral margins with four or
five spinules, more noticeable in smaller specimens, often lacking in adults.
Lateral margins of pleotelson in male with one or two small serrations, in female
with five or six serrations. Uropodal basis longer than wide, with medial lobe
extending furthest distally, tipped with tiny hook; uropods extending beyond
pleotelsonic apex.
Material
Holotype 3 6,0 mm St Paul st.90.
Allotype 2 4,2 mm St Paul st.20.
Paratypes 12 ovig. 99, 5 99, 22 gg. St Paul 22a.
Paratypes 3 ovig. 99, 5 99, 5 g¢. Amsterdam Island st.28.
St Paul Island Amsterdam Island
Station No. Station No.
3.299,1¢ 39: I ovis. 2 lee
7b. 10 ovig. 29, 13 99, 13 gg 4la.19,1¢4
18. 4 ovig. 99, 3 99, 5 3d 41b.1¢
20° Wrovies 2 1eO es 44.19
22c. 9 ovig. 99, 4 22, 12 $s 64a. 1 ovig. 9,1 3
77a. 1 ovig. 2, 1 2, 4 oo 74.4 gd
90. 20 99, 18 3d 94.2 3d
FIVE SPECIES OF JAEROPSIS FROM THE SOUTHERN INDIAN OCEAN 375
Fig. 7. Jaeropsis beuroisi sp. n.
A. Holotype in dorsal view. B. Antenna. C. Maxilliped. D. 2nd maxilla. E. Antennule.
F. Mandible. G. Uropod. H. Ist maxilla.
376 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 8. Jaeropsis beuroisi sp. n.
A. Ist peraeopod. 3B. VIIth peraeopod. C. Ist pleopod, g. D. 2nd pleopod, d.
E. 3rd pleopod, 3. F. 4th pleopod, g. G. Pleotelson. H. Operculum, 9.
FIVE SPECIES OF JAEROPSIS FROM THE SOUTHERN INDIAN OCEAN 377
St Paul Island Amsterdam Island
Station No. Station No.
B/. 1 © 119. 2 ovig. 29, 8 292, 11 $3
B19.19,1¢ 142b. 2 ovig. 29, 3 99,2 gs.
D6. 1 ovig. 2, 2 29,5 dd 147.19,1¢
29/12/1970. 1 juv. 173. 4 ovig. 29, 6 99, 6 ds
eo wi97t. 19 DI2, 1 ovig. 9; 1 932 ss
Remarks
Jaeropsis beuroisi resembles J. intermedius (Nicolet) recorded from
Argentina to southern Chile, the Falkland Islands, and Antarctica, in the
structure of the uropods, antennae (especially the fringed antennae), but differs
in the degree of spination of the cephalic and pleotelsonic margins. The frontal
plate in J. intermedius is more acute than in the present species. Several differences
in the setation and spination of the mouthparts are also apparent.
There is some resemblance to J. marionis Beddard (1886a, 1886), but the
figures of this species are probably somewhat misleading. As the species was
described from a single male, the specific definition needs to be supplemented.
The species is named for Dr J. Beurois of Marseilles, who collected most
of the material.
Jaeropsis stebbingi sp. n.
Figs 9-10
Jaeropsis curvicornis non Nicolet, Barnard, 1914: 224, pl. 20c; 1940: 434, 494; 1965: 202,
fig. 2c. Menzies & Schultz, 1967: 174. Stebbing, 1905: 51, pl. 11 (fig. C).
Diagnosis
Cephalon with frontal plate rounded, medially entire, lateral margins entire.
Lateral margins of pleotelson usually with four very fine serrations on each side,
often difficult to detect. Uropodal basis with broadly rounded medial lobe
tipped with short hook, hardly interrupting pleotelsonic marginal outline;
medio-distal margin serrulate. Cephalon bearing broad curved dark pigment
band.
Material
Holotype SAM-A13649, ¢ 5 mm. Mouille Point, Table Bay.
Allotype SAM-A13648, ovig. 2 3,6 mm. Liideritz, South West Africa.
SAM-A12738 2 ovig. 99, 2 29, 1 ¢ Liideritz, intertidal.
SAM-A12405 1 ¢ Liideritz.
SAM-A12406 1 ¢ Liideritz.
SAM-A12588 a & Liideritz.
SAM-A10381 1 ovig. 2, 1 § Lambert’s Bay.
ie oes S 5 ovig. 22, 7 29, 6 df Mouille Point, Table Bay.
SAM-A2687 1 ovig. 9 St James, False Bay.
378 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 9. Jaeropsis stebbingi sp. n.
A. Holotype in dorsal view. B. Maxilliped. C. Antenna. D. Mandible. E. Antennule.
F. Ist maxilla. G. 2nd maxilla. H. Mandible.
FIVE SPECIES OF JAEROPSIS FROM THE SOUTHERN INDIAN OCEAN 379
Fig. 10. Jaeropsis stebbingi sp. n.
A. Ist peraeopod. B. VIIth peraeopod. C. Uropod. D. Ist pleopod, 3. E. Operculum, ¢.
F. 2nd pleopod, g. G. 3rd pleopod, g. H. 4th pleopod, ¢.
380 ANNALS OF THE SOUTH AFRICAN MUSEUM
Remarks
As noted by Barnard (1965) and Menzies & Schultz (1967), the Jaeropsis
curvicornis described by Stebbing (1905) from Ceylon and later recorded from
South Africa was probably not the same as J. curvicornis described by Nicolet
(1849) from Chile. From the excellent figures provided by Menzies & Schultz
(1967) of J. curvicornis (Nicolet), from the Antarctic, it is immediately apparent
that the South African species is not the same as the Antarctic—Subantarctic
species. The most obvious differences are to be seen in the uropods (curved and
hooked in J. stebbingi, straight and elongate in J. curvicornis), the rostral plate
(evenly convex in J. stebbingi, truncate with small median tooth in J. curvicornis)
and in the lateral margins of the pleotelson (serrate in J. stebbingi, with a single
strong incision in J. curvicornis).
REFERENCES
BARNARD, K. H. 1914. Contributions to the crustacean fauna of South Africa. 1. Additions to
the marine Isopoda.— Ann. S. Afr. Mus. 10: 197-230.
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
identification of the hitherto recorded marine and fresh-water species. — Ann. S. Afr. Mus.
32: 381-543.
BARNARD, K. H. 1965. Isopeda and Amphipoda collected by the Gough Island Scientific
Survey.— Ann. S. Afr. Mus. 48: 195-210.
BEDDARD, E. F. 1886a. Preliminary notice of the Isopoda collected during the Voyage of
H.M.S. ‘Challenger’. Part III.—Proc. zool. Soc. Lond. 1886: 97-122.
BEDDARD, E. F. 1886b. Report on the Isopoda collected by H.M.S. Challenger during the
years 1873-76. Part II.— Rep. scient. Results Voy. Challenger 17: 1-178.
MENZIES, R. J. & SCHULTZ, G. A. 1967. Antarctic isopod Crustacea. II. Families Haploniscidae,
Acanthaspidiidae, and Jaeropsidae, with diagnoses of new genera and species. — Antarctic
Res. Ser. Washington 11: 141-184.
NICOLET, H. 1849. Crustaceos. In: Gay, C. Historia fisica y politica de Chile, Zool. 3: 1-547.
Paris & Santiago: The Author.
RICHARDSON, H. 1909.-Description of a new isopod of the genus Jaeropsis from Patagonia. —
Proc. U.S. natn. Mus. 36: 421-422.
STEBBING, T. R. R. 1905. Report on the Isopoda collected by Professor Herdman at Ceylon
in 1902. In: HERDMAN, W. A. Report to the government of Ceylon on the pearl oyster
fisheries of the Gulf of Manaar, with supplementary reports upon the marine biology of
Ceylon, by other naturalists. Part IV. Supplementary report no. 23: 1-64. London:
Royal Society.
VANHGOFFEN, E. 1914. Die Isopoden der Deutschen Siidpolar-Expedition 1901-1903.— Dt.
Stidpol.-Exped. 15: 449-598.
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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Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
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Leda plicifera A. Adams, 1856 : 50.
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In describing new species, one specimen must be designated as the holotype; other speci-
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not regarded as paratypes should be listed separately. The complete data (registration number,
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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
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‘Revision of the Crustacea. Part VIII. The Amphipoda.’
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to initial capital letter, provided the same generic name is used consecutively.
BRIAN KENSLEY
FIVE SPECIES OF JAEROPSIS
FROM THE SOUTHERN INDIAN OCEAN
(CRUSTACEA, ISOPODA, ASELLOTA)
VOLUME 67 PART 11 OCTOBER 1975 ISSN 0303-2515
an
iz
Con
OF THE SOUTH AFRICAN
MUSEU E
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. ;
Kon, A. J. 19605. Spawning behaviour, egg masses and larval development in Conus from the Indian
Ocean.— Bull. Bingham oceanogr. Coll. 17 (4): 1-51. z
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 67 ~ Band
October 1975 Oktober
Part 11 Deel
A PRELIMINARY CATALOGUE
OF IDENTIFIABLE FOSSIL FISH
MATERIAL FROM SOUTHERN AFRICA
By
R. A. JUBB & B. G. GARDINER
Cape Town Kaapstad
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are issued in parts at irregular intervals as material
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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap
A PRELIMINARY CATALOGUE OF IDENTIFIABLE FOSSIL FISH
MATERIAL FROM SOUTHERN AFRICA
By
R. A. JuBB
Albany Museum, Grahamstown
&
B. G. GARDINER
Queen Elizabeth College, University of London
(With 38 figures)
[MS. accepted 26 March 1975]
ABSTRACT
Records comprising systematics, type and other material and localities of fossil fish
material from the Witteberg, Dwyka, Ecca and Stormberg Series, from Tertiary marine
deposits and from obscure horizons of southern Africa, are listed.
CONTENTS
PAGE
Introduction ; : , : OL
Brief summary of fossil fish records : : . 384
Fossil fish material from the Witteberg Series aoe 4388
Fossil fish material from the Dwyka Series . Pind ee ee
Fossil fish material from the Ecca Series. ; .. 399
Fossil fish material from the Beaufort Series é . 402
Fossil fish material from the Stormberg Series. 2.0) 1423
Species whose horizon is obscure . ; 0 AD
Fossil fish material from Tertiary marine deposits oi, 427
Acknowledgements . é 5 ; ; ‘ ; See: Sif |
FPCIEICHCCS Nee Ee ea Mes a ks eee. SS
INTRODUCTION
The rich fossil beds of the Beaufort Series, South Africa, containing
reptilian and amphibian remains, first discovered by Andrew Geddes Bain in
the fifties of the last century, attracted a great deal of attention from both
professional and amateur palaeontologists. From that time onwards until early
this century a large quantity of fossil material, including fishes, was collected,
much of which was sent overseas. Fortunately, as far as fossil fishes are
concerned, most of the type material not housed in museums in the Republic of
South Africa is to be found in collections in the United Kingdom. Reference to
type material in the British Museum and other institutions in the United
Kingdom, so necessary for the preparation of this catalogue and identification
381
Ann. S. Afr. Mus. 67 (11), 1975: 381-440, 38 figs.
382 ANNALS OF THE SOUTH AFRICAN MUSEUM
of new material, was thus greatly facilitated by the collaboration of a leading
authority on fossil fishes, Dr B. G. Gardiner of Queen Elizabeth College,
London University.
From published data the fossil fish record in this part of Africa goes back
to the Lower Carboniferous. The occurrence of fragmentary fish remains in the
Upper Witteberg Series was known for some time but it was only in the years
1962 to 1963 (Theron 1962; Marais 1963) that the extent of the fossil bed, which
proved to be a strike of some 400 kilometres, was known. In describing this
Upper Witteberg fish fauna Gardiner (1969) recognized eleven new species
representing eight families, and Jubb (1965) described one new species.
To date the Dwyka Series (Upper Carboniferous) and the Ecca Series
(Lower Permian) have produced only a few fossil fish species. During Beaufort
times (Upper Permian to Lower Triassic) fishes appear to have flourished and
twenty-eight species representing twelve families were described (Broom 1909a,
1913a-c; Brough 1931, 1934; Haughton 1934; Woodward 1889). This period
was followed by a decline in the number of species known, there being only
three from the Cave Sandstone of the Stormberg Series. These are Semionotus
capensis Woodward, specimens of which have been found in large numbers in
the Ficksburg and Senekal districts (Jubb 1973), a single specimen of
Daedalichthys formosa (Broom), recorded by Haughton (1924) from Siberia,
Wodehouse District, and a single specimen of Endemichthys likhoeli Forey &
Gardiner, recorded from Mount Likhoeli, western Lesotho (Forey & Gardiner
1973). ;
Referring to Endemichthys likhoeli, Forey & Gardiner (1973) draw attention
to the fact that although the Lower Triassic of South Africa has yielded an array
of early dictyopygids (Brough 1931, 1936) not one of those advanced dictyopygids
so characteristic of the Upper Triassic of North America (Schaeffer 1967) has
so far been recorded from the African continent. The geographical distribution
of the dictyopygids appears to be a discontinuous one with the Lower Triassic
forms coming mainly from South Africa, the Middle Triassic ones from
Australia, and the Upper Triassic forms from North America. It is thus
surprising to find a seemingly advanced dictyopygid in the rather unfossiliferous
Cave Sandstone of Lesotho.
The Cave Sandstone of the Stormberg Series marks the end, in southern
Africa, of the fish fossil record. Apart from sharks’ teeth from marine deposits of
Cainozoic times (Mountain 1962; Davies 1965) no fossil fishes have been found
in Cretaceous, Tertiary or Quaternary deposits. However, in Africa north of the
equator, particularly in the Sahara region (Greenwood 1974) (Fig. 1), fossil
fishes representing extant families and genera have been found at numerous
sites. In Australia all available evidence indicates that the chief genera of extant
freshwater fishes of Australia were present in the continent during much, if not
all of Cainozoic times and for the most part they continue to flourish (Hills
1958).
In southern Africa fossil fish material is housed in the major museums, the
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 383
Fig. 1. Simplified map showing sites where fossil fish remains, representing extant genera of
African freshwater fishes, have been found. (After Greenwood 1974.)
A. Mainly Miocene (Algeria and Egypt) to Upper Pleistocene (Sahara).
© Mainly Lower Pleistocene to Holocene.
The following genera are represented at scattered sites: Protopterus, Polypterus, Hyperopisus,
Hydrocynus, Alestes, Labeo, Barbus, Clarotes, Bagrus, Auchenoglanis, Chrysichthys, Synodontis,
Clarias, Heterobranchus, Lates, Tilapia, Chrysophris.
Geological Survey Department, Pretoria, the Bernard Price Institute for
Palaeontological Research, University of the Witwatersrand, Johannesburg,
and the Rubidge Collection, Wellwood Farm, Graaff-Reinet. In connection
with various studies most of the old and recent material has been studied either
by Gardiner or by Jubb, and overseas material studied by Gardiner and
Hutchinson (1973). All the major museums in the Republic of South Africa and
Rhodesia were visited by Jubb and type specimens and other catalogued
384 ANNALS OF THE SOUTH AFRICAN MUSEUM
material photographed using Kodachrome II 35 mm film. This particular film
was chosen for colour transparencies because of its excellent colour detail and
lack of grain. For taxonomic revision and confirmation of identifications these
colour transparencies were forwarded to Gardiner for comparison with type
and other material in collections in the United Kingdom. As many of the older
descriptions were based on fragmentary samples, mere scales or poorly preserved
material, as well as unsatisfactory locality records, exchange of photographic
slides by air proved far more satisfactory than the exchange of weighty samples
by sea when revising old material, or identifying more recent discoveries. For
guidance on geological horizons from which certain fossil fishes may have come
particular tribute must be paid to Dr J. W. Kitching and Mesdames H. M.
Anderson and J. M. Anderson of the Bernard Price Institute for Palaeontological
Research.
BRIEF SUMMARY OF FOSSIL FISH RECORDS
WITTEBERG SERIES
Family Rhadinichthyidae Romer, 1945
Genus Mentzichthys Jubb, 1965
M. walshi Jubb, 1965
M. jubbi Gardiner, 1969
M. maraisi Gardiner, 1969
M. theroni Gardiner, 1969
Family Holuridae Moy-Thomas, 1939
Genus Australichthys, Gardiner, 1969
A. longidorsalis Gardiner, 1969
Family Atherstoniidae Gardiner, 1967
Genus Aestuarichthys Gardiner, 1969
A. fulcratus Gardiner, 1969
Family Willomorichthyidae Gardiner, 1969
Genus Willomorichthys Gardiner, 1969
W. striatulus Gardiner, 1969
Family Canobiidae Aldinger, 1937
Genus Sundayichthys Gardiner, 1969
S. elegantulus Gardiner, 1969
Family Dwykiidae Gardiner, 1969
C 1s Dwykia Gardiner, 1969
D. anale. sis Gardiner, 1969
Family Amphicentridae Moy-Thomas, 1939
Genus Adroichthys Gardiner, 1969
A. tuberculatus Gardiner, 1969
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 385
Family Platysomidae Young, 1866
Genus Soetendalichthys Gardiner, 1969
S. cromptoni Gardiner, 1969
Family Acanthodidae Agassiz, 1833
Genus Acanthodes Agassiz, 1833
Acanthodes sp. Gardiner, 1973
DWYKA SERIES
Family Elonichthyidae Aldinger, 1937
Genus Namaichthys Girich, 1923
N. schroederi Giirich, 1923
Family Cosmoptychiidae Gardiner, 1963
Genus Watsonichthys Aldinger, 1937
W. lotzi (Giirich, 1923)
Family Palaeoniscidae Aldinger, 1937
Genus Palaeoniscum Agassiz, 1833
P. capensis Broom, 1913
Note
The type specimen of Palaeoniscum capensis, SAM-1061, is stated to come
from the Hantam Mountains, 19 km west of Calvinia, Upper Dwyka. The
original record, crossed out in the register, says Waterval Farm, Prince Albert
District, Cape Province. Under the same name are two specimens SAM-1062
and SAM-1066 with squamation quite different from that of the type specimen,
SAM-1061. These two specimens are recorded as coming from the White Band,
Upper Dwyka Shales, Toren, Hantam Mountains, near Calvinia. They are not
P. capensis and have been provisionally assigned to Watsonichthys lotzi.
Recorded by Giirich (1923) are fragments from the Upper Dwyka Shales,
Ganikobis, South West Africa, representing Elonichthys (?) and Rhadinichthys (?)
species, but no additional material available.
Recorded by Anderson (1972, 1974) are probable fish trails from inter-
glacial sediments in the Dwyka Series, near Vryheid, Natal.
Recorded by Dr V. von Brunn, Natal University, Pietermaritzburg (in Jitt.
1974) are fish trails from the Dwyka on the farm Rooipoort 565, south-east of
Vryheid.
ECCA SERIES
Family Elonichthyidae Aldinger, 1937
Genus Namaichthys Giirich, 1923
N. digitata (Woodward, 1891)
LN. molyneuxi (Woodward, 1903) = N. digitata]
Family Coelacanthidae Agassiz, 1843
Genus Coelacanthus Agassiz, 1844
C. dendrites Gardiner, 1973
386 ANNALS OF THE SOUTH AFRICAN MUSEUM
BEAUFORT SERIES
Lower Beaufort
Tapinocephalus Zone
Endothiodon Zone
Daptocephalus Zone
Family Palaeoniscidae Vogt, 1852
Genus Pteronisculus White, 1933
Pteronisculus sp.
Family Atherstoniidae Gardiner, 1967
Genus Atherstonia Woodward, 1889
A. scutata Woodward, 1889
[(2) A. sculptus (Egerton, 1856) = A. scutata Woodward, 1889]
[(?) A. bainii (Egerton, 1856) = A. scutata Woodward, 1889]
A. minor Woodward, 1889
A. seeleyi Woodward, 1889
A. cairncrossi Broom, 1913 = A. scutata Woodward, 1889
Family Elonichthyidae Aldinger, 1937
Genus Elonichthys Giebel, 1848
E. whaitsi Broom, 1913
Note
There is a remarkable resemblance between E. whaitsi from Droogvoets
Farm, Fraserburg District, and E. browni (Broom) from Bekker’s Kraal fossil
fish bed, Cynognathus Zone.
Middle Beaufort
Lystrosaurus Zone
Family Platysomiidae Young, 1866
Genus Caruichthys Broom, 1913
C. ornatus Broom, 1913
Upper Beaufort
Procolophon Zone
Cynognathus Zone
Family Hybontidae Owen, 1846
Genus Hybodus Agassiz, 1837
HM. africanus Broom, 1909
Family Ceratodontidae Gill, 1872
Genus Ceratodus Agassiz, 1838
C. capensis Woodward, 1889
C. ornatus Broom, 1909
C. kannemeyeri Seeley, 1897
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 387
Family Coelacanthidae Agassiz, 1843
Genus Coelacanthus Agassiz, 1844
C. africanus Broom, 1905
Family Elonichthyidae Aldinger, 1937
Genus Elonichthys Giebel, 1848
E. browni (Broom), 1909 (Originally described as Oxygnathus browni)
Family Dicellopygidae Romer, 1945
Genus Dicellopygae Brough, 1931
D. draperi (Woodward, 1893)
[D. tenuis (Broom, 1909) = D. draperi (Woodward)]
[D. macrodentatus Brough, 1931 = D. draperi (Woodward)]
D. lissocephalus Brough, 1931
Family Redfieldiidae Berg, 1940
Genus Daedalichthys Brough, 1931
D. formosa (Broom, 1909)
[D. higginsi Brough, 1931 = D. formosa (Broom)]
Genus Helichthys Broom, 1909
H. browni Broom, 1909
[H. stegopygae Brough, 1931 = H. browni (Broom)|
[H. obesus Brough, 1931 = H. browni (Broom)]
H. elegans Brough, 1931
H. ctenipteryx Brough, 1931
H. grandipennis Brough, 1931
Family Colobodontidae Stensid, 1916
Genus Meidiichthys Brough, 1931
M. browni (Broom, 1909) (Originally described as Pholidophorus browni)
Family Brookvaliidae Berg, 1940
Genus IJschnolepis Haughton, 1934
I. bancrofti Haughton, 1934
Genus Atopocephala Brough, 1934
A. watsoni Brough, 1934
Family Cleithrolepididae Wade, 1935
Genus Cleithrolepidina Berg, 1940
C. extoni (Woodward, 1888)
C. minor (Broom, 1909)
Family Hydropessidae Hutchinson, 1973
Genus Hydropessum Broom, 1909
H. kannemeyeri Broom, 1909
388 ANNALS OF THE SOUTH AFRICAN MUSEUM
STORMBERG SERIES
Molteno Beds
Family Semionotidae Agassiz, 1832
Genus Semionotus Agassiz, 1832
S. cf. capensis Woodward, 1888
Red Beds
Cave Sandstone
Family Semionotidae Agassiz, 1832
Genus Semionotus Agassiz, 1832
S. capensis Woodward, 1888
Family Dictyopygidae Hay, 1889
Genus Endemichthys Forey & Gardiner, 1973
E. likhoeli Forey & Gardiner, 1973
TERTIARY MARINE DEPOSITS
Family Lamnidae
Genus Carcharodon Miiller & Henle, 1838
C. megalodon Agassiz, 1843
C. angustidens Agassiz, 1843
C. sulcidens Agassiz, 1843
C. carcharias (Linnaeus, 1758)
Genus Oxyrhina Agassiz, 1838
O. desori Agassiz, 1843
O. crassa Agassiz, 1843 = Isurus henedictus Davies, 1964
Family Carchariidae
Genus Carcharius Rafinesque, 1810
C. taurus Rafinesque, 1810
Genus Odontaspis Agassiz, 1838
O. macrota (Agassiz, 1843)
O. elegans (Agassiz, 1843)
Family Galeorhinidae
Genus Galeocerdo Miiller & Henle, 1837
G. cuvier (Le Seuer, 1818)
Genus Hemipristis Agassiz, 1843
H. serra Agassiz, 1843
FOSSIL FISH MATERIAL FROM THE WITTEBERG SERIES
Fragmentary fish remains in the Upper Witteberg Series (Lower Dwyka
Shales) have been known for some time (Haughton et al. 1953: 19; Du Toit
1954: 27). More recently Theron (1962: 263) collected nodules containing fossil
fish from the Dwyka River to Willowmore but more particularly in the Prince
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 389
Albert and the Strydomsvlei-Soetendals Vlei areas. In 1963 Mr G. Walsh, an
official of the Sundays River Irrigation Board, discovered a deposit of fossil fish
in the mountain just south of Lake Mentz and close to the entrance of Karoo-
poort (Jubb 1965). Material from Lake Mentz was sent to Gardiner by the
Geological Survey Department, Pretoria, as well as by the Albany Museum,
Grahamstown (Gardiner 1969).
In the Soetendals Vlei area the so-called “passage beds’ are approximately
500 metres thick and the fish-bearing nodules occur some 200-214 metres from
the top of these (Theron 1962: 264). The ‘passage beds’ were formerly regarded
as Lower Dwyka Shales, but are now considered to represent the Upper
Witteberg Series. The nodules are found scattered in a zone some 9 metres thick
and consist of a black, fine-grained, carbonaceous mudstone somewhat flinty
in character. Within the nodules the fish are only moderately well preserved and
occur either singly or up to five or more in a single nodule.
The Lake Mentz fossils come from the northern slope of the Suurberg
Range on Schiet Hoogte farm about 4,5 km south-west of the lake. The
fossiliferous outcrops are situated 549 metres west of the Sundays River at the
point where it enters the main Suurberg Gorge (Marais 1963: 192, fig. 1). The
Upper Witteberg stage in this area is some 405 metres thick and the fossiliferous
band occurs 225 metres above the base in the Upper Shale zone, or some
183 metres from the top and therefore in the same stratigraphical position as the
fish nodules in the Soetendals Vlei area. The fish are confined to a narrow band
of sandy marl about 125 mm in thickness where they are completely flattened
and occur in vast concentrations, indicating rapid extermination and subsequent
burial.
Fragmentary fish remains have also been collected by Mr J. Loock,
Department of Geology, University of the Orange Free State, at Floriskraal
Dam south of Laingsburg, and also by the Geological Survey Department in the
region of the Dwyka River, the above sites being the same horizon as that of
Soetendals Vlei. Thus these Upper Witteberg fish have been along a strike of
some 400 km.
Family Rhadinichthyidae Romer, 1945
Genus Mentzichthys Jubb
1965 — Mentzichthys Jubb: 270.
Mentzichthys walshi Jubb
1965 — Mentzichthys walshi Jubb: 267-272, figs 1-2, pl. 6A-B.
1969— Menizichthys walshi Jubb. Gardiner: 426.
Locality
Upper Witteberg Series, Lake Mentz. Described in detail by Marais
(1963: 191-202).
390 ANNALS OF THE SOUTH AFRICAN MUSEUM
Holotype
Almost complete fish in the Albany Museum, Grahamstown, AM 4562.
Other material
Parts of fishes representing this species on slabs AM 4563, AM 4564 and
AM 4565 in the Albany Museum. Specimens SAM-K1169 and SAM-K1170
are in the South African Museum, Cape Town.
Mentzichthys jubbi Gardiner
Fig. 2
1963—Species ‘A’, Marais: pl. 3A—D.
1969— Menizichthys jubbi Gardiner: 426-429, figs 1-3.
Locality
Upper Witteberg Series, Lake Mentz.
Holotype
Almost complete fish in Geological Survey Department, Pretoria. No
number available.
Other material
Another specimen on same block as holotype in Geological Survey
Department, one block with two specimens in Albany Museum, Grahamstown,
AM 4562. j
Zo f]
SSS F 5 eS ie: +TTi
STAT
LTP IF TT}
rH
OEE 18 a
: SS LiL] SH 77
See ee
ie Ser 4a
pee ae. “3
Fig. 2. Mentzichthys jubbi Gardiner, total length approximately 38 cm.
Restoration by Gardiner.
Mentzichthys maraisi Gardiner
1969 — Mentzichthys maraisi Gardiner: 430, pl. 2.
Locality
Upper Witteberg Series, Lake Mentz.
Holotype
Almost complete fish with caudal missing in Geological Survey Department,
Pretoria. No number available.
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 391
Other material
One large block with two moderately complete specimens, one block with a
single specimen, and some fragments, all in Geological Survey Department,
Pretoria.
Mentizichthys theroni Gardiner
1969— Mentzichthys theroni Gardiner: 431.
Locality
Upper Witteberg Series, Soetendals Vlei.
Holotype
Almost complete fish with caudal missing in South African Museum,
Cape Town, SAM-13570.
Other material
Block containing three incomplete specimens from Soetendals Vlei in
South African Museum. An almost complete specimen from Lake Mentz in the
collection of the Geological Survey Department, Pretoria.
Family Holuridae Moy-Thomas, 1939
Genus Australichthys Gardiner
1969 — Australichthys Gardiner: 432.
Australichthys longidorsalis Gardiner
1969 — Australichthys longidorsalis Gardiner: 432, figs 4-6.
Locality
Upper Witteberg Series, Lake Mentz.
Holotype
Tolerably complete fish wanting caudal and paired fins, Geological Survey
Department, Pretoria. No number available.
Other material
A well-preserved skull from the same site, Albany Museum, Grahamstown,
AM 4596A.
Family Atherstoniidae Gardiner, 1967
Genus Aestuarichthys Gardiner
1969 — Aestuarichthys Gardiner: 434.
Aestuarichthys fulcratus Gardiner
| Fig. 3
1969 — Aestuarichthys fulcratus Gardiner: 434, figs 7-9.
Locality
Upper Witteberg Series, Soetendals Vlei.
392 ANNALS OF THE SOUTH AFRICAN MUSEUM
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* ms iran ae 1. ( pp :
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: : pa oredg igi t47 +L
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Fig. 3. Aestuarichthys fulcratus Gardiner, total length approximately 10,5 cm.
Restoration by Gardiner.
Holotype
Poorly preserved fish wanting caudal and pectoral fins, South African
Museum, Cape Town, SAM-13562.
Other material
One tolerably complete fish from Soetendals Vlei in the South African
Museum, and two tolerably complete specimens from Lake Mentz in the
Albany Museum, Grahamstown, AM 4596D.
Family Willomorichthyidae Gardiner, 1969
Genus Willomorichthys Gardiner
1969 — Willomorichthys Gardiner: 438.
Willomorichthys striatulus Gardiner
Fig. 4
1969— Willomorichthys striatulus Gardiner: 438, figs 10-12.
Locality
Upper Witteberg Series, Soetendals Vlei.
Holotype
A rather distorted fish with caudal missing, South African Museum,
Cape Town, SAM-13541.
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 393
ua
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: LE TTT EON N
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TT TT EEE og
LEH TT] VLILIL, C7
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Fig. 4. Willomorichthys striatulus Gardiner, total length approximately 25 cm.
Restoration by Gardiner.
Other material
Three tolerably complete specimens in the South African Museum from
Soetendals Vlei. One specimen consisting of head and anterior third of body in
the Albany Museum, Grahamstown, AM 4596B, from Lake Mentz, and
AM 4597 from the same site.
Family Canobiidae Aldinger, 1937
Genus Sundayichthys Gardiner
1969 — Sundayichthys Gardiner: 442.
Sundayichthys elegantulus Gardiner
Fig. 5
1969— Sundayichthys elegantulus Gardiner: 443, fig. 13.
Locality
Upper Witteberg Series, Lake Mentz.
Holotype
Fish with caudal missing, Geological Survey Department, Pretoria. No
number available.
Other material
One specimen consisting of anterior third of fish from Soetensdal Vlei in
the South African Museum, and portion of fish in the Albany Museum,
Grahamstown, AM 4596C, from Lake Mentz.
Family Dwykiidae Gardiner, 1969
Genus Dwykia Gardiner
1969— Dwykia Gardiner: 443.
394 ANNALS OF THE SOUTH AFRICAN MUSEUM
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Fig. 5. Sundayichthys elegantulus Gardiner, total length approximately 10 cm.
Restoration by Gardiner.
Dwykia analensis Gardiner
1969— Dwykia analensis Gardiner: 443, pl. 1.
Locality
Lower Dwyka Shales, Soetendals Vlei.
Holotype
Posterior half of fish, wanting dorsal and pelvic fins, in the South African
Museum, Cape Town, SAM-13561.
Family Amphicentridae Moy-Thomas, 1939
Genus Adroichthys Gardiner
1969 — Adroichthys Gardiner: 445.
Adroichthys tuberculatus Gardiner
Fig. 6
1969 — Adroichthys tuberculatus Gardiner: 445, figs 14-16.
Locality
Upper Witteberg Series, Soetendals Vlei.
Holotype
An incomplete fish, wanting fins and caudal, in the South African Museum,
Cape Town, SAM-13597.
Other material
Five tolerably complete specimens in counterpart, three isolated skulls and
one small specimen in the South African Museum, all from Soetendals Vlei. One
juvenile specimen in the Albany Museum, Grahamstown, from Lake Mentz,
AM 4570.
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 395
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Fig. 6. Adroichthys tuberculatus Gardiner, total length approximately 40 cm.
Restoration by Gardiner.
Family Platysomidae Young, 1866
Genus Soetendalichthys Gardiner
1969 — Soetendalichthys Gardiner: 447.
Soetendalichthys cromptoni Gardiner
1969 — Soetendalichthys cromptoni Gardiner: 449, figs 17-18.
Locality
Upper Witteberg Series, Soetendals Vlei.
Holotype
Incomplete fish, wanting caudal and front of head, in the South African
Museum, Cape Town, SAM-13560.
Other material
Two incomplete specimens from Soetendals Vlei in the South African
Museum.
Family Acanthodidae Agassiz
Fig. 7A—B
1932—Acanthodidae. Zittel: 55, fig. 76.
1937—Acanthodidae. Watson: 49, pls 5-14.
1963 — Acanthodii. Greenwood: 302.
1963 — Acanthoéssidae. Jordan: 591.
1973 — Acanthodes sp. Gardiner: 33, figs 2-3.
Acanthodian remains occur in strata from the Upper Silurian to the Lower
Permian; complete or well-preserved remains are comparatively rare. These
ANNALS OF THE SOUTH AFRICAN MUSEUM
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IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 397
small Palaeozoic fishes, 75 mm to 150 mm in length, have fins of an unusual type.
Both paired and unpaired fins have a strong spine at the anterior edge and in
some species there are additional spines situated ventrally between the pectoral
and ventral spines. In many examples only spines are visible in fossil material.
The first recorded acanthodian spine from Africa (Gardiner 1973) came from
Upper Witteberg Shales 9,6 km south-south-east of Laingsburg, Cape Province.
There are, however, examples of characteristic trails, attributed to the movement
of acanthodian fishes over mud (Haughton 1927) to be found in Dwyka and
Ecca deposits. These have been studied by Dr V. von Brunn, Natal University,
Pietermaritzburg, and Miss Ann Anderson (1972) of the Bernard Price Institute
for Palaeontological Research, Johannesburg. Miss Anderson continued her
research on preserved animal trails and has presented her findings as a Ph.D.
thesis.
FOSSIL FISH MATERIAL FROM THE DWYKA SERIES
Family Elonichthyidae Aldinger, 1937
Genus Namaichthys Giirich, 1923
1923— Namaichthys Giirich: 55.
1962—Namaichthys Giirich. Gardiner: 10.
Namaichthys schroederi Giirich, 1923
1923 — Namaichthys schroederi Giirich: 55, figs 14-16, pl. 2.
1954— Namaichthys schroederi Girich. Du Toit: 280.
1962— Namaichthys schroederi Girich. Gardiner: 10, figs 1-3, pl. 6.
1973— Namaichthys schroederi Giirich. McLachlan & Anderson: 37, table 4.
Locality
Ganikobis, 16 km west of Tses Station, Berseba Reserve, South West
Africa. Shale beds, bottom of Upper Dwyka Shales.
Holotype
The type material described by Giirich (1923) was housed in the Preussische
Geologische Landesanstalt zu Berlin, now known as Sammlung des Zentralen
Geologischen Dienstes der Staatlichen Geologischen Kommission der D.D.R.,
Berlin. As a result of World War II many of the specimens housed in this
museum were lost.
Neotype
Geological Survey Department, Pretoria, Republic of South Africa,
No. 7099 and counterpart, head and anterior part of body. From Ganikobis
South West Africa.
Other material
Fragments of specimens in the British Museum (Natural History), London,
from Ganikobis. Bernard Price Institute for Palaeontological Research,
Johannesburg: specimens P.1-P.6 from Zwartbas, Warmbad Basin, South West
398 ANNALS OF THE SOUTH AFRICAN MUSEUM
Africa; specimens P.11—P.23, P.29-P.42. Alexander McGregor Memorial
Museum, Kimberley: specimens 4902 and 5008 from base of Upper Dwyka
Shales near Kimberley (McLachlan & Anderson 1973 table 4).
Family Cosmoptychiidae Gardiner
1963 —Family Cosmoptychiidae Gardiner: 259.
Genus Watsonichthys Aldinger, 1937
1937—Genus Watsonichthys Aldinger: 254, fig. 72.
1963—Genus Watsonichthys Aldinger. Gardiner: 260.
Watsonichthys lotzi (Giirich, 1923)
1923 —Acrolepis lotzi Giirich: 34, figs 2, 4-9, pl. 1.
1937—Acrolepis lotzi Giirich. Aldinger: 260.
1954—Acrolepis lotzi Giirich. Du Toit: 280 (name only).
1962 — Watsonichthys lotzi (Girich). Gardiner: 16.
1963 — Watsonichthys lotzi (Giirich). Gardiner: 260.
1973 — Watsonichthys lotzi (Girich). McLachlan & Anderson: 37, table 4.
Locality
Upper Dwyka Shales, Ganikobis, near Tses, South West Africa.
Holotype
Incomplete fish, showing underside of head and one third of the body. This
specimen was in the Sammlung des Zentralen Geologischen Dienstes der
Staatlichen Geologischen Kommission der D.D.R., Berlin, but may have been
lost, together with Giirich’s type material of Namaichthys schroederi, during
World War II (Gardiner 1962: 16).
Other material
Specimens in the South African Museum: SAM-1062 and SAM-1066,
collected by Mr A. C. Bain, February 1907, White Band, Upper Dwyka Shales,
Toren, Hantam Mountains, near Calvinia. Recorded previously as Palaeoniscus
capensis Broom but now provisionally assigned to Watsonichthys lotzi. Specimen
SAM-1066, the largest available, is of the caudal section of a fish including the
anal fin. The anal has about 50 rays and is distinctly fringe-like posteriorly. The
caudal fin is heterocercal, inequilobate and deeply cleft. Enlarged fulcral scales
are visible; scales on the sides ornamented with ridges which run longitudinally
along the length of the scale and end in 4 to 7 serrations or teeth posteriorly, the
number of serrations depending on the position of the scale.
There are also fragmented impressions of scales and portions of fishes from,
the White Band, Upper Dwyka Shales, from near Loeriesfontein, in the South —
African Museum, registered number SAM-K1162. Except for one specimen
containing numerous scales, these add little to our knowledge of this species.
These scale impressions are excellent, some showing as many as 8-9 ridges, some
joining posteriorly to end in 5-6 serrations. The fragmented caudal sections are
identical with the caudal section of SAM-1066.
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 399
Family Palaeoniscidae Vogt, 1852
Genus Palaeoniscum Agassiz, 1833
1833 —Palaeoniscum Agassiz: 5, 66.
1891 —Palaeoniscus (Palaeoniscum) Blainville. Woodward: 476.
1967 —Palaeoniscus Agassiz. Gardiner: 198.
Palaeoniscum capensis Broom, 1913
1913a—Palaeoniscus capensis Broom: 1, pl. 2(2).
1923 —Palaeoniscus capensis Broom. Girich: 28, 32.
1926 —Palaeoniscus capensis Broom. Deecke: 122.
1937—Palaeoniscus capensis Broom. Aldinger: 96.
1954 — Palaeoniscus capensis Broom. Du Toit: 279.
1962 —Palaeoniscus capensis Broom. Gardiner: 16.
1973 —Palaeoniscus capensis Broom. McLachlan & Anderson: 37, table 4.
Locality
Hantam Mountains, 19 km west of Calvinia, Upper Dwyka. The original
record, crossed out in the register, states Waterval Farm, Prince Albert District,
Cape Province.
Holotype
Caudal section of a fish in the South African Museum, SAM-1061.
Other material
Specimens SAM-1062 and SAM-1066 appear in previous literature as
representing the same species, P. capensis. They are recorded as coming from the
Hantam Mountains but they are not P. capensis and have been transferred to
Watsonichthys lotzi provisionally. See note under Watsonichthys lotzi.
FOSSIL FISH MATERIAL FROM THE ECCA SERIES
Family Elonichthyidae Aldinger, 1937
Namaichthys digitata (Woodward)
Fig. 8
1891 — Acrolepis (?) digitata Woodward: 508, pl. 15 (fig. 4).
1903 — Acrolepis molyneuxi Woodward: 285, pl. 20.
1909— Acrolepis digitata Woodward. Rogers & Du Toit: 209 (name only).
1910—Acrolepis sp. Woodward: 229, pl. 9 (figs 2-4).
1913b— ?Acrolepis addamsi Broom: 400, pl. 20. (See note below.)
1913¢—Acrolepis ( ?) digitata Woodward. Broom: 391.
1923 — Acrolepis digitata Woodward. Giirich: 32, 51.
1923 — Acrolepis molyneuxi Woodward. Giirich: 32, 51.
1926 —Acrolepis (?) digitata Woodward. Deeke: 105 (name only).
1937—Acrolepis ( ?) digitata Woodward. Aldinger: 258.
1947— Acrolepis molyneuxi Woodward. Macgregor: 50 (additional site).
1954— Acrolepis molyneuxi Woodward. Du Toit: 323 (name only).
1962—Acrolepis ( ?) digitata Woodward. Gardiner: 15 (emended diagnosis).
1962— Acrolepis molyneuxi Woodward. Gardiner: 15 (emended diagnosis).
1973— ?Acrolepis addamsi Broom. McLachlan & Anderson: 56, table 4.
400 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 8. Namaichthys digitata (Woodward). Photograph of Albany Museum specimen AM 263,
the cast of which was used by Woodward as a paratype.
Locality
Acrolepis (?) digitata: Woodward’s type, a group of scales, is recorded as
coming from Graaff-Reinet. Woodward’s paratype, Albany Museum specimen
AM 263, is recorded by Broom (1913c) as coming from Coombs River, near
Grahamstown, but this is also not strictly correct, see note below.
Acrolepis molyneuxi: Sengwe Coalfield, Rhodesia, Upper Ecca Shales.
(?) Acrolepis addamsi Broom, 1913, from the Premier Mine, Wesselton,
Kimberley, 41-metre level, approximately of Dwyka age.
Holotype
Acrolepis (?) digitata; group of scales, in the British Museum (Natural
History), London, No. 47080.
Paratype
In the British Museum (Natural History) a cast of Albany Museum,
Grahamstown, specimen AM 263, cast BM(NH) P.6300. Broom (1913c) records
AM 263 (Fig. 8) as coming from Koomes (Coombs) River, 24 km east of |
Grahamstown, but an older record states ‘Fish River’. As it appeared that the
slab had been washed down the Fish River, and may have been discovered at the
confluence of the Coombs and Fish Rivers, it was sent to Dr J. W. Kitching for
examination. Kitching reported (in litt. 1969): ‘I have examined the slab with
the fossil fish on it and have no doubt in my mind that it is a piece of slaty shale
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 401
Fig. 8A. Scales of Namaichthys cf. digitata (Woodward) approximately 5mm x 5mm
showing ‘downwardly directed denticulations’. Specimen F.36 collected Libertas Farm, annex
Fountains Farm, Pearston, C.P. by J. W. Kitching, Bernard Price Institute, Johannesburg.
402 ANNALS OF THE SOUTH AFRICAN MUSEUM
derived from the Ecca Beds. The slab is badly water-worn and must have been
carried by water for some distance. It compares very favourably with the Ecca
shales in Natal which also yielded fish scales.’ In his “Note on palaeoniscid fish-
scales from the Ecca shales, near Ladysmith’ Woodward (1910) remarked on the
resemblance of these scales to those of Acrolepis (?) digitata and Acrolepis
molyneuxi.
Other material
In the South African Museum: SAM-985, Sengwe Coalfield, Rhodesia;
SAM-K4637, farm Zwartskraal, Prince Albert District, from the lowest part of
the Upper Ecca. Specimens from the same site in the Albany Museum, AM 4774.
In the British Museum (Natural History) a group of scales assigned to Acrolepis
molyneuxi, P.9840.
In the National Museum, Bulawayo, Rhodesia, specimen 5913, a large fish,
lacking detail, some 36 cm in length. Specimens QG 14, fragments, in the Queen
Victoria Museum, Salisbury. Some of these latter have well-preserved
Glossopteris impressions as well.
Family Coelacanthidae Agassiz, 1843
Genus Coelacanthus Agassiz, 1843
Coelacanthus dendrites Gardiner
1973 — Coelacanthus dendrites Gardiner: 33, fig. 1.
Locality
Coal Measures of Somkele, KwaZulu (northern Natal), South Africa.
Holotype
In the British Museum (Natural History) specimen P.10510.
Other material
Identical scales occur in the Madumabisa Shales, Chimwar Ranch, Gwacei
Valley, Rhodesia, British Museum (Natural History) specimens P.27563-5. Like
the Coal Measures of Somkele the Madumabisa Shales are considered to be
Lower Beaufort.
Similar scales have also been recorded from nodules well below the White
Band in the lower parts of the Dwyka sediments at Klipneus in the Warmbad
basin, South West Africa.
FOSSIL FISH MATERIAL FROM THE BEAUFORT SERIES
Family Atherstoniidae Gardiner, 1967
1967 —Atherstoniidae Gardiner: 176, 198.
Genus Atherstonia Woodward, 1889
1889a— Atherstonia Woodward: 242.
1928— Broometta Chabakov: 1291.
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 403
Atherstonia scutata Woodward, 1889
Fig. 9
1856—(?) Palaeoniscus ‘sculptus Egerton: 227, pl. 28 (figs 28-30, 32, 35-36, 39-42).
1856—(?) Palaeoniscus bainii Egerton: 227, pl. 28 (figs 26-27, 31, 33-34, 37-38).
1876—(?) Hypterus bainii Owen: 9.
1889a—Atherstonia scutata Woodward: 242, figs 1-3.
1891 —(?) Palaeoniscus sculptus Egerton. Woodward: 485.
(?) Palaeoniscus bainii Egerton. Woodward: 485.
Atherstonia scutata Woodward. Woodward: 514.
1913a—Atherstonia cairncrossi Broom: 3, pl. 1 (fig. 2).
1913c—Amblypterus capensis Broom: 392, pl. 28.
1923 — Atherstonia scutata Woodward. Girich: 32, 53, figs 12, 13a—b.
(?) Palaeoniscus bainii Egerton. Girich: 32.
(?) Palaeoniscus sculptus Egerton. Giirich: 32.
1926—(?) Palaeoniscus sculptus Egerton. Deeke: 122.
(?) Palaeoniscus bainii Egerton. Deeke: 122.
1928 — Broometta cairncrossi Chabakov: 1281.
1937—(?) Palaeoniscus sculptus Egerton. Aldinger: 96.
(?) Palaeoniscus bainii Egerton. Aldinger: 96.
1946—(?) Palaeoniscus sculptus Egerton. Bond: 128, pl. 10.
1962—(?) Namaichthys sculptus (Egerton). Gardiner: 14.
Locality
The type specimen of Atherstonia scutata Woodward is one of four speci-
mens which came from Colesberg, and which was sent to the British Museum
(Natural History) by Dr W. Guyborn Atherstone of Grahamstown. The fossil
site lies within the Daptocephalus Zone (Kitching 1972) of the Beaufort Series.
Fig. 9. Atherstonia scutata Woodward. Illustration of type specimen P.4735 in the British
Museum (Natural History). Total length approximately 25 cm.
Holotype
An almost complete specimen BM(NH) P.4735 in the British Museum
(Natural History), London.
Other material
Specimens AM 260, AM 261 and AM 262 in the Albany Museum, Grahams-
town, which came from the type locality, Colesberg.
404 ANNALS OF THE SOUTH AFRICAN MUSEUM
The type specimen of Amblypterus capensis Broom, 1913, which came from
a quarry near Alice, Cape Province. The counterpart of this fish fossil is in the
British Museum (Natural History), P.46007, labelled originally as Hypterus
bainii Owen, but identified as Atherstonia scutata by Woodward.
Note
The species Palaeoniscus sculptus and Palaeoniscus bainii described by
Egerton (1856) were described from scanty and poorly preserved scales collected
on Styl Krantz (Steilekrans) Farm near New Bethesda, north of Graaff-Reinet.
The syntypes are in the British Museum (Natural History), P.12192, P.12193 and
P.12194. Both the material and the description are unsatisfactory for identifi-
cation purposes. The species Hypterus bainii Owen, 1876, was not described,
only the name was submitted (BM(NH) P.46007).
The Albany Museum has fragments of fish from Styl Krantz, mostly
scales, recorded AM 4149, some of which have been examined by Gardiner.
These scales can be referred to Atherstonia scutata. Kitching has informed Jubb
(in litt. 1973) that he has collected at Styl Krantz and, apart from fish scales, has
found no other fish material. The horizon where fish scales were found can be
referred to the Daptocephalus Zone (Kitching 1972).
Atherstonia seeleyi Woodward, 1893
Figs 10, 12-13
1893 —Atherstonia seeleyi Woodward: 393, pl. 17 (figs 3—3a).
Locality
Farm Klip Fontein, south-west of Fraserburg, Nieuwveldt Range, Cape
Colony. Daptocephalus Zone, Lower Beaufort.
Syntype
Fragment of a fish with distinctive scales, in the British Museum (Natural
History) P.8613 (Fig. 10).
Other material
A number of fine specimens, V 55—V 64 in the Victoria West Museum,
V 63 and V 64 being almost complete, from Blourug 16 km east of Victoria
West (Figs 12-13).
Atherstonia minor Woodward, 1893
Fig. 11, 14
1893 — Atherstonia minor Woodward: 395, pl. 17 (figs 22a).
Locality
Farm Klip Fontein, south-west of Fraserburg, Nieuwveldt Range, Cape
Colony. Daptocephalus Zone, Lower Beaufort.
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 405
Fig. 10. Holotype of Atherstonia seeleyi Woodward,
British Museum (Natural History) P.8613. x 2.
By courtesy British Museum (Nat. Hist.), London.
Syntype
Scales only; middle portion of small fish with distinctive scales, in the
British Museum (Natural History) P.8614 (Fig. 11).
Other material
Almost complete specimen in the Albany Museum, AM 4290, from
Plaatjiesfontein near Dwaal, Hanover District. Also fragments from Blouhoogte,
New Bethesda District.
406 ANNALS OF THE SOUTH AFRICAN MUSEUM
Both the above sites are considered to be in the Daptocephalus Zone
(Kitching 1972: 309-311; Kitching in itt. 1973).
Family Elonichthyidae Aldinger, 1937
Genus Elonichthys Giebel, 1848
1848 — Elonichthys Giebel: 249.
1937 —Elonichthys Giebel. Aldinger: 16.
1967 — Elonichthys Giebel. Gardiner: 197.
Fig. 11. Holotype of Atherstonia minor Woodward. British
Museum (Natural History) P.8614. x 2. By courtesy British
Museum (Nat. Hist.), London.
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 407
Fig. 12. Atherstonia seeleyi Woodward. Specimen V.63, Victoria West Museum. Total length
approximately 19 cm. By courtesy of the Victoria West Museum.
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the Victoria West Museum. Total length approximately 17 cm.
408 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 13. Atherstonia seeleyi Woodward. Specimen V.64, Victoria West Museum. Total length
approximately 17 cm. By courtesy of the Victoria West Museum.
Elonichthys whaitsi Broom, 1913
1913a—Elonichthys whaitsi Broom: 2, pl. 2 (fig. 2).
1923 —Elonichthys whaitsi Broom. Giirich: 32.
1926—Elonichthys whaitsi Broom. Deeke: 111.
1962—Elonichthys whaitsi Broom. Gardiner: 17.
Locality
Droogvoets Farm, Fraserburg District, Lower Beaufort Series, Cistecephalus
Zone.
Holotype
Almost complete fish, well preserved, in the South African Museum,
SAM-2358.
Other material
None.
Note =
There is a remarkable resemblance between Elonichthys whaitsi and
Elonichthys browni (Broom) from Bekker’s Kraal, Rouxville District, Orange
Free State, a species from the Cynognathus Zone, Upper Beaufort Series.
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 409
Fig. 14. Atherstonia minor Woodward. Specimen AM 4290 in the Albany Museum. Total
length approximately 12 cm.
Family Platysomidae Young, 1866
Genus Caruichthys Broom
1913a—Caruichthys Broom: 4.
1967—Caruichthys Broom. Gardiner: 197.
Caruichthys ornatus Broom
1913a—Caruichthys ornatus Broom: 4, pl. 1 (fig. 1).
Locality
Lystrosaurus Zone of the Beaufort Series. Described as coming from the
Doorn River, Cradock District.
Holotype
Part of a fish, poorly preserved, with the caudal section missing, in the
South African Museum, SAM-—748.
Other material
None.
Note
In a letter from Mr J. van Rensburg, Secretary, Cradock Divisional
Council, C/2, dated 9 October 1974, he states: ‘As far as I can establish the farm
Doorn River where a fish fossil was discovered by a Mr J. S. du Plessis in 1905
is farm No 232/2. It adjoins the farm Lorraine and is about twenty-seven miles
from Cradock on the road to Graaff-Reinet.’
410 ANNALS OF THE SOUTH AFRICAN MUSEUM
Family Hybontidae Owen, 1846
Genus Hybodus Agassiz
1837— Aybodus Agassiz: vol. 3: 41.
1932— Hybodus Agassiz. Zittel: 68, figs 94-95.
Hybodus africanus Broom
Fig. 15
1909a— Hybodus africanus Broom: 252, pl. 12 (fig. 1).
Locality
Lower Cynognathus Zone of the Beaufort Series at Bekker’s Kraal,
Rouxville, Orange Free State.
Holotype
Almost complete fish in the South African Museum, SAM-1082.
Other material
Fragments SAM-1084, SAM-1183, SAM-1185, SAM-1186, SAM-2776,
SAM-6020, SAM-6039.
Fig. 15. Hybodus africanus Broom. Total length approximately 21 cm.
Reconstruction by Broom.
Family Ceratodontidae Gill, 1872
Genus Ceratodus Agassiz, 1837
1837—Ceratodus Agassiz: vol. 3: 129.
1891 —Ceratodus Agassiz. Woodward: pt 2: 264.
1932—Ceratodus Agassiz. Zittel: 104, figs 164-165.
Ceratodus capensis Woodward, 1889
1889— Ceratodus capensis Woodward: 243, pl. 14 (fig. 4).
1891 —Ceratodus capensis Woodward. Woodward: 269.
1909a— Ceratodus capensis Woodward. Broom: 253.
Locality
Recorded as coming from Smithfield, Orange Free State, but no doubt part
of the fish fauna of Bekker’s Kraal, Lower Cynognathus Zone.
Holotype
Dental plate wanting one or perhaps two anterior denticles in the British
Museum (Natural History), specimen P.4807.
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 411
Ceratodus kannemeyeri Seeley, 1897
1897 — Ceratodus kannemeyeri Seeley: 543.
1909a— Ceratodus kannemeyeri Seeley. Broom: 253.
Locality
Given as Kraai Fontein, Indwe District, above the coal seams, and thus
probably within the Molteno Beds.
Holotype
No record.
Ceratodus ornatus Broom, 1909
1909a— Ceratodus ornatus Broom: 253, pl. 12 (fig. 4).
Locality
Found at Vaalbank near Burghersdorp, Cynognathus Zone of the Beaufort
Series.
Holotype
Dental plate, specimen SAM-3601 in the South African Museum.
Family Coelacanthidae Agassiz, 1843
Coelacanthus africanus Broom
1905 — Coelacanthus africanus Broom: 338.
1909a— Coelacanthus africanus Broom. Broom: 253, pl. 12 (fig. 3).
1931 — Coelacanthus africanus Broom. Brough: 238, pl. 1 (fig. 1).
Locality
Bekker’s Kraal Farm, Rouxville District, Cynognathus Zone, Upper
Beaufort Series.
Holotype
In the South African Museum SAM-6027, Bekker’s Kraal. Rouxville.
Caudal section of a fish.
Other material
In the South African Museum SAM-6028, from same locality as SAM-6027.
Small section of a fish.
Family Elonichthyidae Aldinger, 1937
Genus Elonichthys Giebel, 1848
1848 — Elonichthys Giebel: 249.
1937 — Elonichthys Giebel. Aldinger: 16.
1967 — Elonichthys Giebel. Gardiner: 197.
Elonichthys browni (Broom, 1909)
1909a— Oxygnathus browni Broom: 259, pi. 13 (fig. 8).
1931 — Oxygnathus browni Broom. Brough: 236.
1932—Oxygnathus browni Broom. Zittel: vol. 2: 120.
412 ANNALS OF THE SOUTH AFRICAN MUSEUM
Locality
Bekker’s Kraal Farm, Rouxville District, Orange Free State. Cynognathus
Zone, Upper Beaufort Series.
Holotype
Nearly complete fish with Cleithrolepidina and other species on the same
slab in the South African Museum, SAM-—5978.
Other material
South African Museum, SAM-001, no locality, is probably the same
species. Attention is drawn to the similarity between this species and Elonichthys
whaitsi Broom from Droogvoets Farm, Fraserburg, Cistecephalus Zone.
Family Dicellopygidae Romer, 1945
Genus Dicellopygae Brough, 1931
1931 — Dicellopygae Brough: 238.
1945 — Dicellopygae Brough. Romer: 579.
1967 — Dicellopygae Brough. Gardiner: 198.
Dicellopygae draperi (Woodward, 1893)
Figs. 16-17
1893 — Dictyopygae (?) draperi Woodward: 393, pl. 17 (fig. 1).
1909a— Helichthys draperi (Woodward). Broom: 257, pl. 13 (fig. 6).
1909a— Helichthys tenuis Broom: 258.
1909b— Helichthys draperi (Woodward). Broom: 286.
1931 — Dicellopygae macrodentatus Brough: 239, fig. 1.
1931— Helichthys draperi (Woodward). Brough: 247.
1931 — Dictyopygae (?) draperi (Woodward). Brough: 247.
Locality
Stated by Woodward (1893) to be Rouxville, Orange Free State, Stormberg
Series, Upper Karoo. This undoubtedly refers to Bekker’s Kraal, Rouxville
District, Cynognathus Zone, Upper Beaufort Series.
Holotype
A complete fish except for the caudal fin. Housed in the National Museum,
Bloemfontein, specimen QR 1507. This specimen was housed in the Natal
Museum, Pietermaritzburg, for a while with the registered number NM 20.
Other material
Specimen SAM-6022, South African Museum, described and figured by
Broom (1909a), which came from Bekker’s Kraal, Rouxville District, |
Cynognathus Zone, Upper Beaufort Series; SAM-2762, which came from the
same site.
The Natal Museum, Pietermaritzburg, has two incomplete specimens,
registered number NM 21, which were described by Woodward (1893) as
representing an ‘Unidentified Palaeoniscid Fish’. The locality recorded is
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 413
Fig. 16. Dicellopygae draperi (Woodward). Specimen SAM-6022 in the South African Museum.
Total length approximately 13 cm.
Fig. 17. Dicellopygae draperi (Woodward). Reconstruction of Dicellopygae macrodentatus
Brough, by Brough. Total length approximately 11,5 cm.
‘Molteno Beds, 150 feet above main coal seam, Sutherlands Quarry, Biggars-
berg’. These specimens have been examined and identified as representing
Dicellopygae draperi. As Biggarsberg lies well within the Ecca Series this matter
was referred to Mr M. H. P. Rilett, the geologist who was responsible for
mapping the Biggarsberg area. Mr Rilett has confirmed (in litt. 1973) that the
recorded locality is in doubt.
Identical to NM 21 above is fossil fish 1201D in the Port Elizabeth Museum
(Jubb 1973). This fossil is recorded as coming from near a coal mine, Stormberg,
Cape Province. This specimen, like QR 1507 above, was sent to Gardiner who
identified it as Dicellopygae draperi. Here again the recorded locality requires
careful investigation.
414 ANNALS OF THE SOUTH AFRICAN MUSEUM
Dicellopygae lissocephalus Brough, 1931
1931 — Dicellopygae lissocephalus Brough: 242, fig. 2.
Locality
Lower Cynognathus Zone, Lower Beaufort Series, Bekker’s Kraal, Roux-
ville District, Orange Free State.
Holotype
In the Watson Collection, London University, P.13.
Note
Poor preservation does not enable definite diagnosis but it is highly probable
that D. lissocephalus is a synonym of D. draperi (Woodward, 1893).
Family Redfieldiidae Berg, 1940
Genus Daedalichthys Brough, 1931
1931 — Daedalichthys Brough: 245.
1973 — Daedalichthys Brough. Hutchinson: 271.
Daedalichthys formosa (Broom, 1909)
Fig. 18
1909a— Dictyopyge formosa Broom: 262.
1931 — Daedalichthys higginsi Brough: 245, fig. 3, pl. 1 (fig. 4).
1934— Daedalichthys higginsi Brough: 559, fig. 1, pl. 1 (fig. 1).
1973 — Daedalichthys higginsi Brough. Hutchinson: 273, figs 19-20.
Locality
Lower Cynognathus Zone of the Beaufort Series at Bekker’s Kraal, Roux-
ville, Orange Free State.
Holotype
Almost complete fish in the South African Museum, SAM-2761.
Holotype of Daedalichthys higginsi Brough, University Museum of
Zoology, Cambridge, GN 301.
ies rae a: Pg ee oS a
fee aS g ae ie
Fig. 18. Daedalichthys formosa (Broom). Reconstruction of Daedalichthys higginsi Brough,
by Brough. Total length approximately 10,5 cm.
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 415
Other material
British Museum (Natural History): P.17532, P.17533, P.17538.
Genus Helichthys Broom, 1909
1909a— Helichthys Broom: 254.
1931 — Helichthys Broom. Brough: 246.
1932—Helichthys Broom. Zittel: 123.
1967 — Helichthys Broom. Gardiner: 200.
1973 — Helichthys Broom. Hutchinson: 274.
Helichthys browni Broom, 1909
Fig. 19
1909a— Helichthys browni Broom: 254, pl. 12 (fig. 7).
1931 — Helichthys stegopygae Brough: 252, fig. 6.
1931 — Helichthys obesus Brough: 254, fig. 7.
Locality
Lower Cynognathus Zone, of the Beaufort Series at Bekker’s Kraal,
Rouxville, Orange Free State.
Holotype
Almost complete fish with caudal fin missing, in the South African Museum,
specimen SAM-2767.
Other material
South Africgn Museum: specimens SAM-1086, SAM-1182, SAM-5980,
SAM-6021.
Holotype of Helichthys stegopygae Brough, P 12D in the D. M. S. Watson
Collection, University College, London. Holotype of Helichthys obesus Brough,
P 12B and P 12C in the D. M. S. Watson Collection, University College,
London.
a ET SO ee oe se
—— 5 ae a oa oe ee
= Sop oe eg a _
ALTA LH
a
.
SLIT
=.
LEAL TI FH
a aay ane
Fig. 19. Helichthys browni Broom. Reconstruction of Helichthys stegopygae Brough, by
Brough. Total length approximately 10,5 cm.
416 ANNALS OF THE SOUTH AFRICAN MUSEUM
Helichthys elegans Brough, 1931
Fig. 20
1931 — Helichthys elegans Brough: 248, figs 4-5, pl. 2 (figs 1-2).
1973 — Helichthys elegans Brough. Hutchinson: 275, figs 21-22.
Locality
Lower Cynognathus Zone of the Beaufort Series, Bekker’s Kraal, Rouxville
District, Orange Free State.
Holotype
Represented by three nearly complete specimens in the D. M. S. Watson
Collection, housed in the University Museum of Zoology, Cambridge, P 13A,
P 13B, P 19A.
Other material
Specimen GN 316, in the University Museum of Zoology, Cambridge,
England.
Helicthys ctenipteryx Brough, 1931
1931 — Helicthys ctenipteryx Brough: 256, fig. 8.
Locality
Lower Cynognathus Zone of the Beaufort Series, Bekker’s Kraal, Rouxville,
Orange Free State.
Holotype
An imperfect specimen lacking caudal fin, dorsal fin and posterior part of
body, P 12A in the D. M. S. Watson Collection, University College, London.
Other material
Counterpart of above, P 11A.
WE LI TE: RE
i wa B
Fig. 20. Helichthys elegans Brough. Reconstruction by Brough. Total length about 10,5 cm.
Helichthys grandipennis Brough, 1931
Fig. 21
1931 —Helichthys grandipennis Brough: 259, fig. 10.
Locality
Lower Cynognathus Zone of the Beaufort Series, Bekker’s Kraal, Rouxville,
Orange Free State.
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 417
Holotype
Almost complete specimen, lacking tail and part of head, in the D. M. S.
Watson Collection, University College, London, P 14 and P 24.
—. S — : = I a aga
* a. Fo > FASS tt Ff SS+A TH} S++ : So,
*S Sh £2 ESS Sy IPI SS-S TIT FI FF ~~—~—f-_ fr Z? a, —
“ = ~~ Pee 47 so say ag ty BG ED BPG Gy SY BE EY BS SR ee ED 2 ; fi —
N 4 Renee at a Z 7 AS en By A / } .
_ be = : os f-r— f+ 4 ‘mo a ig ae + dies ie er Se dae b+ ee 8 y es, SLEEP. %
‘ ‘ Pf. PIII IIIT ht by a tty
‘ : See eo Es ae ae 1) tigi, *
i PITPPIT EPIL II EES i HAY fd y BA A wy
/ AREA aan a:
4 PTH
,
, on 222 eee eo
anf 8
Pi a Pon
¢ * > “saa 5
— Litt: == 7g
a iy ==
Fig. 21. Helichthys grandipennis Brough. Reconstruction by Brough. Total length about 11 cm
Family Colobodontidae Stensid, 1916
Genus Meidiichthys Brough, 1931
1931 — Meidiichthys Brough: 261.
1973 — Meidiichthys Brough. Hutchinson: 292.
Meidiichthys browni (Broom, 1909)
Fig. 22
1909a—Pholidophorus browni Broom: 267, pl. 13 (fig. 10).
1931 — Meidiichthys browni (Broom). Brough: 262, figs 11-12, pl. 4 (fig. 1).
1973 — Meidiichthys browni (Broom). Hutchinson: 293, figs 32-33.
Locality
Lower Cynognathus Zone of Beaufort Series at Bekker’s Kraal, Rouxville,
Orange Free State.
Fig. 22. Meidiichthys browni (Broom). Reconstruction by Brough. Total length approximately
10 cm.
418 ANNALS OF THE SOUTH AFRICAN MUSEUM
Holotype
In the South African Museum, specimen SAM-6030.
Other material
South African Museum: SAM-6034. British Museum (Natural History):
P.16074, P.16075. University Museum of Zoology, Cambridge: GN 304-305,
GN 311-314, GN 317, GN 348, GN 355.
Family Brookvaliidae Berg, 1940
Genus Atopocephala Brough
1934— Atopocephala Brough: 564.
1973 — Atopocephala Brough. Hutchinson: 249.
Atopocephala watsoni Brough
1934— Atopocephala watsoni Brough: 564, figs 2-3, pl. 1.
1973 — Atopocephala watsoni Brough. Hutchinson: 250, fig. 7.
Locality
Lower Cynognathus Zone, Beaufort Series, Bekker’s Kraal, Rouxville,
Orange Free State.
Holotype
A specimen with counterpart, the only specimen known to date, P.16079-80,
British Museum (Natural History), London.
Family Dictyopygidae Hay, 1889
Genus Ischnolepis Haughton
1934—Ischnolepis Haughton: 97.
1973 —Ischnolepis Haughton. Hutchinson: 239.
Ischnolepis bancrofti Haughton
Figs 23-24
1934—Ischnolepis bancrofti Haughton: 97, figs 1-2, pl. 29.
1973 —Ischnolepis bancrofti Haughton. Hutchinson: 240, figs 1-6.
Locality
Madumabisa shales, 132 km from Kabwe (Broken Hill), Lunsempfwa
Valley, Zambia. Upper Beaufort Series (Hutchinson 1973: 237).
Holotype
Two almost complete fish on slab SAM-9338 together with counterpart
SAM-9339 in the South African Museum.
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 419
Other material
Material in the British Museum (Natural History) consists of two blocks,
part and counterpart, P.27577-8, and several fragments recorded P.27579;
all contain specimens of Jschnolepis together with an unidentified palaeoniscoid.
From the same locality there are in the South African Museum specimens
SAM-9340, SAM-9350, SAM-9352 and SAM-9353 which contain fragments of
possibly two species of unidentified palaeoniscoid fishes.
Fig. 24. Ichnolepis bancrofti Haughton. Reconstruction by Hutchinson. Total length
approximately 7 cm.
420 ANNALS OF THE SOUTH AFRICAN MUSEUM
Family Cleithrolepididae Wade, 1935
Genus Cleithrolepidina Berg, 1940
1940—Cleithrolepidina Berg: 405.
1973 — Cleithrolepidina Berg. Hutchinson: 302.
Cleithrolepidina extoni (Woodward)
Figs 25-26
1888 — Cleithrolepis extoni Woodward: 141, pl. 6 (figs 6-7).
1895 — Cleithrolepis extoni Woodward. Woodward: 156.
1909a— Cleithrolepis extoni Woodward. Broom: 264.
1973 — Cleithrolepidina extoni (Woodward). Hutchinson: 305, figs 40-42.
Locality
Lower Cynognathus Zone, Beaufort Series at Bekker’s Kraal, Rouxville,
Orange Free State.
Holotype
In the British Museum (Natural History) specimen P.5455.
Other material
British Museum (Natural History): specimens P.5455a, P.16043 with
counterpart P.16044, P.16045, P.16105 with counterpart P.16106. South
Fig. 25. Cleithrolepidina extoni (Woodward). Specimen SAM-1335, counterpart SAM-1085
in the South African Museum. Total length approximately 10 cm.
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 421
Fig. 26. Cleithrolepidina extoni (Woodward). Reconstruction by Hutchinson.
Total length 10 cm.
African Museum: specimens SAM-1059 (Caledon River), SAM-1085, SAM-
1335 and SAM-5981.
Cleithrolepidina minor (Broom)
Fig. 27
1909a—Cleithrolepis minor Broom: 266, pl. 13 (fig. 9).
1931 —Cleithrolepis minor Broom. Brough: 267, fig. 13.
1940—Cleithrolepidina minor (Broom). Berg: 405.
1973 — Cleithrolepidina minor (Broom). Hutchinson: 303, figs 38-39.
Locality
Lower Cynognathus Zone, Beaufort Series, at Bekker’s Kraal, Rouxville,
Orange Free State.
Holotype
In the South African Museum, SAM-6037.
Other material
British Museum (Natural History): P.12594, P.16048-51, P.16067-8 and
counterpart P.16069. University Museum of Zoology, Cambridge: GN 194,
GN 293, GN 328, GN 330 and counterpart GN 331. South African Museum:
SAM-1081, SAM-1084, SAM-5963, SAM-5983, SAM-5985, SAM-5987 and
SAM-5988 all from Bekker’s Kraal, Rouxville; SAM-—2763 from the Caledon
River, Orange Free State.
422 ANNALS OF THE SOUTH AFRICAN MUSEUM
:
Pt the
a 3
v Soa
ee
4 ad
Fig. 27. Cleithrolepidina minor (Broom). Reconstruction by Brough.
Total length approximately 5,3 cm.
Family Hydropessidae Hutchinson, 1973
Genus Hydropessum Broom, 1909
1909a— Hydropessum Broom: 266, pl. 12 (fig. 5).
1967 — Hydropessum Broom. Gardiner: 200.
1973 — Hydropessum Broom. Hutchinson: 316, figs 45-47.
Hydropessum kannemeyeri Broom, 1909
Fig. 28
1909a— Hydropessum kannemeyeri Broom: 266, pl. 12 (fig. 5).
1931 — Hydropessum kannemeyeri Broom. Brough: 236.
1973 — Hydropessum kannemeyeri Broom. Hutchinson: 316, figs 45-47.
Locality
Lower Cynognathus Zone of the Beaufort Series, Bekker’s Kraal, Rouxville
District, Orange Free State.
Holotype
South African Museum specimen SAM-1334.
Other material
British Museum (Natural History): P.16042, P.16180, P.16181. University
Museum of Zoology, Cambridge: No. 358. See Figure 28 for reconstruction by
Hutchinson.
423
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA
ONG
IK OS
iG .
KK SOIT he
EK Mt iN
(=¥ iS Ae
ve} Wines
oe \ ae See
. vi Wa Se
N j a ‘
Nt Ng
\ ~ es /
SA (| LA /
QU PRN /
SS a
Fig. 28. Hydropessum kannemeyeri Broom. Reconstruction by Hutchinson.
Total length approximately 9 cm.
FOSSIL FISH MATERIAL FROM THE STORMBERG SERIES
Family Dictyopygidae Hay, 1889
Genus Endemichthys Forey & Gardiner, 1973
Endemichthys likhoeli Forey & Gardiner
1973 —Endemichthys likhoeli Forey & Gardiner: 29, fig. 1.
Locality
Specimen found lying on a Cave Sandstone plateau at the base of the north-
eastern tip of Mount Likhoeli, 5 km south by south-east of Mafeteng, Mafeteng
District, Lesotho.
424 ANNALS OF THE SOUTH AFRICAN MUSEUM
Holotype
University College, London, catalogue number UC 10000. Posterior
section of a fish showing dorsal and anal fins.
Other material
None.
Family Semionotidae Agassiz, 1832
Genus Semionotus Agassiz, 1832
1832—Semionotus Agassiz: 140.
1895 — Semionotus Agassiz. Woodward: 55.
1960— Semionotus Agassiz. Gardiner: 355.
Semionotus capensis Woodward, 1888
Figs 29-30
1888 — Semionotus capensis Woodward: 138, pl. 6 (figs 1-5).
1895 — Semionotus capensis Woodward. Woodward: 58.
1909a— Semionotus capensis Woodward. Broom: 262.
1915—Semionotus capensis Woodward. Hennig: 49, pl. 3.
1924— Semionotus capensis Woodward. Haughton: 388.
1960— Semionotus capensis Woodward. Gardiner: 355, fig. 71.
1973 —Semionotus capensis Woodward. Jubb: 17, figs 1-5.
Locality
Base of Cave Sandstone, Stormberg Series, Ficksburg, Orange Free State.
Holotype
Four nearly complete fishes on one block, British Museum (Natural
History) P.4089.
Other material
Abundant material in state and provincial museums, some particularly good
specimens being:
South African Museum: SAM-K 436, Clocolan, O.F.S.; SAM-1065, Ficksburg,
O.F.S.
National Museum, Bloemfontein: C 286, Senekal, O.F.S.; C 292, Senekal,
O.F.S.; C 2606, Ficksburg, O.F.S.; C 2821, Ficksburg, O.F.S.; C 2827,
Ficksburg, O.F.S.; C 2842, Witteberge, north-west of Fouriesburg, O.F.S.
Natal Museum, Pietermaritzburg: NM 741, NM 821, Ficksburg, O.F.S.
McGregor Memorial Museum, Kimberley: No. 428, Ficksburg, O.F-.S.
Transvaal Museum, Pretoria: No. 2829, two slabs from Ficksburg; No. 2830,
one small block from Rouxville, O.F-.S.
Albany Museum, Grahamstown: No. 477, two slabs from National Museum, —
Bloemfontein, recorded as coming from Senekal, O.F:S.
As described by Jubb (1973) there are some interesting specimens, B.P.I.
Nos 355, 750 and 752, from the Molteno Stage of the Stormberg Series, in the
Bernard Price Institute for Palaeontological Research, which have been assigned
provisionally to this species.
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 425
Z*
t
Fig. 29. Semionotus capensis Woodward. Specimen No. 428 in the McGregor Memorial
Museum, Kimberley. Total length of upper fish 12 cm.
aS
Ry Cag
eute ss <<
SSSR
WEES SS WES
Fig. 29A. Reconstruction of Semionotus capensis Smith Woodward. Details taken from
several specimens.
SPECIES WHOSE HORIZON IS OBSCURE
Family Holuridae Moy-Thomas, 1939
Genus Disichthys Broom, 1913
1913b— Disichthys Broom: 400.
1967 — Disichthys Broom. Gardiner: 197.
Disichthys kimberleyensis Broom, 1913
1913b—Disichthys kimberleyensis Broom: 400, pl. 21 (figs 1-2).
426 ANNALS OF THE SOUTH AFRICAN MUSEUM
Locality
From De Beers Wesselton Diamond Mine, Kimberley. Horizon obscure.
Holotype
A poorly preserved specimen, almost complete fish, McGregor Memorial
Museum, Kimberley, No. 535.
Other material
Portion of a fish, poorly preserved, with the same number in the McGregor
Memorial Museum, Kimberley.
Fig. 30. Semionotus capensis Woodward. Specimen No. 428 in the McGregor Memorial
Museum, showing concentration of fishes.
Family Holuridae Moy-Thomas, 1939
Genus Peleichthys Broom, 1913
1913b—Peleichthys Broom: 401.
1967 —Peleichthys Broom. Gardiner: 197.
Peleichthys kimberleyensis Broom, 1913
1913b—Peleichthys kimberleyensis Broom: 401, pl. 22 (fig. 1).
Locality
De Beers Mine, Kimberley. Horizon obscure.
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 427
Holotype
Almost complete fish, poorly preserved, in the McGregor Memorial
Museum, Kimberley, No. 536.
Other material
None.
FOSSIL FISH MATERIAL FROM TERTIARY MARINE DEPOSITS
ELASMOBRANCHII
The elasmobranchs listed are those which appear in current literature.
Family Lamnidae
Genus Carcharodon Miiller & Henle, 1838
1838 — Carcharodon Miiller & Henle: 70.
1932—Carcharodon Miiller & Henle. Zittel: 78, fig. 120.
Carcharodon megalodon Agassiz, 1843
Figs 31-32
1843 —Carcharodon megalodon Agassiz: 247, pl. 29.
18895 — Carcharodon megalodon Agassiz. Woodward: pt 1: 417.
1932—Carcharodon megalodon Agassiz. Zittel: 78, fig. 120.
1954—Carcharodon megalodon Agassiz. Du Toit: 434.
1962—Carcharodon megalodon Agassiz. Mountain: 9.
1964—Carcharodon megalodon Agassiz. Davies: 47, fig. 23.
1973 — Carcharodon megalodon Agassiz. Case: 36, figs 135-137.
Locality
Eocene and Phosphate Beds: Alabama and South Carolina, U.S.A.
Miocene: France, Spain, Portugal, Malta, Sicily, Corsica, Germany, Austria
and Maryland, U.S.A. Pliocene: Italy, Belgium and south-east England.
Various Tertiary deposits: Arabian Desert, Central and South America, South
Australia, New Zealand and South Africa.
Type specimens
Various detached teeth in the museums of Paris, Carlsruhe and Strassburg
(Woodward 18895).
Other material
Teeth from early Miocene: Uloa, Zululand. Mid to Upper Eocene:
Bogenfels, South West Africa; Alexandria Beds in South Africa, especially
Birbury Farm, Bathurst District, Albany Museum AM 903, AM 1905-1906,
AM 2912, AM 2915, AM 2917-2918, AM 2920, AM 2924, AM 4255.
428 ANNALS OF THE SOUTH AFRICAN MUSEUM
AAR nn h ‘ip
ni ala igh ga
= =
> 7
J i
ry
8
lai
I
Fig. 31. Teeth of Carcharodon megalodon Agassiz from Uloa, Zululand. By courtesy of the
Oceanographic Research Institute, Durban.
Carcharodon angustidens Agassiz, 1843
Fig. 33
1843 —Carcharodon angustidens Agassiz: 255, pl. 28 (figs 20-25), pl. 30 (fig. 3).
18895 —Carcharodon angustidens Agassiz. Woodward: 412.
1954 — Carcharodon auriculatus (Blainville). Du Toit: 434.
1962—Carcharodon angustidens Agassiz. Mountain: 9.
1973 — Carcharodon angustidens Agassiz. Case: 35, figs 134, 138-139.
Locality
Middle and Upper Eocene: South-east England, northern France, Belgium,
Bavaria, South Carolina and Alabama, U.S.A. Miocene: Western France,
Belgium, north Germany, Italy, Sicily and Maryland, U.S.A. Pliocene:
Antwerp, Belgium. Lower and Middle Tertiary: Egypt, Arabia, Australia,
New Zealand and Alexandria Beds of South Africa.
Holotype
No record.
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 429
Fig. 32. Teeth of Carcharodon megalodon Agassiz from Birbury Farm, Bathurst District,
south-east Cape. Albany Museum specimens AM 4255 and AM 903, the length of the
former 70 mm.
Fig. 33. Teeth of Carcharodon angustidens Agassiz from Birbury Farm, Bathurst District,
south-east Cape. Albany Museum specimens AM 1902 and AM 902, the former being 65 mm
in length.
430 ANNALS OF THE SOUTH AFRICAN MUSEUM
Other material
See Woodward (1889b: 411-415). In South Africa known from Birbury
Farm, Bathurst District. Albany Museum specimens AM 902, AM 1898,
AM 1902, AM 2913-2914, AM 4029.
Carcharodon sulcidens Agassiz, 1843
Fig. 34
1843 —Carcharodon sulcidens Agassiz: 254, pl. 30a (figs 3-7).
1889b—Carcharodon sulcidens Agassiz. Woodward: 420.
1964—Carcharodon sulcidens Agassiz. Davies: 12, fig. 9.
1965 — Carcharodon sulcidens Agassiz. Davies: 48, fig. 24.
1973 — Carcharodon sulcidens Agassiz. Case: 35, fig. 134.
Locality
Pliocene: Italy, Sicily and England. Miocene and Eocene: South Carolina,
U.S.A. Tertiary: Chili and South Africa, Uloa and Sapolwana Beds (Miocene),
northern Zululand.
Holotype
Detached teeth, Palaeontological Museum, Munich (Miinster Collection).
Other material
See Woodward (18895: 420-421). Detached teeth in the ice
Research Institute, Durban (Fig. 34).
Carcharodon carcharias (Linnaeus, 1758)
Fig. 35
1925 — Carcharodon carcharias (Linnaeus). Barnard: 33, pl. 1 (fig. 7).
1932 — Carcharodon rondeleti Miiller & Henle. Zittel: 78.
1961 — Carcharodon carcharias (Linnaeus). Smith: 49, pl. 1 (fig. 26).
1964 — Carcharodon carcharias (Linnaeus). Davies: 9.
1973 — Carcharodon carcharias (Linnaeus). Case: 44, figs 178-180.
This is a widely distributed species which dates back to the Miocene.
Although specimens were not recorded by Davies (1964), material has since been
collected at Sapolwana, Zululand. Two teeth from the collection in the Oceano-
graphic Research Institute, Durban, are illustrated in Figure 35.
Genus Oxyrhina Agassiz, 1838
1838—Oxyrhina Agassiz: vol. 3: 86.
1889b— Oxyrhina Agassiz. Woodward: 376 (1).
1932—Oxyrhina Agassiz. Zittel: 77.
Oxyrhina desori Agassiz, 1843
1843 —Oxyrhina desori Agassiz: vol. 3: 282, pl. 37 (figs 8-13).
1889b— Oxyrhina desori Agassiz. Woodward: 382.
1932—Oxyrhina desori Agassiz. Zittel: 77.
1962—Oxyrhina desori Agassiz. Mountain: 9.
1973 — Oxyrhina desori Agassiz. Case: 36, fig. 151.
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 431
Fig. 34. Teeth of Carcharodon sulcidens Agassiz from Uloa, Zululand. By courtesy of the
Oceanographic Research Institute, Durban. Largest tooth 60 mm in length.
Fig. 35. Teeth of Carcharodon carcharias (Linnaeus) from Uloa, Zululand. By courtesy of the
Oceanographic Research Institute, Durban. Larger tooth 52 mm in length.
432 ANNALS OF THE SOUTH AFRICAN MUSEUM
Locality
Upper Eocene: Prussia, Italy, Alabama and South Carolina, U.S.A.
Miocene: France, Belgium, Switzerland, Germany, Malta, Italy, Sicily and
Corsica. Pliocene: Belgium and Italy. Alexandria Beds, South Africa.
Holotype
Detached teeth.
Other material
See Woodward (18895: 383). Albany Museum material identified by
Dr E. I. White of the British Museum (Natural History) cannot be traced.
Oxyrhina crassa Agassiz, 1843
Fig. 36
1843 —Oxyrhina crassa Agassiz: vol. 3: 283, pl. 37 (fig. 16).
1871—Oxyrhina benedeni Le Hon: 6.
1889b5—Oxyrhina crassa Agassiz. Woodward: 389.
1964 —Jsurus benedictus = (Oxyrhina benedeni) Davies: 12, fig. 11.
1965 —Isurus benedictus = (Oxyrhina benedeni) Davies: 48, fig. 25.
1973 — Oxyrhina (benedeni) crassa Agassiz. Case: 36, fig. 149.
Locality
Eocene: South Carolina and Alabama, U.S.A. Eocene or Miocene: Valley
of the Rhine. Miocene: Sicily. Pliocene: Italy and Belgium.
Holotype
Detached tooth—no other record.
Other material
See Woodward (1889b: 390). Davies (1964), Oceanographic Research
Institute, Durban, records one tooth from the Lower Miocene, Uloa, Zululand.
Family Carchariidae
Genus Carcharias Rafinesque, 1810
1810—Carcharias Rafinesque: 10.
1961 —Carcharias Rafinesque. Smith: 48.
1963 — Carcharias Rafinesque. Jordan: 77, 599.
Carcharias taurus Rafinesque, 1810
Fig. 37, 38C
1810—Carcharias taurus Rafinesque: 10.
1961 — Carcharias taurus Rafinesque. Smith: 25, pl. 1 (fig. 25).
1964—Carcharias taurus Rafinesque. Davies: 11, fig. 10C.
1965 — Carcharias taurus Rafinesque. Davies: 48, fig. 26C.
1973 —Carcharias taurus Rafinesque. Case: 44, fig. 184.
Locality
Upper Cretaceous to present day, widely distributed.
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 433
Fig. 36. Tooth of Oxyrhina crassa Agassiz (Oxyrhina benedeni
Le Hon) from Uloa, Zululand. By courtesy of the Oceanographic
Research Institute. Total length 50 mm.
Holotype
Muséum National d’Histoire Naturelle, Paris.
Other material
Detached teeth from Miocene fossil beds, Sapolwana, South Africa, and
recent material in the Oceanographic Research Institute, Durban. Detached
teeth from Birbury Farm, Bathurst District in Albany Museum AM 4773A
(Fig. 37).
Note
In some literature the genus Carcharias Rafinesque has been replaced by
Odontaspis Agassiz, 1838.
Genus Odontaspis Agassiz, 1838
1838 — Odontaspis Agassiz: vol. 3: 87.
18895— Odontaspis Agassiz. Woodward: 360.
1932 —Odontaspis Agassiz. Zittel: 76.
434 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 37. Tooth of Carcharias taurus Rafinesque from Birbury Farm, Bathurst District,
south-east Cape. Total length 35 mm. See also Figure 38C showing teeth of C. taurus from —
Sapolwana, Zululand.
Odontaspis macrota (Agassiz, 1843)
1843 —Otodus macrotus Agassiz: vol. 3: 273, pl. 32 (figs 29-31).
18895—Lamna macrota (Agassiz). Woodward: 402.
1932 —Odontaspis macrota (Agassiz). Zittel: 76.
1962 —Odontaspis macrota (Agassiz). Mountain: 9.
1973 — Odontaspis macrota (Agassiz). Case: 32, fig. 119.
Locality
Lower Eocene: South-east England and South Carolina, U.S.A. Middle
Eocene: London and Hampshire Basins, and northern France. Upper Eocene:
Southern Germany. Lower Miocene: Belgium and West Germany, Oamaru and
Waipara Systems, New Zealand. Alexandria Beds, South Africa.
Holotype
Detached teeth, Muséum National d’Histoire Naturelle, Paris.
Other material
See Woodward (18895: 402-403). Albany Museum AM 4773B, Birbury
Farm, Bathurst District.
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 435
Note
Quoting Woodward (18895: 360): ‘Teeth of all but the few hindermost
series with a high, narrow, compressed coronal eminence, flanked by one or two
pairs of small pointed denticles; the fourth tooth from the symphysis upon each
side of the upper jaw very small; the teeth of the most anterior pair in the lower
jaw small and relatively very slender.’
Odontaspis elegans (Agassiz, 1843)
1843 — Odontaspis elegans (Agassiz): vol. 3: 289, pl. 35 (figs 1-5), pl. 37 (fig. 59).
1889b—Odontaspis elegans (Agassiz). Woodward: 361.
1932 — Odontaspis elegans (Agassiz). Zittel: 76.
1962—Odontaspis elegans (Agassiz). Mountain: 9.
1973 — Odontaspis elegans (Agassiz). Case: 32, fig. 118.
Locality
Eocene: south-east England, France, Belgium and Germany, also Alabama
and South Carolina, U.S.A. Lower Miocene: Belgium and Corsica, also
Oamaru and Waireka Series, New Zealand.
Holotype
Detached teeth, no other record.
Other material
See Woodward (18896: 363-366). Albany Museum specimen AM 4118
identified by Dr E. I. White, British Museum (Natural History), with reservation.
Family Galeorhinidae
Genus Galeocerdo Miiller & Henle, 1837
1837—Galeocerdo Miiller & Henle: 308.
1889—Galeocerdo Miiller & Henle. Woodward: 443.
1932—Galeocerdo Miiller & Henle. Zittel: 79, fig. 121.
1961 —Galeocerdo Miiller & Henle. Smith: 43.
Galeocerdo cuvier (Le Seuer, 1818)
Fig. 38A
1818—Galeocerdo cuvier (Le Seuer): vol. 1: 222-235
1961 —Galeocerdo cuvier (Le Seuer). Smith: 44, pl. 1 (fig. 14).
1964—Galeocerdo cuvier (Le Seuer). Davies: 11, fig. 10A.
1965 —Galeocerdo cuvier (Le Seuer). Davies: 48, fig. 26A.
1973 —Galeocerdo cuvier (Le Seuer). Case: 44, fig. 187.
Locality
From Eocene marine beds to present day where found in all warm seas.
Holotype
Muséum National d’Histoire Naturelle, Paris.
436
ANNALS OF THE SOUTH AFRICAN MUSEUM
apoio
Fig. 38. A. Teeth of Galeocerdo cuvier (Le Sueur) from Uloa, Zululand. Larger tooth 25 mm.
B. Teeth of Hemipristis serra Agassiz from Uloa, Zululand. Larger tooth 25 mm.
C. Teeth of Carcharias taurus Rafinesque from Uloa, Zululand. Largest tooth 25 mm.
By courtesy of the Oceanographic Research Institute, Durban.
IDENTIFIABLE FOSSIL FISH MATERIAL FROM SOUTHERN AFRICA 437
Other material
Detached teeth from Miocene deposits, Sapolwana, Zululand, South Africa,
as well as recent material housed in the Oceanographic Research Institute,
Durban.
Genus Hemipristis Agassiz, 1843
1843 — Hemipristis Agassiz: 237.
18895 — Hemipristis Agassiz. Woodward: 448.
1932 — Hemipristis Agassiz. Zittel: 79.
Hemipristis serra Agassiz, 1843
Fig. 38B
1843 — Hemipristis serra Agassiz: vol. 3: 237, pl. 27 (figs 18-30).
18895— Hemipristis serra Agassiz. Woodward: 449.
1964 — Hemipristis serra Agassiz. Davies: 11, fig. 10B.
1965 — Hemipristis serra Agassiz. Davies: 48, fig. 26B.
1973 — Hemipristis serra Agassiz. Case: 36, fig. 146.
Locality
Eocene: South Carolina, U.S.A. Miocene: Germany, Austria, Italy,
Sicily, Malta, Corsica, Switzerland, France, Maryland and Virginia, U.S.A.
Pliocene: Tuscany. Miocene beds: Uloa, Zululand, South Africa.
Holotype
Detached teeth, Muséum National d’Histoire Naturelle, Paris.
Other material
See Woodward (18895: 450-451). Detached teeth in the Oceanographic
Research Institute, Durban.
ACKNOWLEDGEMENTS
In August 1974 Dr A. J. D. Meiring of Alice presented fossil fish material
to the Albany Museum. This came from a quarry (Lower Beaufort) near the
town. On one slab and counterpart, together with a specimen of Atherstonia
scutata, there is the anterior portion of a large palaeoniscid. From photographs
and description sent to him Gardiner has provisionally identified this fossil as
representing an undescribed species of the genus Preronisculus White, 1933, of
the Triassic of Madagascar.
Current research by Mr J. Loock of the University of the Orange Free
State, and Dr J. N. Theron of the Geological Survey Department, also involves
undescribed fish fossil material. Theron’s specimens from the Bokkeveld Beds
date back to the Devonian. Both these workers are collaborating with Gardiner.
Miss Ann Anderson, Mrs Seidi Anderson, Dr J. W. Kitching and Dr I. R.
McLachlan of the Bernard Price Institute for Palaeontological Research have
been unstinting in their supply of material and data. Illustrations of sharks’
438 ANNALS OF THE SOUTH AFRICAN MUSEUM
teeth from Uloa, Zululand, were kindly supplied by the Director of the
Oceanographic Research Institute, Durban. i
During his tour of the state and provincial museums to examine fish fossil
material Jubb was provided with every assistance by the respective directors and
staff members concerned. This co-operation is gratefully acknowledged, as well
as that of the Curator and staff of the Bulawayo Museum, Rhodesia.
This project, proposed by Dr T. H. Barry, Director of the South African
Museum, and made possible by the collaboration of Dr B. R. Gardiner of the
Queen Elizabeth College, London, is part of a research programme sponsored
by the South African Council for Scientific and Industrial Research, Pretoria.
By courtesy of the Director, Mr C. F. Jacot Guillarmod, this catalogue was
prepared at the Albany Museum, Grahamstown.
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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.,
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
e.g. Du Toit but A. L. du Toit
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(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.
R. A. JUBB & B. G. GARDINER
A PRELIMINARY CATALOGUE
OF IDENTIFIABLE FOSSIL FISH
MATERIAL FROM SOUTHERN AFRICA
you UME 67 PART 12 DECEMBER 1975 ISSN 0303-2515 —
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{ANNALS
INSTRUCTIONS TO AUTHORS
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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.
Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.—
Ann. Mag. nat. Hist. (13) 2: 309-320. F
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.
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 67 ~~ Band
December 1975 Desember
Part. “12°” Deel
A NEW PALAEONISCID FROM THE LOWER
BEAUFORT SERIES OF SOUTH AFRICA
By
B. G. GARDINER & RR. A. JUBB
Cape Town Kaapstad
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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap
A NEW PALAEONISCID FROM THE LOWER BEAUFORT SERIES
OF SOUTH AFRICA
By
B. G. GARDINER
Queen Elizabeth College, University of London
&
R. A. JUBB
Albany Museum, Grahamstown
(With 1 figure)
[MS. accepted 26 March 1975]
ABSTRACT
Pteronisculus meiringi sp. nov. from Alice, Cape Province, is described. This is the first
record of this species from South Africa and extends the range of the genus from the Lower
Trias into the Upper Permian.
CONTENTS
PAGE
Introduction . : : : : : : , . 441
Systematic description . : : ; : : . 442
Description of a new species of Pteronisculus . . 443
Conclusions . 2 E ‘ : : : : . 444
Acknowledgements : : : s : : . 444
meicrences " -. : , : : : : . 445
INTRODUCTION
In 1974 Professor A. J. D. Meiring, recently retired from the Department
of Zoology, University of Fort Hare, Alice, presented some fish fossil material
to the Albany Museum. This material had been discovered by him amongst rock
taken from a quarry situated on the immediate outskirts of Alice in an outcrop
of the Lower Beaufort Series. As both before and after this discovery the quarry
had been used extensively for crushed stone required for road-making, the
search for additional material at this particular site was abandoned.
As far back as 1876 Owen recorded, without formal description, a fossil fish
under the name Hypterus bainii from Alice. This fossil was presented to the
British Museum (Natural History), London, by the Trustees of the Albany
Museum. Under the registered number P.46007 it was included by Woodward
with the material used for his description of Atherstonia scutata in 1889. The
counterpart of P.46007 is in the Albany Museum with the registered number
AM 264, the site recorded being a quarry near Alice. In 1913 Broom described
441
Ann. S. Afr. Mus. 67 (12), 1975: 441-445, 1 fig.
442 ANNALS OF THE SOUTH AFRICAN MUSEUM
AM 264 as a new species Amblypterus capensis but this confusion has been
resolved. By comparing AM 264 with other material it is agreed that it is a
specimen of Atherstonia scutata.
Meiring’s material has been examined and found to represent two species,
several specimens of Atherstonia scutata, a species first described from the
Daptocephalus zone of the Lower Beaufort Series near Colesberg, and a new
species identified by the senior author as belonging to the genus Pteronisculus
White, 1933, which is described below.
SYSTEMATIC DESCRIPTION
Order PALAEONISCIFORMES
Family Palaeoniscidae Aldinger, 1937
Diagnosis
See Aldinger 1937: 229.
Genus Pteronisculus White, 1933
1921 —Glaucolepis Stensié: 200.
Diagnosis
See Nielsen 1942: 265.
Type species
Pteronisculus cicatrosus White, 1933.
Remarks
The genus was erected by White (1933: 118) to include the two species
P. cicatrosus White and P. macropterus White from the Lower Trias of Madagas-
car. Later Nielsen (1936: 1942) showed that Preronisculus was synonymous with
the genus Glaucolepis of Stensié (1921), but failed to note that the name
Glaucolepis was preoccupied.
Other species included within this genus are: P. arambourgi Lehman
(1952: 39) and P. broughi Lehman (1952: 39) also from the Lower Trias of
Madagascar; P. gyrolepidoides (Stensidé, 1921: 200) from the Lower Trias of
Spitsbergen; P. artica (Stensiéd, 1932: 119), P. stensioi (Nielsen, 1942: 271),
P. magna (Nielsen, 1942: 272), P. gunnari (Nielsen, 1942: 273), and P. aldingeri
(Nielsen, 1942: 274) all from the Lower Trias of East Greenland.
It has been suggested by Aldinger (1937: 96) that Palaeoniscum crassus
Woodward (1908: 9) from the late Middle or early Upper Trias of New South
Wales, Australia bears a closer resemblance to Pteronisculus than to Palaeonis-
cum. However, all the fins are small and quite unlike those of Preronisculus,
while the enlarged ridge scales in front of the dorsal fin is a primitive feature and
should not be taken to indicate a relationship to Pteronisculus as suggested by
Aldinger. It is thought that Woodward’s fish will probably prove to be a
redfieldid.
A NEW PALAEONISCID FROM THE LOWER BEAUFORT SERIES 443
DESCRIPTION
Pteronisculus meiringi sp. nov.
Fig. 1
Diagnosis
A large, slender species with an estimated length of 40 cm. Dermal bones of
head ornamented with tubercles and striae. The suspensorium is oblique.
Pectoral fin large, expanded in the horizontal plane and with a well-developed
scaly lobe; there are some 20 to 22 stout lepidotrichia unsegmented in their
proximal third. In each of the anterior 10 transverse scale rows there are a
number of flank scales higher than broad, remaining trunk scales rhombic.
res YUEAY DEY
4 ek
:
Fig. 1. Type specimen of Pteronisculus meiringi sp.n.
Lower Beaufort Series, Alice, Cape Province.
444 ANNALS OF THE SOUTH AFRICAN MUSEUM
Immediately behind the cleithrum there are approximately 20 scales in each
transverse row. Scales ornamented with stout, diagonal ridges of ganoine which
end in delicate serrations posteriorly.
Holotype
Specimen AM 4770A in the Albany Museum, Grahamstown; anterior
one-third of fish with snout missing, from the Lower Beaufort Series of Alice,
Cape Province.
Other material
Specimen AM 4770B, counterpart of holotype AM 4770A, in the Depart-
ment of Zoology, University of Fort Hare, Alice.
Specimen AM 4772 in the Albany Museum; part of the caudal fin.
Remarks
This species is named in honour of Professor A. J. D. Meiring of the Fort
Hare University for Bantu.
CONCLUSIONS
This is a large fish only approached in size by P. magna (Nielsen) and from
which it may easily be separated by the relatively small number of its pectoral fin
rays (P. magna has 45 to 50).
Until the discovery of this new species from South Africa all known
members of the genus Pteronisculus came from the Lower Trias. If the horizon
of the type locality of P. meiringi at Alice is Lower Beaufort then the range of
this genus has been pushed back into Upper Permian.
ACKNOWLEDGEMENTS
We wish to thank the Director of the British Museum (Natural History),
London, and the Director of the Albany Museum, Grahamstown, Mr C. F.
Jacot-Guillarmod, for making material and literature available for this research
work; Dr J. W. Kitching of the Bernard Price Institute for Palaeontological
Research for collaboration regarding the zoning of fossil beds of the Beaufort
Series, as well as the supply of fossil fish material, and Dr T. H. Barry, Director
of the South African Museum, Cape Town, for publishing this paper. The
illustration, Fig. 1, was supplied by Professor A. J. D. Meiring together with
the fossils from Alice.
The study of the fossil fishes of southern Africa is part of a research
programme sponsored by the Council for Scientific and Industrial Research,
Pretoria.
A NEW PALAEONISCID FROM THE LOWER BEAUFORT SERIES 445
REFERENCES
ALDINGER, H. 1937. Permische Ganoidfische aus Ostgrénland.—Meddr Grnland 102 (3):
i-xliv, 1-393.
Broom, R. 1913. On some fossil fishes in the collection of the Albany Museum.— Rec. Albany
Mus. 2: 391-393.
LEHMAN, J. P. 1952. Etude complémentaire des poissons de l’Eotrias de Madagascar.—
K. svenska Vetensk Akad. Hand. (4) 2 (6): 1-201.
NIELSEN, E. 1936. Some few preliminary remarks on Triassic fishes from East Greenland. —
Medadr Grnland 112 (3): 1-55.
NIELSEN, E. 1942. Studies on Triassic fishes from East Greenland. I. Glaucolepis and Boreo-
somus.— Meddr Grnland 138: 1-403.
OweEN, R. 1876. Descriptive and illustrated catalogue of the fossil Reptilia of South Africa in the
collection of the British Museum. London: The Trustees.
STENSIO, E. A. 1921. Triassic fishes from Spitzbergen. Part 1. Wien: Holzhausen.
STENSIO, E. A. 1932. Triassic fishes from East Greenland collected by the Danish Expeditions
in 1929-1931.—Meddr Grnland 83 (3): 1-305.
WuitE, E. I. 1933. New Triassic palaeoniscids from Madagascar.— Ann. Mag. nat. Hist. (10)
11: 118-128.
WoopwaprbD, A. S. 1889. On Atherstonia, a new genus of palaeoniscid fishes from the Karoo
Formation of South Africa.— Ann. Mag. nat. Hist. (6) 4: 239-242.
WoopDwaprD, A. S. 1908. The fossil fishes of the Hawkesbury Series at St Peter’s.— Mem. geol
Surv. N.S.W. Palaeont. 10: i-v, 1-30.
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6. SYSTEMATIC papers must conform with the International code of zoological nomenclature
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Nuculana (Lembulus) bicuspidata (Gould, 1845)
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