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ANNALS OF THE ANNALE VAN DIE
SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

VOLUME 83 BAND 83

ais

a
ff
a,

ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

VOLUME 8&3 BAND

tHE TRUSTEES OF THE DIE TRUSTEES VAN DIE

SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM
CAPE TOWN KAAPSTAD

1980-1981

SET, PRINTED AND BOUND IN THE REPUBLIC OF SOUTH AFRICA BY
THE RUSTICA PRESS (PTY.) LTD., WYNBERG, CAPE
C914

LIST OF CONTENTS

Best, P. B. & SHAUGHNESSY, P. D.
First record of the melon-headed whale Peponocephala electra from South Africa.
PEIGHE OPM IGCh OSI) Sec eke nate (haar case be ake dell ea ee Dee Maden SE RE RUS
CLuver, M. A. & Horton, N.
The genera Dicynodon and Diictodon and their bearing on the classification of the
Dicynodontia (Reptilia, Therapsida). (Published February 1981.) ............
Cooper, M. R.
Revision of the Late Valanginian Cephalopoda from the Sundays River Formation
of South Africa, with special reference to the genus Olcostephanus. (Published
ALUVETIST JIG os See ota ae Pe erie ors ee ce ere ne Cea er ee
GriFFiTHS, C. L.
The freshwater Amphipoda (Crustacea) of South and South West Africa. (Pub-
Pieeem me RURAy AO Gilbert hen yeni Sted kein Cee Munem ow 20 valaw fae
Hortron, N. see CLUVER, M. A.
KENSLEY, B.
Decapod and isopod crustaceans from the west coast of southern Africa, including
Scamounts Vema and Tripp. (Published November 1980.).............+.2.--
KENSLEY, B.
The South African Museum’s Meiring Naude cruises. Part 12. Crustacea Decapoda
mmineio77, 1978, 1979 cruises. (Published February 1981.) ...........0:....4:
SHAUGHNESSY, P. D. see BEsT, P. B.
WINTERBOTTIOM, R.
A new genus and three new species of the family Congrogadidae (Pisces, Perci-
formes) from Natal, South Africa. (Published November 1980.) ..............

Page

147

NEW GENERIC NAMES PROPOSED IN THIS VOLUME

Natalichthys Winterbottom, 1980

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OF THE SOUTH AFRICAN
MUSEUM

CAPE ‘TOWN

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

FISCHER, PH. 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. Zen. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 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.

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 83 Band
November 1980 November
Part 1 = Deel

A NEW GENUS AND THREE NEW SPECIES OF THE
FAMILY CONGROGADIDAE (PISCES, PERCIFORMES)
FROM NATAL, SOUTH AFRICA

By |

RICHARD WINTERBOTTOM

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material
becomes available

Obtainable from the South African Museum, P.O. Box 61, Cape Town

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

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beskikbaarheid van stof

Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad

OUT OF PRINT/UIT DRUK

1, 2(1-3, 5-8), 3(1-2, 4-5, 8, t.-p.i.), 5(1-3, 5, 7-9),
6(1, t.—p.i.), 71-4), 8, 9(1-2, 7), 10(1-3),
11(1-2, 5, 7, t-—-p.i.), 15(4—-5), 24(2), 27, 3101-3), 32(5), 33

Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 003 6

Printed in South Africa by In Suid-Afrika gedruk deur
The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk.,
Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

A NEW GENUS AND THREE NEW SPECIES OF THE
FAMILY CONGROGADIDAE (PISCES, PERCIFORMES)
FROM NATAL, SOUTH AFRICA

By
RICHARD WINTERBOTTOM
Royal Ontario Museum, Toronto
(With 3 figures)
[MS. accepted 12 June 1980]

ABSTRACT

Examination of two lots and six specimens of a congrogadid from Natal which had been
identified as Halidesmus scapularis Giinther, 1871, has led to the proposal of a new genus,
Natalichthys, with three new species, N. leptus, N. ori and N. sam. The range of H. scapularis
is restricted to the coast between False Bay and the Umtata River mouth, Transkei.

CONTENTS

PAGE
Introduction . R 5 ‘ : : : 1
Methods : : : : ‘ : : 2
Descriptions . : : : x , : 2
Discussion : : : : . : : 11
Acknowledgements . : 2 : : ' i2
References. ; 3 : i : : ie

INTRODUCTION

Halidesmus scapularis is a well-known South African intertidal congrogadid,
ranging from False Bay to Coffee Bay, Transkei. Giinther (1871: 668, 669)
described the genus and species based on two specimens from Port Elizabeth,
adding in parentheses “Port Natal’. A misinterpretation of Port Natal for Natal
could explain why Gilchrist & Thompson (1917a: 220) placed H. scapularis
in Part 1 of their ‘A catalogue of sea fishes recorded from Natal’, and why, in
Part 2 (19176: 416) they recorded it as being from ‘(Port Natal)’. Later, how-
ever, Barnard (1927: 871) gave the locality for this species as ‘False Bay,
Algoa Bay, Natal coast, down to 50 fathoms’. It was presumably this notation
that was used by Smith (1961) and Smith & Smith (1966) to report the range
of H. scapularis as being from False Bay to Natal, although Smith (1952: 100),
in his earlier review of south and east African congrogadids, restricted the
range from False Bay to East London. A list of the South African Museum’s
holdings of H. scapularis included two lots with a total of six specimens from
off Natal. Extensive collecting by personnel from the J.L.B. Smith Institute of
Ichthyology during the last decade in the area between Port Elizabeth and

Ann. S. Afr. Mus. 83 (1), 1980: 1-12, 3 figs.

2 ANNALS OF THE SOUTH AFRICAN MUSEUM

Sodwana Bay, KwaZulu, failed to produce specimens of H. scapularis north
of the Umtata River mouth, Transkei. Therefore, the two lots from Natal
were of considerable interest, and were borrowed for examination. The six
specimens proved to represent a new genus comprised of three new species.

METHODS

Counts and measurements are conventional (Hubbs & Lagler 1964)
except that head length is measured from the tip of the snout to the tip of the
opercular spine, and only the pored lateral-line scales are counted. Since the
spine of the first dorsal fin is nearly or barely connected to the base of the
first dorsal-fin ray, the condition is expressed by a slash between spine and ray
counts (e.g. DI/42-43). The abbreviation SAM is an acronym for the South
African Museum, Cape Town.

DESCRIPTIONS

Natalichthys gen. nov.
Type species

Natalichthys ori sp. nov.

Diagnosis

The new genus is placed in the family Congrogadidae because, among
other reasons, it lacks anal spines, has a single dorsal spine not (or barely)
attached to the base of the first dorsal-fin ray, lacks palatine teeth and has a
single, spur-like opercular spine. Natalichthys primarily differs from other
congrogadid genera in a combination of characters. The following three
characters, taken together, will distinguish the genus from all other described
congrogadids: pelvic fin of one spine and two rays (I,2), gill membranes united
but free from isthmus, a single short lateral line.

Comparison

Only two other congrogadid genera possess a I,2 pelvic fin (Blennodesmus
and Halidesmus, both monotypic). Natalichthys differs from the eastern
Australian Blennodesmus in having the gill membranes free of the isthmus
(v. fused), and in the presence of a supraotic sensory canal pore (v. absent).
Halidesmus (which appears to be closely related to the monotypic Pholioides
from Pakistan and India) possesses at least three complete lateral-lines (v. a
single incomplete lateral-line), seven (v. six) preopercular sensory canal pores,
and a sensory canal pore between the first suborbital and first preopercular
sensory canal pores (vy. absent). Pholioides agrees with Halidesmus in all these
characters, but differs from that genus in lacking pelvic fins (v. present) and
in having eight (v. 10) pectoral rays. Anatomical studies presently being under-
taken will hopefully allow monophyletic supraspecific taxa to be proposed

NEW GENUS AND THREE NEW SPECIES OF FAMILY CONGROGADIDAE ~- 3

for the congrogadids, and this may result in some reduction of the number of
genera (presently ten genera, seventeen species, including this study).

Etymology

Named for Natal, the province off which all specimens were collected,
and ichthys, a fish. Gender: masculine.

KEY TO THE SPECIES OF NATALICHTH YS

la. DI/42, A 32, P 10, two dorsal and a single ventral procurrent caudal-fin rays...... N. sam
Pin CICMnOne vA oO? OF MOTE, PO... sc. kw cw eo ooh ech eee ee ciedbieeancnuucenewseceus 2
2a. DI/52-53, A 42-43, two dorsal and two ventral procurrent caudal-fin rays, cheeks naked

N. ori

2b. DI/48, A 39, a single dorsal and ventral procurrent caudal-fin ray, cheeks scaled N. Jeptus

Natalichthys ori sp. nov.
Fig. 1
Holotype

SAM-17340, 60,2 mm SL (67,8 mm TL), Umhlangakulu River, Natal,
South Africa (30°04'10’S 30°21'47”E ? ). No further data, but see discussion.

Paratype
SAM-28993, 53,7 mm SL (60 mm TL), collected with holotype.

Diagnosis

Differs from the other two species in the genus in the higher number of
dorsal- and anal-fin rays (DI/52-53, A 42-43 v. DI/42 or 48, A 32 or 39). It
can further be separated from N. sam in having one less pectoral-fin ray (nine
v. ten), two ventral procurrent caudal-fin rays (v. one), more lower gill rakers
(seven to eight v. five), and the absence of vomerine teeth (v. present). Additional
characters separating N. ori from N. leptus include the first ray of the dorsal
fin unbranched (v. branched), two dorsal and ventral procurrent caudal-fin
rays (v. one each) and the absence of scales on the cheek (v. present).

Description

A small (60,2 mm maximum recorded SL) congrogadid known only from
two specimens collected off Natal. The following.counts and measurements
are given for the holotype, with values for the paratype in parentheses where
different. As per cent standard length: soft dorsal-fin base 77,2 (78,2); anal-fin
base 59,6 (59,4); snout tip dorsal-fin spine origin 21,1 (20,9); snout tip to first
dorsal-fin ray origin 23,6 (23,1); snout tip to first anal-fin ray origin 40,9 (39,5);
head length 15,0 (15,6). As per cent head length: head depth at parietal commis-
sure 51,1 (52,4), body depth at anal-fin origin 50,0 (51,2); eye diameter 25,6

ANNALS OF THE SOUTH AFRICAN MUSEUM

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NEW GENUS AND THREE NEW SPECIES OF FAMILY CONGROGADIDAE. 5

(26,2); snout length 22,2 (23,8); bony interorbital 7,7 (7,1); upper-jaw length
37,8 (38,1); lower-jaw length 54,4 (52,4); pectoral-fin length 44,4 (45,2); pelvic-
fin length 23,3 (25,0); length of first dorsal-fin ray 24,4 (27,4); tenth 38,9 (44,0);
twentieth 50,0 (46,4); thirtieth 48,9 (47,6); fortieth 47,8 (50,0); penultimate
47,8 (50,0); length of first anal-fin ray 22,2 (23,8); tenth 33,3 (35,7); twentieth
34,4 (36,9); thirtieth 35,6 (38,1); penultimate 38,9 (44,0).

Dorsal fin I/52-53, first five rays of holotype and first ray only of paratype
unbranched; A 42-43, first ray of holotype and no rays of paratype unbranched;
pectoral fin 9; pelvic fin I,2, spine rudimentary; caudal fin with two dorsal
procurrent rays, the posteriormost ray with a few segmentations, five dorsal
and five ventral principal rays (the dorsalmost and ventralmost unbranched
in the holotype, branched in the paratype), two ventral procurrent rays, the
posteriormost ray with a few segmentations in the paratype but not in
the holotype. Caudal fin connected to dorsal and anal fins by a membrane
which reaches about half-way along the length of the last dorsal- and anal-fin
rays.

Sensory canal openings (Fig. 1B—all pores bilateral except where other-
wise stated): nasal double, with one just posterior to upper lip and the other
just behind posterior nostril; an anterior interorbital and a single, median
posterior interorbital; a supraotic (absent on left side of paratype); eight
suborbitals; six in preopercular canal; four in dentary. An intertemporal;
anterior and posterior post-temporal; and two parietal pores.

Gill membranes fused to each other in ventral midline, but free from
isthmus. Six branchiostegal rays; gill rakers on first arch with two to three
epibranchial, one in angle, and seven to eight ceratobranchial = (2—3) + 1 +
(7—8). A single, short, incomplete lateral line ending beneath the third ray of
soft dorsal fin, consisting of 22-23 pored scales. Olfactory capsule with two
nostrils, anterior a short tube, posterior pore-like. Cheeks and opercles naked,
body scaled almost to parietal commissure with small, elliptical, cycloid scales.
Pseudobranch with five lobes.

Vomer and palate edentate. Jaw teeth conical, slightly recurved, decreasing
in size posteriorly. A short inner row of about four teeth in upper jaw, two
in lower jaw.

Colour pattern (alcohol-preserved specimens)—plain yellow-brown with
a diffuse, somewhat elliptical dark blotch (about half eye diameter) on shoulder
above opercle. The blotch was apparently unocellated—although the material
appears faded.

Etymology

The specific name ori (treated as a noun in apposition) is the acronym of
the Oceanographic Research Institute, Durban, whose Directors (Drs D. H.
Davies and A. E. F. Heydorn) and staff have contributed greatly to the know-
ledge of South Africa’s coastal marine fauna.

6 ANNALS OF THE SOUTH AFRICAN MUSEUM

Natalichthys sam sp. nov.
Fig. 2

Holotype

SAM-21915, 42,7 mm SL (48,5 mm TL), Indian Ocean, off Port Shepstone,
south coast of Natal, South Africa (30°47'06’S 30°29'06’E), 44 m, stony
bottom. Collected by University of Cape Town (NAD 2x), 17 May 1958.

Paratype
SAM-28940, 40,3 mm SL (44,5 mm TL), collected with holotype.

Diagnosis

Differs from the other two species in the genus in the lower number of
dorsal and anal-fin rays (DI/42, A 32 v. DI/48-53, A 39-43), one more pectoral-
fin ray (10 v. 9), and four to six teeth on the vomer (v. two or none). It can be
further separated from N. ori in having a single ventral procurrent caudal-fin
ray (v. two), and fewer lower gill rakers on the first gill arch (five v. seven to
eight). Additional characters separating N. sam from N. leptus include first
ray of dorsal fin unbranched (v. branched), one more dorsal procurrent caudal-
fin ray (two v. one) and naked cheeks (v. scaled cheeks).

Description

A short, small (42,7 mm maximum recorded SL) congrogadid known
only from two specimens collected off Natal. The following counts and measure-
ments are given for the holotype, with values for the paratype in parentheses
where different. As per cent standard length: soft dorsal-fin base 73,1 (69,5);
anal-fin base 55,3 (51,1); snout tip to dorsal-fin spine origin 23,4 (26,1); snout
tip to first dorsal-fin ray origin 26,0 (27,8); snout tip to first anal-fin ray origin
44,5 (46,4); head length 18,3 (18,9). As per cent head length: head depth at
parietal commissure 59,0 (56,6); body depth at anal-fin origin 57,7 (59,2);
eye diameter 25,6 (25,0); snout length 21,8 (22,4); bony interorbital 7,7 (6,6);
upper-jaw length 38,5 (34,2); lower-jaw length 51,3 (53,9); pectoral-fin length
53,8 (50,0); pelvic-fin length 24,4 (26,3); length of first dorsal ray 26,9 (26,3);
tenth 44,9 (44,7); twentieth 47,4; thirtieth 51,3; penultimate 52,6; length of
first anal-fin ray 26,9 (22,4); tenth 38,5 (36,8); twentieth 39,7 (39,5); penultimate
51,3 (42,1).

Dorsal fin 1/42, only first ray unbranched; A 32, all rays branched; pectoral
fin 10, pelvic fin I,2, spine rudimentary; caudal fin of two procurrent and five
principal dorsal rays, and one procurrent and six principal ventral rays. The
second (more posterior) dorsal procurrent ray has a few segmentations near its
tip in the paratype, but is unsegmented in the holotype. Caudal fin connected
to dorsal and anal fins by a membrane which reaches to about the midpoint of
the length of the last dorsal- and anal-fin rays.

NEW GENUS AND THREE NEW SPECIES OF FAMILY CONGROGADIDAE

Fig. 2. A. Natalichthys sam, \eft lateral view of holotype (42,7 mm SL,
SAM-21915). B. N. sam (holotype), left lateral view of head to show
sensory canal openings. Abbreviations as for Figure 1. Drawn by A. Odum.

POP 1-6

8 ANNALS OF THE SOUTH AFRICAN MUSEUM

Sensory canal openings (Fig. 2B—all pores bilateral except where stated
otherwise): nasal double, with one just posterior to the upper lip and the other
just behind the posterior nostril; an anterior interorbital and a single, median
posterior interorbital; a supraotic; eight suborbitals; six in preopercular canal;
four in dentary. An intertemporal; anterior and posterior posttemporal; and
two parietal pores. In addition, an extra pore between the dorsalmost pre-
opercular pore and the intertemporal pore is present on the left, but not the
right, sides of both specmens. |

Gill membranes fused to each other in ventral midline, but free from
isthmus. Six branchiostegal rays; gill rakers on first arch with two epibranchial,
one in angle and five ceratobranchial = 2 + 1 + 5. A single, short, incomplete,
lateral line ending beneath the sixth dorsal-fin ray, consisting of 22-27 pored
scales. Olfactory capsule with two nostrils, anterior a short tube, posterior
pore-like. Cheeks and opercles naked, body scaled up to the parietal commissure
with small, elliptical cycloid scales. Pseudobranch short, with five lobes.

A few (four to six) conical teeth on vomer, none elsewhere on palate. Jaw
teeth conical, slightly curved, anterior largest, decreasing irregularly in size
posteriorly. Inner patch of small teeth behind symphysis in both Jaws, an
inner row continuing posteriorly half as far as the outer row.

Colour pattern (alcohol-preserved specimens)—plain yellow-brown with
a somewhat elliptical dark blotch (about three-quarters eye diameter) on
shoulder above opercle. The blotch was apparently unocellated (ocellated in
many congrogadids), but the material is somewhat faded.

Etymology

The specific name sam (treated as a noun in apposition) is fie acronym
for the South African Museum, Cape Town, whose ichthyologists, Dr P. A.
Hulley and Miss E. Louw, have always been most co-operative and good
company.

Natalichthys leptus sp. nov.
Fig. 3

Holotype

SAM-28938, 57,2 mm SL (63,9 mm TL), Umblaasaieal River, Natal,
South Africa (30°04'10’S 30°21’47"E ? ). No further data, but see discussion.

Paratype
SAM-28939, 49,7 mm SL (56,9 mm TL), collected with holotype.

Diagnosis

Differs from the other two species in the genus in fin-ray counts (DI/48,
A 39 v. DI/42 or 52-53, A 32 or 42-43), in having the first dorsal-fin ray
branched (v. unbranched), in a single dorsal procurrent caudal-fin ray (v. two),

NEW GENUS AND THREE NEW SPECIES OF FAMILY CONGROGADIDAE

133
NIH

ail Wy} i:

yw aly i,
Se Lye

1. Drawn by A. Odum.

left lateral view of head to show

hthys leptus, \eft lateral view of holotype (57,2 mm SL,

ic
SAM-28938). B. N. leptus (holotype),

sensory canal openings. Abbreviations as for Figure

~—s

_—

=
a
©
Lt
a
re)
jo
£
Ele

rio z
Ww
a

SOB 1-8

10 ANNALS OF THE SOUTH AFRICAN MUSEUM

and in possessing scales on the cheeks. It differs from N. sam in having nine
(v. ten) pectoral-fin rays, and in having two or no vomerine teeth (v. four to
six). Additional differences from WN. ori include a single ventral procurrent
caudal-fin ray (v. two) and a lower number of gill rakers on the lower limb
(five to six v. seven to eight).

Description

A small (57,2 mm SL maximum recorded SL) congrogadid known only
from two specimens collected off Natal. The following counts and measure-
ments are given first for the holotype, with values for the paratype in paren-
theses where different. As per cent standard length: soft dorsal-fin base 69,2
(74,8); anal-fin base 55,6 (59,6); snout tip to dorsal-fin spine origin 23,0 (22,5);
snout tip to first dorsal-fin ray origin 25,3 (25,2); snout tip to first anal-fin
origin 41,4 (41,6); head length 17,5 (16,9). As per cent head length: head depth
at parietal commissure 54,0 (54,8); body depth at anal-fin origin 49,0 (53,8);
eye diameter 26,0 (26,2); snout length 25,0; bony interorbital 7,0 (7,1); upper-
jaw length 36,0 (38,1); lower-jaw length 55,0 (54,8); pectoral-fin length 53,0
(51,2); pelvic-fin length 21,0 (25,0); length of first dorsal-fin ray 21,0 (23,8);
tenth 45,0 (42,6); twentieth 47,0 (46,4); thirtieth 54,0 (53,6); fortieth 55,0
(53,6); penultimate 46,0 (54,8); length of first anal-fin ray 21,0 (25,0); tenth
29,0 (34,5); twentieth 32,0 (38,1); thirtieth 35,0 (41,7); penultimate 40,0 (47,6).

Dorsal fin I/48, all rays branched; A 39, all rays branched; pectoral fin 9;
pelvic fin I,2, spine rudimentary; caudal fin with one dorsal procurrent ray,
five branched principal dorsal and five branched principal ventral rays, and a
single ventral procurrent ray. Both dorsal and ventral procurrent rays with a
few striations in holotype, no striations in paratype. Caudal fin connected to
dorsal and anal fins by a membrane which reaches about one-third along the
length of the last dorsal- and anal-fin rays.

Sensory canal openings (Fig. 3B—all pores bilateral except where other-
wise stated): nasal double, with one just posterior to upper lip and the other
just behind posterior nostril; an anterior interorbital, and a single, median
posterior interorbital (paratype only); a supraotic; eight suborbitals; six in
preopercular canal; four in dentary. An intertemporal; anterior and posterior
posttemporal; and two parietal pores.

Gill membranes fused to each other in ventral midline, but free from
isthmus. Six branchiostegal rays, gill rakers on first arch with two to three epi-
branchial, one in angle and five to six ceratobranchial (= 2 — 3 + 1 + 5 — 6).
A single, short, incomplete lateral line ending beneath the third to fourth ray
of soft dorsal fin; consisting of 20-24 pored scales. Olfactory capsule with two
nostrils, anterior a short tube, posterior pore-like. Cheeks, but not opercles,
with small, elliptical, cycloid scales, body scaled to parietal commissure with
similar scales. Pseudobranch with six lobes in holotype, five in paratype.

A single minute tooth on either side of the head of the vomer in paratype,
none in holotype; rest of palate edentate. Jaw teeth stoutly conical, decreasing

NEW GENUS AND THREE NEW SPECIES OF FAMILY CONGROGADIDAE - 11

in size posteriorly. A short inner row of five teeth in upper jaw, three in lower
jaw.

Colour pattern (alcohol-preserved specimens)—plain yellow-brown, with
a diffuse, elliptical dark blotch (about three-quarters eye diameter) on shoulder
above opercle. Blotch apparently unocellated.

Etymology

From the Greek /eptos, like a scale or peel, thin, fine, small, delicate; in
allusion to the scaled cheeks of the new species. Treated as a noun in apposition.

DISCUSSION

Two of the species, N. ori. and N. leptus, were in the same lot. The only
data accompanying the lot consisted of the catalogue number, and ‘Umhlanga-
kulu River, Natal’ Congrogadids have not previously been recorded from fresh
or estuarine waters. It seems probable that these specimens were collected off
the mouth of the river, in the sea. Additional circumstantial evidence for this
comes from Barnard’s statement (1927: 871) that Halidesmus scapularis had
been collected from the Natal coast at 50 fathoms. This specific statement of
depth and place indicates that he had specimens with that data. There are no
specimens of H. scapularis from that depth and locality at the South African
Museum (where Barnard worked). The South African Museum specimens
labelled as H. scapularis from Natal represent the material forming this paper.
The specimens here described as N. sam were collected in 1958 (31 years after
publication of Barnard’s monograph), and therefore cannot represent his
material. Thus it is probable that the lot on which Barnard based his statement
of range and depth is the one now labelled as being from Umhlangakulu River.
The original catalogue entry for this lot (SAM-—17340) is in Barnard’s hand-
writing (E. Louw, pers. comm.).

A further point is of interest here. Although Gilchrist & Thompson’s 19175)
statement of Port Natal as a locality for H. scapularis appears to be taken
directly from Giinther’s (1871) description of the types (see introduction), it is
possible that they may have had additional specimens identified as H. scapularis
from Natal. In this context, P. Heemstra (pers. comm.) informed the author
that there is an ‘Umhlangankulu’ estuary served by a short (6,5 km) river
located at 30°56’45”S 30°18’E, which thus lies some 34 km south-west of Port
Shepstone. P. A. Hulley (pers. comm.) searched the original catalogues of the
R.S. Pieter Faure and found the following: ‘14 March, 1901. Umhlangakulu
[sic] River NW by N, 74 miles. Dredge, 12.50-1.00 p.m.’. The co-ordinates for
this station would then be 31°04’10’S 30°21’47’E. The evidence, although
circumstantial, appears to indicate strongly that N. ori and N. leptus were taken
at this station.

In summary, then, it appears probable that (i) Gilchrist & Thompson’s
(19176) record of H. scapularis was taken directly from Giinther’s description

12 ANNALS OF THE SOUTH AFRICAN MUSEUM

and (ii) that the specimens of N. ori and N. leptus were collected at 30°04’10’S
30°21'47”E off the Natal south coast near Port Edward at a depth of 90m
(50 fm.) on 14 March 1901.

ACKNOWLEDGEMENTS

I thank the South African Museum for allowing me to report on the
specimens and Dr P. Hulley and Miss E. Louw for the information on the
Faure stations, catalogue data and geography. Dr P. Heemstra of the J.L.B.
Smith Institute of Ichthyology, Grahamstown, kindly sent me the co-ordinates
for the Umhlangankulu River. Drs E. J. Crossmand and A. R. Emery of the
Royal Ontario Museum commented on the manuscript, and Mr Anker Odum
(ROM) expertly prepared the figures for me.

REFERENCES

BARNARD, K. H. 1927. A monograph of the marine fishes of South Africa. Part II. Ann.
S. Afr. Mus. 21: 419-1065.

GILcurisT, J. D. F. & THompson, W. W. 1917a. A catalogue of the sea fishes recorded from
Natal. Part I. Ann. Durban Mus. 1(4): 255-290.

GILCHRIST, J. D. F. & THomMPpson, W. W. 1917b. A catalogue of the sea fishes recorded from
Natal Part II. Ann. Durban Mus. 1(4): 291-431.

GUNTHER, A. 1871. Report on several collections of fishes recently obtained for the British
Museum. Proc. zool. Soc. Lond. 1871: 652-675.

Husss, C. L. & LAGLER, K. F. 1964. Fishes of the Great Lakes Region. Ann Arbor: University
of Michigan Press.

SMITH, J. L. B. 1952. The fishes of the family Haliophidae. Ann. Mag. nat. Hist. (12) 5: 85-101.

SMITH, J. L. B. 1961. The Sea Fishes of Southern Africa. 4th ed. South Africa: Central News
Agency.

SmiTH, J. L. B. & SmitH, M. M. 1966. Fishes of the Tsitsikama Coastal National Park. South
Africa: National Parks Board of Trustees.

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6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.

An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15SA
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:
comma separates author’s name and year
“semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.

In describing new species, One specimen must be designated as the holotype; other 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°51’S 25°39’E), collected by A. ‘Smith, 15 January 1973.

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

7. SPECIAL HOUSE RULES

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

’ .

e.g. . the Figure depicting C. namacolus ...’; *. . . in C. namacolus (Fig. 10) .

*(b) The prefixes of prefixed surnames in all languages, whet used in the text, if not preceded
by initials or full names
e.g. DuToit but A.L.du Toit; Von Huene but F. von Huene

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.

Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.

RICHARD WINTERBOTTOM

A NEW GENUS AND THREE NEW SPECIES OF
THE FAMILY CONGROGADIDAE (PISCES,
PERCIFORMES) FROM NATAL, SOUTH AFRICA

OF THE SOUTH AFRICAN
MUSEUM

CAPE ‘TOWN

INSTRUCTIONS TO AUTHORS

1. MATERIAL should be original and not published elsewhere, in whole or in part.
2. LAYOUT should be as follows:

(a) Centred masthead to consist of
Title: informative but concise, without abbreviations and not including the names of new genera or species
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The number of the figure should be lightly marked in pencil on the back of each illustration.

5. REFERENCES cited in text and synonymies should all be included in the list at the end of
the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable):

(a) Author’s name and year of publication given in text, e.g.:

‘Smith (1969) describes...’

‘Smith (1969: 36, fig. 16) describes...’

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“As described (Haughton e¢ al. 1927)...’

Note: no comma separating name and year
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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. gen. 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.

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 83 Band
November 1980 November
Part 2 °#£Deel

DECAPOD AND ISOPOD CRUSTACEANS FROM THE
WEST COAST OF SOUTHERN AFRICA, INCLUDING
SEAMOUNTS VEMA AND TRIPP

BRIAN KENSLEY

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material
becomes available

Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000

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word uitgegee in dele op ongereelde tye na gelang van die
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Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad 8000

OUT OF PRINT/UIT DRUK

1, 2(1=3, 5-8), 31-2, 4-5, 8; tpi), 5(t-3; 57-9
6(1, t.-p.i.), 71-4), 8, 9(1-2, 7), 10(1-3),
11(1-2, 5, 7, t--p.i.), 15(4-5), 24(2), 27, 31(1-3), 32(5), 33

Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 002 8

Printed in South Africa by In Suid-Afrika gedruk deur
The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk.,
Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

DECAPOD AND ISOPOD CRUSTACEANS FROM THE WEST COAST
OF SOUTHERN AFRICA, INCLUDING SEAMOUNTS VEMA AND
TRIPP

By
BRIAN KENSLEY

Smithsonian Institution, Washington, D.C.
(With 9 figures)

[MS. accepted 15 July 1980]

ABSTRACT

Seven species of isopods (including Stenetrium vemae sp. nov. and Jaeropsis monsmarinus
sp. nov.) and nineteen species of decapods (including Pseudodromia cacuminis sp. nov. and
Macropodia cirripilus sp. nov.) are recorded from seamounts Vema and Tripp, and the Liideritz
area. Zoogeographically, the isopods show strong affinities with the South African fauna,
while the decapods include mainly South African and west African forms, with single Indo-
Pacific and Austral species.

CONTENTS

PAGE
Introduction . : : : : : : 13
Systematic discussion : : . ‘ ‘ 15
Isopoda . ; ‘ é : : ‘ 15
Decapoda : ‘ : ‘ : ‘ 21
Zoogeographic discussion : ; : : 29
Acknowledgements . : : ‘ : ‘ 31
References. 2 : : : : é 31
INTRODUCTION

The benthic fauna of the continental shelf and seamounts off the west
coast of South Africa has barely been investigated. What information exists
is to be found in scattered reports, and we are still a long way from even a
superficial overview.

The material dealt with in this report comes from several sources and
emphasizes the fragmentary state of our knowledge. It was thought useful,
however, to publish several new records and species, and to summarize the
little that is known about Seamount Vema’s crustacean fauna.

_ Seamount Vema, first discovered in 1957, was visited by personnel from
the University of Cape Town and the South African Museum in 1964 and
1966. In 1978 the University of Cape Town did further collecting on the summit
peak. Lying about 650 km off the west coast of South Africa at 31°38’S 08°02’E
(Fig. 1), and rising steeply from the 5 000 m deep sea-floor, the summit plateau
averages about 40 m below the surface. Collecting on this plateau has been
done both with air-lift dredge and by scuba divers (see Grindley 1967). Most

Ann. S. Afr. Mus. 83(2), 1980: 13-32, 9 figs.

35°

14 ANNALS OF THE SOUTH AFRICAN MUSEUM

Lideritz

Atlantic Ocean

e
302 Tripp Seamount

e@
Vema Seamount

wCape Town

5° 10° 15° 20°

Fig. 1. Map showing localities.

of this material has been deposited in the South African Museum, while a
preliminary account of the fauna was given by Berrisford (1969). The Decapoda
in this latter report were given preliminary identifications by J. Forest and
D. Guinot of the Paris Museum.

Material from Seamount Tripp was collected in the late 1960s by the
then Division of Sea Fisheries and the South African Museum, and a few
specimens came from commercial fishing boats on the west coast. Seamount
Tripp (20°36’S 14°15’E) has received even less attention than Vema, the three
specimens mentioned here having been accidentally caught during hydro-
graphic operations. The summit is about 150 m below the sea surface.

Abbreviations used

CL—carapace length

CW —carapace width

IK —Isaacs Kidd trawl

juv.—juvenile(s)

ovig.— ovigerous

SAM-—South African Museum

USNM-— United States National Museum
VEM—Vema station numbers

DECAPOD AND ISOPOD CRUSTACEANS 15

SYSTEMATIC DISCUSSION

Order ISOPODA
SPECIES LIST
Family Idoteidae Material Station No. Locality Distribution
Glyptidotea lichtensteini(Krauss) 2 juvs VEM 2.3 Vema, 39m _ £Liideritz to Transkei
3 juvs VEM 4.3 Vema, 40 m
Paridotea ungulata (Pallas) 1¢3 VEM 2.2 Vema, 39m Walvis Bay to East London;
Australia; New Zealand; Chile;
Argentina
Family Cirolanidae
Cirolana saldanhae Barnard 1 ovig. 2 VEM 2.3 Vema, 39 m ishee River mouth to Saldanha
ay
1 damaged
1 juv. VEM 4. Vema, 40 m

19 VEM 4. Vema, 42 m

3
6
Family Sphaeromatidae
VEM AS Vema,42m_ Liideritz to False Bay

Cymodoce unguiculata Barnard 192
Cymodocealla sublevis Barnard 12 VEM 4 Vema,40m_ Liideritz to East London
Family Stenetriidae :
Stenetrium vemae sp. nov. 2 ovig. 2 VEM 2.1 Vema,39m —
2 ovig. 2 VEM 2.2 Vema, 39 m
2236
4 ovig. 2 VEM 2.3 Vema, 39 m
4°86
13 VEM 4.1 Vema, 40 m
12 VEM 4.2 Vema, 40 m
1¢ VEM 4.6 Vema, 42 m
Family Jaeropsidae
Jaeropsis monsmarinus sp. Nov. 121¢ VEM 4.3 Vema,40m —

Family Stenetriidae

Stenetrium vemae Sp. nov.
Figs 2-3
Description

Male

Body about three and one-half times longer than wide, with scattered setae
dorsally. Cephalon broader than long, with well-developed dorsal reniform
eyes; anterolateral corners produced, acute; antennal spine of frontal margin
triangular, acute; rostrum wider than long, pentagonal, two anterior margins
with tiny teeth, apex an obtuse angle somewhat dorsally flexed. Pereonites
1-4 with anterolateral corners acute, posterolateral corners rounded; mid-
ventral keel hardly developed, with tiny denticle on pereonites 1-3, absent
on 4; pereonites 5—7 with anterolateral corners rounded, posterolateral corners
of 5 rounded, 6 bluntly produced, 7 acute; midventral keel with strong posterior
spine on 6 and 7. First pleonite short, reduced; pleotelson wider than long,
with single strong lateral tooth in posterior half of margin, followed by sinuous
margin leading to rounded apex; middorsal region gently convex, barely
demarked from lateral regions.

Antennular peduncle 3-segmented, basal segment broader and longer
than two distal segments, second segment shorter than third, bearing elongate
simple setae; flagellum of 29-30 articles. Basal antennal segment produced
into spinose process on outer distal angle; second segment shorter than first;
third segment outer distal margin deeply excavate for insertion of large setiferous
scale; fourth segment less than half length of third. Mandibular palp
3-segmented, basal segment with single, strong fringed seta; second segment

16 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 2. Stenetrium vemae.
A. Holotype in dorsal view. B. Maxilliped. C. Maxilla 2. D. Mandible. E. Maxilla 1.

DECAPOD AND ISOPOD CRUSTACEANS : 17

Fig. 3. Stenetrium vemae.
A. Pereopod 2. B. Antennular peduncle. C. Operculum 2. D. Pleopod 1 g. E. Pleopod 3 ¢.
F. Pleopod 2 3.

with row of short spines and single, strong fringed seta; terminal segment
strongly curved, with row of spines on inner margin, several elongate setae
on distal narrowed part; left mandible with incisor of four large cusps, sclero-
tized lacinia of two strong cusps and serrate spine; spine row of six serrate
spines; molar bearing short marginal spines, roughened distal surface; right
mandible lacking lacinia; spine row of sixteen serrate spines. Maxilla 1 outer
ramus with eleven strong dentate spines; inner ramus distally with two strong
and two slender setae. Maxilla 2 both lobes of outer ramus each with five
elongate fringed spines; inner ramus with eight fringed spines and several
setae. Maxilliped exopod apically acute; palp of five segments, each with
numerous simple setae; endite with seven coupling hooks on median margin,
several fringed spines and seven or eight flattened fringed scales. Pereopod
1 almost equal to entire body in length; dactylus strongly curved, longer than
propodal palm; propodus widening distally, palm with strong tooth at about

18 ANNALS OF THE SOUTH AFRICAN MUSEUM

midlength, outer half of palm somewhat produced into lobe bearing two strong
teeth separated by three tiny teeth; carpus, merus, and ischium together
shorter than propodus, merus and ischium with triangular acute process on
upper margin; basis elongate-cylindrical. Pereopods 2-7 similar, ambulatory;
dactylus biunguiculate; propodus and carpus with slender spines on ventral
margin. Pleopod | rami distally rounded-truncate, with short setae. Remaining
pleopods typical of genus. Uropod with basis shorter than rami; latter terete,
setose, inner ramus longer than outer. .

Female

Pereopod 1 strongly setose, much shorter than in male; dactylus with
row of short serrate spines on cutting edge; propodal palm with row of curved
serrate spines. Operculum triangular, lateral margins slightly sinuous, apex
narrowly rounded.

Material

Holotype SAM-A16780 ¢ TL 7,5 mm VEM 2.2
Paratypes SAM-A16781 4 ovig. &, 4 9, 8 6 VEM 2.3
Paratypes USNM 173516 2 ovig. 9, 1 9, 2 ¢ VEM 2.2
Additional material SAM-A16782 2 ovig. 9 VEM 2.1
SAM-A16783 1 3 VEM 4.1
SAM-A16784 1 9 VEM 4.2
SAM-A16785 1 ¢ VEM 4.6
Remarks
Using Wolff’s (1962: 22) key to the genus Stenetrium the present material
can be run down to S. diazi Barnard, 1920, from South Africa. The two species
are undoubtedly similar, especially in the elongate form and structure of
pereopod 1 of the mature male. Several differences separate the two species:
the rostrum of S. diazi has a concave anterior margin, the rami of pleopod
1 3 are acutely rounded on the inner distal angle, while pereopod 1 d, although
having a somewhat elongate propodus, does not have the outer distal lobe
bearing two large teeth separated by three tiny teeth; instead it possesses two
blunt, closely opposed teeth near the articulation.

Etymology
The specific name is derived from the type locality, Seamount Vema.

Family Jaeropsidae

Jaeropsis monsmarinus sp. nov.
Figs 4—5
Description
Male
Body about three times longer than wide, with numerous scattered, short
setules dorsally. Cephalon with lateral margins entire, apex rounded.

DECAPOD AND ISOPOD CRUSTACEANS 19

gae=-—

os
NY

Fig. 4. Jaeropsis monsmarinus.
A. Holotype in dorsal view. B. Antennule. C. Maxilliped. D. Antenna. E. Maxilla 1.
F. Mandible. G. Maxilla 2.

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

Antennule with broad basal segment equal in length to following six
segments; three distal articles bearing aesthetascs. Antennal peduncle
5-segmented; two basal segments short; flagellum of nine very short articles.
Mandibular palp 3-segmented, two distal segments bearing several fringed
setae; incisor of six cusps; spine row of nine strong spines; molar elongate-
slender with tiny denticles on lower margin. Maxilla 1 outer ramus with twelve
strong serrate spines; inner ramus with four distal setae. Maxilla 2, two outer
lobes each with four elongate fringed spines; inner lobe with three distal simple
setae. Maxillipedal palp 5-segmented, segments 1 and 2 broad, three distal seg-
ments narrow; endite broad, inner distal angle excavate, with strong delimiting
spine, four flattened denticles, and two short fringed spines; median margin with
three coupling hooks. Pereopod 1 dactylus biunguiculate, remaining pereopods
triunguiculate. Pleopod 1 two rami fused for two-thirds of length, terminal
setose part narrowly triangular. Uropod with apically rounded hook on inner
distal angle of basis; reduced rami bearing elongate simple setae.

Material

Holotype SAM-A 16786 ¢ TL 3,1 mm VEM 4.3
Allotype USNM 173517 2 TL 3,0 mm VEM 4.3

Fig. 5. Jaeropsis monsmarinus.
A. Pleopod 1 ¢. B. Pleopod 2 g. C. Pleopod 3 2. D. Uropod. E. Pereopod 2.

DECAPOD AND ISOPOD CRUSTACEANS ; 21

Remarks

The present species belongs to the groups of species possessing well-
separated obliquely-inserted uropods (Barnard 1965: 200), which includes
J. stebbingi Kensley, 1975, J. paulensis Vanhoéffen, 1914, and J. waltervadi
Kensley, 1975. Both J. waltervadi and J. stebbingi each possess a pleon with
serrate margins. J. monsmarinus most closely resembles J. paulensis, especially
in rostral shape and mouthparts. VanhOffen’s species, however, does not have
a hook on the inner distal angle of the uropod, while the triangular terminal
part of the pleopod 1 ¢ is broader than in the present species. Considering
the isolated nature of Vema, J. monsmarinus possibly represents a population
of J. paulensis (known from Gough, St Paul and Amsterdam Islands) which
has become genetically isolated.

Etymology
The specific name is the Latin for ‘seamount’.

Order DECAPODA
SPECIES LIST
Material Station No. Locality Distribution

* material not seen

Family Penaeidae

Funchalia villosa (Bouvier) 11372 Vema, from eastern and western North
40 juv. tuna stomach Atlantic, Caribbean, South

and Central Atlantic to Natal
Family Oplophoridae

Notostomus auriculatus Barnard 192 IK 52 Vema off Cape Point
Family Alpheidae
* Alpheus macrocheles (Hailstone) Vema Mediterranean, Great Britain,
Antilles, Guinea, Sao Tome
Synalpheus huluensis africanus 5 $5ovig.2 VEM2.3 Vema, 39 m Guinea, Sao Tome, Cape
Crosnier & Forest 12 juv. Verde Is., Principe, Annobon
19 VEM 4.6 Vema, 42 m
Family Hippolitidae
Eualus ctenifera (Barnard) 13 VEM 2.2 Vema, 39m Port Elizabeth to Natal,
113162 Walter’s Shoal
1 ovig. 2
ile VEM 3.2 Vema, 48 m
19 VEM 3.3 Vema, 50m

Family Crangonidae
* Pontophilus sculptus (Bell) Vema False Bay to Durban, Mediter-
ranean, North Atlantic

Family Palinuridae
Jasus tristani Holthuis Vema Tristan da Cunha
Family Paguridae
* Pagurus chevreuxi Bouvier Vema Mediterranean
Pagurus cuanensis (Bell) 13 VEM 4.5 Vema, 42 m False Bay to Port Elizabeth,
North Atlantic, west Africa,
Mediterranean
Family Galatheidae
Eumunida picta Smith 19 Off Liideritz North-western Atlantic, Cuba,
1312 Seamount Tripp Elonca, New Zealand, Austra-
ia
Galathea sp. 13 VEM 2.3 Vema, 39 m
Family Lithodidae
Lithodes murrayi Henderson 23 Off Liideritz St Paul and Amsterdam Is.
; 33 Off South West Prince Edward Is., Crozet Is.,

Africa off Natal
Family Dromiidae

Pseudodromia cacuminis sp.nov. 1¢ VEM 3.2 Vema, 48 m
13 VEM 4.2 Vema, 40 m
192 VEM 4.4 Vema, 40 m
19 VEM 4.6 Vema, 42 m
Family Homolidae
Paromola alcocki (Stebbing) 1° Off Liideritz Port Elizabeth, Mozambique,
800 m Maldives
Paromola cuvieri (Risso) 13 Off Liideritz, eastern North Atlantic, Medi-
800 m terranean, west Africa

1 ovig. 2 Seamount Tripp

22 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material Station No. Locality Distribution
Family Marjidae
Macropodia cirripilus sp. nov. 192 VEM 4.4 Vema, 40 m
13 VEM 15M _ Vema, 40m
Family Xanthidae
Pilumnus sp. 1 3 4 juv. VEM 2.3 Vema, 39 m
1¢3 VEM 3.3 Vema, 50m
* Pseudactaea corallina (Alcock) Vema Indo-Pacific
Family Grapsidae
Plagusia chabrus (Linnaeus) 1 3 1 ovig. 2 Vema South West Africa to Natal,

Chile, Juan Fernandez, Austra-
lia, New Zealand

Family Lithodidae.
Lithodes murrayi Henderson

Lithodes murrayi Henderson, 1888: 43, pl. 4. Hale, 1941: 272, pl. 3 (figs 3-4). Yaldwyn &
Dawson, 1970: 275, figs 1-3. Arnaud, 1971: 167; Kensley, 1977: 166, fig. 3.

Previous records

Possession Is., Prince Edward Is. 620 m; Macquarie Is. 120 m; Crozet Is.,
New Zealand, 764 m; Zululand to Durban, South Africa, 600-810 m.

Material

SAM-A 16206 3 CL 89 mm CW 90 mm off Liideritz, 800 m

SAM-—A 15358 3 CL 101 mm CW 102 mm off Liideritz, 800 m |

SAM-A 16211 3 gg CL 109-118 mm CW 106-120 mm off South West
Africa

Remarks

These first Atlantic records of L. murrayi represent a considerable extension
in the range of what was regarded as a southern Indian Ocean species.

Family Dromiidae
Pseudodromia cacuminis sp. nov.
Figs 6-7

Description
Female

Carapace, abdomen, and pereopods covered with short spiky hairs,
becoming dense in supraorbital, rostral, and abdominal margins. Carapace
wider than long, dorsaJly convex; front bluntly trilobed, median lobe set at
lower level than lateral lobes, ventrally keeled, dorsally visible; single, rounded
lateral lobe present; anterolateral and supraorbital areas bearing numerous
short spines; scattering of tiny spines in rostral area. Abdomen 7-segmented,
terminal segment broadly rounded; no trace of uropods. Sternal grooves
ending together on broad rounded-truncate sternal plate between bases of
chelipeds.

Eyestalk with scattered spinules. Antennular peduncle segments with few
scattered spinules; flagellum of six articles, barely extending to distal end of

DECAPOD AND ISOPOD CRUSTACEANS 23

antennal peduncle. Antennal peduncle segments with scattered spinules;
flagellum of about fifteen articles, extending well beyond orbit. Maxilliped 1
with triangular epipodite. Maxilliped 2 with narrow epipodite and gill.
Maxilliped 3 segments 3 to 6 with tiny spinules on outer surface, slender epipo-
dite and gill present. Chelipeds equal; palm of chela longer than finger and
thumb; dactylus with cutting edge of seven rounded cusps; terminal teeth
fitting between two terminal teeth of propodal finger; outer surface of propodus,
carpus, merus, and ischium bearing scattered spinules. Pereopods 2 and 3

Fig. 6. Pseudodromia cacuminis.
A. Holotype carapace in dorsal view. B. Ventral view of orbit and antennae. C. Sternum 9.
D. Cheliped. E. Pereopod 2. F. Pereopod 4. G. Pereopod 5.

24 ANNALS OF THE SOUTH AFRICAN MUSEUM

Bet hes |
7, Vi),

<
hoa SSE —
V Foe
: hos

Fig. 7. Pseudodromia cacuminis.
A. Maxilliped 1. B. Maxilliped 2. C. Maxilliped 3.

ambulatory, similar; dactyli with strong corneous unguis and five spines on
ventral margin; dactylus, propodus, carpus, and merus with scattered spinules.
Pereopod 4, dactylus forming pincer with strong curved terminal spine of
propodus; spinules on carpus, merus, and ischium. Pereopod 5 slightly longer
and more slender than pereopod 4; curved dactylus forming pincer with strong
straight spine of propodus; spinules on carpus, merus, and ischium. Branchial
formula: 8 gills (on maxillipeds 2 and 3, and pereopods 1-5) + 3 epipods (on
maxillipeds 1-3).

Material

Holotype SAM-A 16787 2 CL 6,0 mm CW 6,6 mm VEM 4.4

Paratype SAM-A 16788 immature ¢ CL 2,9 mm CW 3,0 mm VEM 4.2

Paratypes USNM 173518 2 CL 4,4 mm CW 4,8 mm VEM 4.6
immature 3 CL 2,9 mm CW 3,1 mm VEM 3.2

Remarks

The lack of epipodites on the pereopods, the tridentate rostral area of the
carapace, and a fifth pereopod longer than the fourth, place this material in
the genus Pseudodromia Stimpson.

DECAPOD AND ISOPOD CRUSTACEANS 25

Of the five species of Pseudodromia described, the Vema material most
closely resembles P. spinosissima Kensley, 1977, from deep water off the east
coast of South Africa. However, the overall carapace shape of the two species
differs, as does the carapace ornamentation (uniformly scattered spinules and
long hairs in P. spinosissima, patchy spinules and short spiky hairs in
P. cacuminis).

Etymology
The specific name “‘cacuminis’, meaning pointed as in a peak, refers to the
type locality, viz. the summit of Seamount Vema.

Family Homolidae
Paromola alcocki (Stebbing)
Thelxiope (Moloha) alcocki: Barnard, 1950: 341.
(See Gordon 1950 for full synonymy.)
Previous records
Algoa Bay, South Africa, 80 m; Mozambique, 312 m; Maldive Islands,
229 m.

Material
SAM-A16207 © CL (excluding rostrum) 42 mm, rostral length 8,5 mm,
supraorbital spine length 12,5 mm, west of Liideritz, about 800 m.

Remarks

This specimen closely resembles the type from South Africa, especially
in the relatively elongate pereopodal spination. The supra-orbital spines,
however, are relatively longer and more slender. The specimen differs markedly
from the type of P. alcocki faughni Serene & Lohavanijaya, 1973, from the
South China Sea, especially in its lack of strong setation, and in its stronger
carapace and pereopodal spination.

Paromola cuvieri (Risso)
Paromola cuvieri: Monod, 1956: 79, fig. 89.
Previous records
Eastern North Atlantic and southern Scandinavia; Mediterranean to
west Africa.

Material
SAM-A16789 ovigerous 2 CL 105 mm, CW 83 mm, Seamount Tripp,
150 m.
SAM-A16790 ¢g CL 120 mm, CW 102 mm, off Liideritz,
19°55’S 11°43’E.
Remarks

Although not previously recorded from southern Africa. P. cuvieri is
being commercially fished in the Liideritz area.

26 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Majidae
Macropodia cirripilus sp. nov.
Figs 8-9
Description
Male

Carapace piriform, dorsally convex. Scattered curved hairs over entire
carapace and abdomen. Rostrum of two relatively short parallel spines, reaching
distal end of third antennal peduncle segment. Supra-orbital eaves with four
or five short spines; strong nuchal spine present; hepatic region with few
scattered spine-tubercles; strong dorsolateral spine on protogastric region;
metagastric region convex with medial tubercle; convex bulbous branchial
region with several scattered tubercles; cardiac region convex, lacking tubercles.
Abdomen 6-segmented; third segment widest, with convex lateral areas; distal
margin of terminal segment evenly convex; all segments with weakly-raised
middorsal longitudinal ridge.

Eyestalk with curved anterior margin, produced into rounded papilla,
posterior margin straight; cornea oval, as wide as eyestalk base. Outer margin
of antennular fossa spinose; interantennular spine situated at end of acutely
triangular grooved process; basal peduncular segment of antennule inflated,
with row of four or five small spines. Basal antennal peduncular segment
narrow, with few small proximal spines and two more elongate spines distally;
second segment with single small distal spine; third segment longer than second,
unarmed. Epistome broader than long, flattened. Maxilliped 3 ischium wider
than merus, medial margin with several small tubercle-spines, exterior surface
with scattered tubercles; outer distal margin of merus with five spines, few
tubercles on external surface. Chelipeds subequal, only slightly longer than
middorsal carapace length (including rostrum); finger and thumb shorter
than palm; both cutting edges shallowly serrate, single spine at dactylar base;
upper surface of palm with few scattered tubercles; lower margin with row of
seven small spines; carpus shorter than palm, with few strong dorsal spines
and strong proximal tubercle on outer surface; merus one and one-half times
length of carpus, with row of spines on upper and lower margins; ischium
about one-third length of merus, with row of spines on ventral margin. Ambu-
latory pereopods decreasing in length posteriorly; pereopod 2 dactylus almost
straight, unarmed ventrally, almost as long as propodus. Dactylus of pereopod
3 with subapical secondary spine. Pereopods 4 and 5, dactyli curved, with
strong subapical secondary spine plus row of smaller spines. Pleopod 1 ¢
basally broad, tapering distally to curved rounded apex.

Female as in male, except for abdomen.

Material

Holotype SAM-A 16791 ¢ CL 5,0 mm CW 3,1 mm (across branchial areas)
VEM 15M
Allotype USNM 173519 2 CL 6,0 mm CW 3.9 mm VEM 4.4

DECAPOD AND ISOPOD CRUSTACEANS 27

Fig. 8. Macropodia cirripilus.
Holotype in dorsal view.

Remarks

Of the seven species of Macropodia mentioned by Monod (1956), the
present species resembles M. rostrata (Linnaeus), and that only to a limited
extent. This similarity lies in the shape of the male abdomen and pleopod 1,
and to a lesser degree, in the maxilliped 3. The carapace of M. rostrata, however,
is much more strongly spinose, while the ambulatory pereopods do not become
as strongly armed as in M. cirripilus. Further, the basal antennal segment is
unarmed. None of the five species from the Mediterranean (Forest & Zariquiey
Alvarez 1964) bears any close resemblance to M. cirripilus. The species referred
to by Barnard (1950: 15, fig 2j) as Macropodia formosa var., from off the Natal
coast, shows a similarity in the antennal and antennular spination and in the
rostrum, but the carapace is less spinose and the proportions are quite different.
The dactyli of the fourth and fifth pereopods, although possessing ventral
spines, lack the strong subterminal spine seen in M. cirripilus.

Etymology

The specific name is derived from the two Latin words ‘cirrus’, a curl,
and ‘pilus’ a hair, and refers to the characteristic curled hairs of the integument
of this species.

Family Xanthidae

Pilumnus sp.

Pilumnus hirsutus non Stimpson, Barnard, 1950: 263, fig. 49 (d—g).

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 9. Macropodia cirripilus.
A. Maxilliped 3. B. Abdomen 3. C. Ventral view of orbit and antennae. D. Cheliped
E. Pleopod 1 g. F. Pereopod 2. G. Pereopod 3. H. Pereopod 4. I. Pereopod 5.

DECAPOD AND ISOPOD CRUSTACEANS 29

Previous records
False Bay, Port Elizabeth, Port Shepstone, Durban.

Material
SAM-—A 16792 1 3 4 juv. 39 m Vem 2.3
SAM-A 16793 1 ¢ 50 m Vem 3.3
Remarks

The six small specimens from Vema agree well with the South African
material with which it has been compared. The true identity of this species,
however, is an open question. Barnard (1950) was careful to note that his
figures and description were based solely on South African material, which he
suspected differed from the Indo-Pacific P. hirsutus Stimpson. Comparison with
material from the Indo-Pacific shows some distinct differences. The Vema/
South African species possesses stiff hairs as well as longer flexible hairs. The
larger chela is proportionally squatter, with a shorter fixed finger in the Vema/
South African species. Unfortunately, the latter material is dry and in poor
condition, making further comparison difficult.

Comparison with the most closely related west African form, Pilumnus
inermis A. Milne Edwards & Bouvier, also reveals several differences, especially
in the carapace hairs.

ZOOGEOGRAPHIC DISCUSSION

Although the summit plateau of Seamount Vema is of somewhat limited
area (about 8 km in diameter), it supports a relatively rich fauna dominated
by encrusting and cryptic forms. Berrisford (1969, table 1) summarized the
affinities of the 105 species of invertebrates identified. Of these, 25 per cent
were South African species, 27 per cent had a scattered (cosmopolitan) distribu-
tion, 28 per cent were endemic, and 10 per cent had Indo-Pacific affinities.
Millar (1968) found the ascidian fauna of Vema to have strong affinities with
South Africa and no components in common with Tristan da Cunha.

Vema is about 11 x 10° years old (Simpson & Heydorn 1965), and older
than Tristan. Apart from the fish and the spiny lobster species, there are few
species in common, even though the prevailing oceanic conditions need not
reinforce isolation. Vema is bathed in South Atlantic Central Water, with what
was initially interpreted as local subsurface upwelling of Antarctic Intermediate
water having a strong northerly-flowing component (Simpson & Heydorn
1965: 251). Welsh & Visser (1970), however, suggest that this apparent upwelling,
which also occurs further south away from any topographical features such as
seamounts, is really cyclonic upwelling caused by a ‘dying’ eddy moving north-
ward. These eddies are formed in the area where the Agulhas Current meets
the West Wind Drift.

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

The fish, being mainly pelagic forms, can be regarded as part of the Vema
fauna only in the widest sense, while Penrith (1967) has recorded only one
endemic species. The problem of recruitment of Jasus tristani is less simple,
but with a planktonic life of several months, phyllosomata originating at
Tristan would need to be transported in a north-easterly direction, perhaps
by offshoots of the West Wind Drift encountering the north-flowing Benguela
System, for successful colonization. This obviously happens, judging from the
population discovered in the later 1950s. Since then, Vema’s spiny lobster
population has been heavily exploited. By 1967 Heydorn reported the summit
almost denuded of Jasus, while the divers of the 1978 cruise did not see any
lobsters, neither were any specimens of the grapsid crab Plagusia chabrus
noted. This latter species was fairly commonly seen on the earlier visits. How
long a time is required for this population to recover will probably be answered
only by a theoretical exercise in population dynamics.

The isopods, with their strong South African affinity and lacking plank-
tonic larvae for dispersal, perhaps reached Vema clinging to drifting kelp. The
major alga of the summit is Ecklonia biruncinata, which also occurs off the
southern Cape coast, and would provide ideal shelter for clinging animals.

Seventeen species of decapods are included in this brief discussion of
zoogeography (the two mesopelagic species mentioned being excluded). Of
these seventeen, six have been recorded from South Africa, including three
from the east coast only (Euvalus ctenifera, Paromola alcocki, Lithodes murrayi);
Pontophilus sculptus, known from False Bay to Natal and also from the
Mediterranean, north-western Atlantic, and Angola; Pilumnus sp. recorded
as P. hirsutus from the east and south coast; and Plagusia chabrus, an essentially
cold-temperate austral species known from South West Africa to Natal,
Australia, New Zealand, Chile, and Juan Fernandez. Lithodes murrayi was
previously regarded as an austral form, but has been recorded from deep water
off Natal (Kensley 1977).

Pseudactaea corallina is a true Indo-Pacific species and has not been
recorded from the east coast of South Africa.

The two new species described here, viz. Pseudodromia cacuminis and
Macropodia cirripilus, are the only ‘endemics’.

Five species have been recorded from West Africa; Alpheus macrocheles,
Synalpheus huluensis africanus, and Pagurus cuanensis (also known from the
Mediterranean) are regarded as true West African forms; Paromola cuvieri
and Eumunida picta have a much wider range. Pagurus chevreuxi 1s known
only from the Mediterranean.

With an age of eleven million years, it is not difficult to envisage coloni-
zation of Seamount Vema by West African/Mediterranean species. The species
from the Indo-Pacific and the east coast of South Africa, however, must have
been faced with greater problems of colonization. Perhaps the most feasible
explanation is that planktonic larval forms of these species, present in
southward-flowing Agulhas water, were caught in the pockets of Agulhas

DECAPOD AND ISOPOD CRUSTACEANS 31

water eddying northward in the South Atlantic as previously mentioned. The
temperature regime in this series of events would not be a barrier to colonization.
Although there is so-called upwelling of Antarctic Intermediate water in the
vicinity of Vema, above the 75 m depth line the temperatures are fairly uniform
(Welsh & Visser 1970: 2), being between 18° and 21° C, and comparable with
east coast shallow-water temperatures.

ACKNOWLEDGEMENTS

My thanks are due to Prof. J. Field of the Department of Zoology,
University of Cape Town, for making the present collection available for study;
Messrs C. Beyers and G. Fridjhon of the Sea Fisheries Branch, Cape Town,
for data and donation of material to the South African Museum; the South
African Museum for making the material available for study; Prof. J. R.
Grindley of the School of Environmental Studies, University of Cape Town,
for information on Vema; and Blue Continent Products of Cape Town for
allowing me to examine specimens of Paromola cuvieri. I am grateful to
Dr D. Guinot of the Paris Museum, for examining the xanthid material and
for her valuable comments; and to Drs T. E. Bowman and R. B. Manning of
the Department of Invertebrate Zoology, Smithsonian Institution, for reading
the manuscript and for their useful criticisms.

REFERENCES

ARNAUD, P. M. 1971. Lithodes murrayi Henderson, 1888 (Crustacea, Decapoda, Anomura)
dans les eaux cétiéres des iles Crozet (SW de |l’Océan Indien). Tethys 3: 167-172.
BARNARD, K. H. 1920. Contributions to the Crustacean Fauna of South Africa. No. 6. Further
additions to the list of marine Isopoda. Ann. S. Afr. Mus. 17: 319-438.

BARNARD, K. H. 1950. Descriptive catalogue of South African Decapod Crustacea (Crabs
and Shrimps). Ann. S. Afr. Mus. 38: 1-837.

BARNARD, K. H. 1965. Isopoda and Amphipoda collected by the Gough Island Scientific
Survey. Ann. S. Afr. Mus. 48: 195-210.
BERRISFORD, C. D. 1969. Biology and zoogeography of Vema Seamount: a report on the
first biological collection made on the summit. Trans. R. Soc. S. Afr. 38: 387-398.
Forest, J. & ZARIQUIEY, R. A. 1964. Le genre Macropodia Leach en Méditerranée. 1. Descrip-
tion et étude comparative des espéces (Crustacea Brachyura Majidae). Bull. Mus. natn.
Hist. nat., Paris (2) 36: 222-244.

Gorpon, I. 1950. Crustacea: Dromiacea. Part I. Systematic account of the Dromiacea
collected by the ‘John Murray’ Expedition Part II. The morphology of the spermatheca
in certain Dromiacea. Scient. Rep. John Murray Exped. 9: 201-253.

GRINDLEY, J. 1967. Research on the Vema Seamount. Comm. Fish. News, S. Afr. 2: 14-19.

HAte, H. M. 1941. Decapod Crustacea. Rep. B.A.N.Z. antarct. Res. Exped. (B) 4: 259-285.

HENDERSON, J. R. 1888. Report on the anomura collected by H.M.S. Challenger during the
years 1873-1876. Rep. Voy. Challenger 27: 1-221.

Heyporn, A. E. F. 1967. Research on the Vema Seamount. S. Afr. Ship. News Fish. Ind.
Rey. 12: 79-83.

KENSLEY, B. 1975. Five species of Jaeropsis from the southern Indian Ocean. (Crustacea,
Isopoda, Asellota). Ann. S. Afr. Mus. 67: 367-380.

KENSLEY, B. 1977. The South African Museum’s Meiring Naude cruises. Part 2. Crustacea,
Decapoda, Anomura and Brachyura. Ann. S. Afr. Mus. 72: 161-188.

MILLAR, R. H. 1968. A collection of Ascidians from the Vema Seamount. Trans. R. Soc.
S. Afr. 38: 1-22.

32 ANNALS OF THE SOUTH AFRICAN MUSEUM

Monop, T. 1956. Hippidea et Brachyura Ouest-africains. Mem. Inst. fr. Afr. Noire 45: 1-674.

PENRITH, M. J. 1967. The fishes of Tristan da Cunha, Gough Island, and the Vema Seamount.
Ann. S. Afr. Mus. 48: 523-548.

SERENE, R. & LOHAVANIJAYA, P. 1973. The Brachyura (Crustacea: Decapoda) collected by
the Naga Expedition including a review of the Homolidae. Naga Rep. 4 (4): 1-187.

SIMPSON, E. & HEYDORN, A. E. F. 1965. Vema Seamount. Nature, Lond. 207: 249-251.

VANHOFFEN, E. 1914. Die Isopoden der Deutschen Siidpolar-Expedition 1901-1903. Dt.
Siidpol.-Exped. 15: 447-598.

WELSH, J. G. & ViSsER, G. A. 1970. Hydrological observations in the south-east Atlantic
Ocean. 2. The Cape Basin. Invest] Rep. Div. Fish. Rep. S. Afr. 83: 1-5.

WoLrFr, T. 1962. The systematics and biology of the bathyal and abyssal Isopoda Asellota.
Galathea Rep. 6: 1-320.

YALDWyYN, J. C. & Dawson, E. W. 1970. The stone crab Lithodes murrayi Henderson: the
first New Zealand record. Rec. Dominion Mus. 6: 275-284.

6. SYSTEMATIC papers must conform to the J/nternational code of zoological nomenclature
(particularly Articles 22 and 51).

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
- not acceptable.

In describing new species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes shou!d 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°51’S 25°39’E), collected by A. Smith, 15 January 1973.

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

7. SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. *. . . the Figure depicting C. namacolus ...’; *. . . in C. namacolus (Fig. 10)...

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. DuToit but A.L.du Toit; Von Huene but F. von Huene

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.

Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.

BRIAN KENSLEY

DECAPOD AND ISOPOD CRUSTACEANS FROM
THE WEST COAST OF SOUTHERN AFRICA,
INCLUDING SEAMOUNTS VEMA AND TRIPP

Di? .
VOLUME 83 PART 3 MARCH 1981 ISSN 0303-2515

{ ;
|

OF THE SOUTH AFRICAN |
” MUSEUM J

CAPE ‘TOWN.

INSTRUCTIONS TO AUTHORS

1. MATERIAL should be original and not published elsewhere, in whole or in part.
2. LAYOUT should be as follows:

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(c) Table of contents giving hierarchy of headings and subheadings
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(i) Abbreviations, where these are numerous

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The number of the figure should be lightly marked in pencil on the back of each illustration.

5. REFERENCES cited in text and synonymies should all be included in the list at the end of
the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable):

(a) Author’s name and year of publication given in text, e.g.:

‘Smith (1969) describes...’

‘Smith (1969: 36, fig. 16) describes .. .’

“As described (Smith 1969a, 19695; Jones 1971)’
‘As described (Haughton & Broom 1927)...’
‘As described (Haughton et al. 1927)...’

Note: no comma separating name and year
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For books give title in italics, edition, volume number, place of publication, publisher.

Examples (note capitalization and punctuation)

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

FISCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.

FiscHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634. \

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume $3 Band
March 1981 Maart
Part 3 Deel

FIRST RECORD OF THE MELON-HEADED WHALE
PEPONOCEPHALA ELECTRA
FROM SOUTH AFRICA

PELER Be BEST
&
PETER D. SHAUGHNESSY

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material
becomes available

Obtainable from the South African Museum, P.O. Box 61, Cape Town

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word uitgegee in dele op ongereelde tye na gelang van die
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Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad

OUT OF PRINT/ UIT DRUK

1, 2(1-3, 5-8), 3(1-2, 4-5, 8, t.-p.i.), 51-3, 5, 7-9),
6(1, t.—p.i.), 7-4), 8, 9(1-2, 7), 101-3),
11(1-2, 5, 7, t.—p.i.), 15(4-5), 24(2), 27, 311-3), 32(5), 33, 45(1)

Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 004 4

Printed in South Africa by In Suid-Afrika gedruk deur
The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk.,
Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

FIRST RECORD OF THE MELON-HEADED WHALE
PEPONOCEPHALA ELECTRA FROM SOUTH AFRICA

By
PETER B. BEST
&
PETER D. SHAUGHNESSY

Sea Fisheries Institute, Cape Town

(With 7 figures and 4 tables)
[MS. accepted 17 July 1980]

ABSTRACT

An adult male Peponocephala electra that stranded at Hout Bay (34°03’S 18°21’E) in
July 1976 was the first record of the species from South Africa and the South Atlantic. Details
of the animal’s external appearance, body measurements, organ weights, parasites, stomach
contents, skull measurements and skeletal characters are given. Differences in coloration and
head and flipper shape between this species and Feresa attenuata are demonstrated.

CONTENTS
PAGE
Introduction ‘ : : : Senos
External appearance . f : Be
Life history data : : : hes)
Skeleton . : ; ; : ie SAS
Discussion ; : j : i “45
Acknowledgements. : . 46
References ; : : ‘ | *46
INTRODUCTION

At about 07h40 on 16 July 1976 a single small whale stranded alive on
the beach at Hout Bay (34°03’S 18°21’E). It was placed in the water four
times by the harbour master, but each time the animal returned to the beach.
Finally it was taken to the harbour where it was placed in the water from a
jetty, but the animal then stranded on a near-by slipway, where it died half
an hour later (at about 10h00).

Later the same day the whale was examined by staff of the marine mammal
laboratory of the Sea Fisheries Institute, and was found to be a melon-headed
whale! (Peponocephala electra), the first such record for South Africa. A cast
of the head was prepared and the whole skeleton was presented to the South
African Museum (ZM 38245).

1 As there appears to be no Afrikaans (or Dutch— Van Bree 1975) name for this species,
the authors suggest ‘bolkopdolfyn’.

813)

Ann. S. Afr. Mus. 83 (3), 1981: 33-47, 7 figs, 4 tables.

34 ANNALS OF THE SOUTH AFRICAN MUSEUM
EXTERNAL APPEARANCE

The animal was a male, 248 cm long, and was generally undamaged apart
from a few superficial cuts and abrasions caused by stranding. It was photo-
graphed about an hour after death, while the colour pattern was still rather
prominent (Figs. 1-6).

The general body coloration was bluish-black overall (Fig. 1). Both upper
and lower jaws, however, were irregularly edged with white as far back as the
angle of the gape (Fig. 2). On the belly of the animal there was also a greyish-
white ‘blaze’ in the midline extending from the throat to the anal slit (Fig. 3).
The shape of this blaze was similar to the ‘throat chevron-genital patch pattern’
(Mitchell 1970) seen in pilot whales (especially Globicephala melaena), false
killer whales (Pseudorca crassidens), pygmy killer whales (Feresa attenuata),
and Risso’s dolphins (Grampus griseus), particularly juveniles. On the throat
this blaze was roughly bracket-shaped, rapidly narrowing posteriorly to a thin
mid-ventral streak between the flippers. The blaze gradually widened on the
abdomen to form a lozenge-shaped mark extending approximately from the
umbilicus to the anus. Only at its posterior extremity (where the blaze formed
a V terminating at the anterior end of the anal slit) were its outer margins
well defined.

The animal had a healed scar roughly elliptical in outline under the left
flipper (probably attributable to the small shark Jsistius (Jones 1971)). The
only other feature of the body coloration was a well-defined pale band on the
dorsal midline of the head from the blowhole to the tip of the snout, which
broadened anteriorly to cover most of the front of the head (Fig. 4). This band
appears equivalent to the ‘apex of melon to blowhole stripes’ described by
Mitchell (1970) for Tursiops and other species of delphinid cetaceans. It is
just apparent in figures of P. electra provided by Nishiwaki & Norris (1966),
being masked by highlights in most pictures, but has not been described pre-
viously for this species, nor for F. attenuata.

In other respects the animal was very similar in coloration to the pygmy
killer whale. However it lacked the pale grey lateral coloration described for
F. attenuata by Nishiwaki et al. (1965). As this feature fades rapidly after
death, its absence on this P. e/ectra carcass cannot be taken unequivocally as
applying to the animal in life. Bryden et al. (1977b), however, assert that this
coloration is not present in Peponocephala, and it could not be detected on a
Captive animal (seen by P.B.B.) at Sea Life Park, Hawaii, on 23 March 1980.

The dorsal fin of the Hout Bay specimen had presumably been damaged,
as the trailing edge was ragged (Fig. 5).

The external measurements of the whale were taken as recommended by
the Committee on Marine Mammals, American Society of Mammalogists
(Norris 1961). As only ten specimens of this species appear to have been
measured previously the data from all eleven animals (expressed as proportions
of the total body length) are presented in Table 1. The specimen measured by

FIRST MELON-HEADED WHALE FROM SOUTH AFRICA 35

Fuh PAE.
we a oie.
pic:

pty

BS oi
baa

Fig. 1. General body coloration of the Peponocephala electra that stranded at Hout Bay.

Fig. 2. Head of the Peponocephala electra that stranded at Hout Bay showing white edging
to the upper and lower jaws.

36 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 3. Belly of the Peponocephala electra that stranded at Hout Bay showing a greyish-white
‘blaze’ along the midline.

Fig. 4. Head of the Peponocephala electra that stranded at Hout Bay showing a pale band
from the blowhole to the snout.

FIRST MELON-HEADED WHALE FROM SOUTH AFRICA 37

Fig. 5. Dorsal fin of the Peponocephala electra that stranded at Hout Bay.

Pilleri & Gihr (1973-4) was a mounted skin and hence the measurements may
not be strictly comparable.

At least some of the measurements given for specimen C15 appear to be
erroneous, particularly those concerning the position of the genital aperture
and anus, which (although the animal was a male) are placed further posteriorly
than in any of the females measured. The data for this animal have therefore
been excluded from subsequent analyses. Although the remaining sample
is small (six males, three females and one of unknown sex), a comparison
between sexes of the proportional measurements indicates some apparent
differences (apart from those concerned with the position of the anus and
genital aperture). In males the anal girth appears larger (29,1-38,6% cf
26,1—28,8%), the flippers longer (17,5-20,4% cf 16,2-17,0%, or 13,0-15,8%
cf 11,7-12,5°%), the dorsal fin greater in height (8,4-10,9% cf 7,3-8,0°%) and
the tail flukes broader from the notch to their anterior margin (6,1-7,2% cf
5,1-5,5%) than in females. These apparent differences cannot all be accepted
as evidence of sexual dimorphism until ontogenetic changes in body proportions
of P. electra have been investigated: the present sample is too small for such
an analysis, but it may be significant that only one of the three females was
sexually mature, while four of the six males could be classified as mature
(Bryden et al. 1977b). Differences in the anal girth may reflect the more anterior
position of the opening in males. Alternatively, they may be indicative of real

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FIRST MELON-HEADED WHALE FROM SOUTH AFRICA 39

sexual dimorphism. The largest male P. electra yet measured (273 cm long
(Goodwin 1945)) had a pronounced protuberant keel posterior to the anus,
similar to that seen in adult males of some stocks of delphinid species (e.g.
eastern Pacific Stenella longirostris (Perrin 1972)). There is also a suggestion
of a similar but smaller keel in the male 267,7 cm long illustrated by Bryden
et al. (1977a). The apparently greater anal girth in male P. electra could reflect
the development of this protuberant keel, if it should be found to be a con-
sistent feature of the males of this species.

By comparison with the only species with which it is likely to be confused,
F. attenuata, the melon-headed whale has a relatively longer head section. Measure-
ments from the tip of the snout to the angle of gape, eye and blowhole as a pro-
portion of the total body length are all greater in P. electra than in F. attenuata
(Table 2). This distinction does not extend as far as the anterior insertion of the
flipper, suggesting that the real difference between the two species lies in the
length of the rostrum. No other distinctions in body proportions could be
found between the two species.

TABLE 2

Comparison of body measurements (expressed as a proportion of body length) between
Peponocephala electra and Feresa attenuata.

P. electra’ F. attenuata*
Measurement mn mean range n mean range
Mipior snout to centre ofeye . . .- 10 13,7 12,7-14,9 19 10,1 8,1-12,0
Tip of snout to angle of gape . . : 9 10,4 9,7-11,3 17 69 5,4— 8,9°
iiponsnoeut to blowhole... . =~ ©. 10 13,9. 12,9-14,9 19s 9:8e5 67-119
Tip of snout to anterior insertion of
mippewy Cs Oe 20 TnI 5=23.4 18 19,8 16,8-22,6

1 From Table 1 (C15 excluded).

2 From Best (1970), Bryden (1976), Nishiwaki et al. (1965), Perrin & Hubbs (1969), and
Pryor et al. (1965).

3 Excluding a value of 11 per cent (Bryden 1976) which appears atypical as it places the angle
of gape posterior to the blowhole, a situation not recorded in any of the other specimens
measured to date.

From a photographic comparison, however, the flippers of P. electra
(Fig. 6) appear to be more pointed at the tip and with a straighter trailing
edge than those of F. attenuata (Best 1970; Bryden et al. 1977a; Nakajima &
Nishiwaki 1965; Nishiwaki et al. 1965; Perrin 1976; Pilleri & Gihr 1973-4;
Pryor et al. 1965; Rancurel 1974; Yamada 1954). This distinction has been
noted by Bryden et al. (1977b), and may arise from differences in the arrange-
ment of bones in the flipper (see below). To illustrate the distinction, outline
drawings of flippers of P. electra and F. attenuata are shown in Figure 7.

LIFE HISTORY DATA

The animal weighed 206 kg entire on a platform scale, but its weight in
parts totalled 209,8 kg (Table 3). As dissection of a large cetacean normally
creates significant weight-loss (Lockyer 1976), one of the two weighings must

40 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 6. Flipper of the Peponocephala electra that stranded at Hout Bay.

TABLE 3
Weight of Peponocephala electra in parts and weights of various organs.
Weight
Part Ib kg

Blubber_ . ‘ 5 : ; ; : : 2 : ; 97 (44,0)

Axial muscle, dorsal ee Se ae ee es 91 (41,3)

WENCH wa cilee 28 el | abi Sees ne we 52 (23,6)

Head? . oe ae alaghe 65 (29,5)

Vertebral column Gncluding tail) . : ‘ : : : , 46 (20,9)

Ribcage . : : ; : : ; : A : : 46 (20,9)

Flippers & scapulae : . F : : ’ ' : é 18 (8,1)

WVisceta:.-* +. , : - : ; : : F ; : : ATs (21-5)

Total . ‘ ge we Me ge : 2 ; ae 3) G 4625 (209,8)

Organ

Heart (minus clots) . : : : : : eA ie : ; 1,025

Lungs & trachea. ; ' , : ; : é ; e 5,245

Liver . 2 3 : z : . ; ; 2 : : 3,310

Kidneys, left : : é : ; Pen raeh By ; ; ; 0,575

right . : : , ‘ 5 : : ; : : 0,560

Spleen ; . : . ‘ : e , . : : ‘ 0,075

Intestines . ; ; : E ; : p : : F : 3,750
Adrenals, left . . : 3 ; : Pius ‘ : : 0,0068
right . : : : : 2 : ae : 0,0092

Diaphragm : : / : ; : : ‘ . , : 1,455

Oesophagus . : : : : : ; : : ; 0,360

Bladder’ 2%: i. Oe egies ese At. Bea 0,130

Stomach (plus contents) . : : : : : : : 2,455

1 Parts were weighed on a spring balance erate in pounds.
2 Including associated blubber and muscle.

FIRST MELON-HEADED WHALE FROM SOUTH AFRICA 4]

Fig. 7. Outlines of left flipper.

A. Feresa attenuata (from Nishiwaki et al. 1965).
B. Peponocephala electra (SAM-ZM38245).

be incorrect. The platform scale belonged to a commercial fishing company in
Hout Bay and was regularly assayed, and hence its accuracy should perhaps
be considered more reliable. Repeat weighings on this scale, however, produced
differences of up to 2,5 kg depending on the position of the animal on the
platform. The highest weight obtained (206 kg) has been adopted as the most
accurate, given the apparent weight of the animal in parts.

This is the largest P. electra weighed to date. Bryden et al. (1977b) provided
data for a foetus (3,9 kg) and three immature animals (78, 100 and 100 kg),
while a calf from the eastern tropical Pacific Ocean weighed 15 kg (Perrin 1976).

No external parasites were found, despite an examination of the body
surface, eye, blowhole, mouth, anal and genital slits, and appendages. ‘Whale
lice’ (Miyazaki & Wada 1978) and traces of barnacle attachments on the tail
flukes (Bryden et al. 1977a) have been recorded previously.

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

Numerous internal parasites were found. Longitudinal strips of blubber
were cut transversely at 10 to 13 cm intervals, and the number of cestode
cysts (probably Phyllobothrium sp.) counted. These totalled 58. ofwhich about
90 per cent were in the posterior half of the body. Cestode cysts (Phyllobothrium
chamissonii) were also found in the abdomen grouped around the rectum (32),
in the mesentery of the small intestine (2), in the diaphragm (2), and in the
muscle (1). Nematodes (Anisakis simplex) were found in the oesophagus (36)
and in the stomach (chiefly the second), where they weighed 20 g. The large
and small intestines were opened for about 25 cm at 2 m intervals: one incom-
plete acanthocephalan (probably Bolbosoma sp.) was found in the rectum and
2 nematode fragments (probably Anisakis simplex) at about the midlength of
the small intestine. Parasites were not found in the heart, liver, lungs, kidneys,
or bladder.

Parasitic cysts in the blubber (unidentified) and Phyllobothrium chamissonii
(= Monorygma sp.) cysts in the stomach wall or between the peritoneum and
abdominal muscles in the inguinal region of P. electra have been recorded
previously (Bryden et al. 1977a; Cannon 1977; Dailey & Brownell 1972).
Nakajima & Nishiwaki (1965) reported the presence of unidentified stomach
nematodes, while Cannon (1977) described Anisakis simplex and A. typica
from the stomach of P. electra. Bryden et al. (1977a), Dawbin et al. (1970),
and Nakajima & Nishiwaki (1965) all record the presence of small thread-like
worms or nematodes in the air sinuses of the head, identified by Bryden et al.
(1977a) as Stenurus sp. and by Cannon (1977) as S. globiocephalae. The air
sinuses of the Hout Bay specimen were unfortunately not examined. Dailey
& Brownell (1972) also list the trematode Nasitrema sp. and the nematode
Halocercus sp. as ‘new host records’ for this species, without specifying the
host tissue.

The stomach contained 2 upper beaks and | lower beak of Loligo reynaudi
and 1 lower beak of a juvenile ommastrephid squid, a squid pen, 2 fish otoliths
(Merluccius sp.) and some sand. There are no previously identified stomach
contents for this species. As indicated by Leatherwood & Walker (1979),
however, stomach contents of stranded cetaceans should be interpreted with
a great deal of caution: in Lissodelphis borealis these authors found many
near-shore fish species not representative of the normal known distribution of
the dolphin and which were probably ingested just prior to stranding.

The faeces were bright green with much mucus, indicating that the animal
was either sick or had not fed for some time (Ridgway 1972).

The testes (without epididymides) weighed 760 g (left) and 1 035 g (right),
and measured 34 x 7,4 x 3,9 cm (left) and 38 x 8,8 x 4,6 cm (right). Bryden
et al. (1977b) have summarized the known reproductive data for male P. electra,
comprising a total of five animals. A male with a combined testis weight of
1 359 g was shown histologically to be sexually mature, so it is reasonable to
assume that the Hout Bay animal was also mature.

FIRST MELON-HEADED WHALE FROM SOUTH AFRICA 43

SKELETON

The skull dimensions of the Hout Bay specimen were measured using a
pair of 55 cm vernier calipers. To avoid ontogenetic differences, comparison
with previously measured skulls (Table 4) has been confined to animals with a
condylobasal length (CBL) exceeding 440 mm: this limit was chosen on the
basis of the determination of skulls with a CBL of 415 mm as adolescent and
a CBL of 456 mm as adult (Dawbin et al. 1970), and the description of a skull
with a CBL of 440,2 mm as ‘approaching adulthood’ (Bryden et al. 1977a).
The Hout Bay specimen could clearly be classified as adult from the robust
rostrum, strongly developed supraoccipital crest, the degree of fusion of sutures,
the advanced ossification of the mesethmoid, the degree of filling-in of the
cranial hiatus and the posterior extension of the vomer in the basicranial
trough (Dawbin et al. 1970).

TABLE 4
Skull measurements of Peponocephala electra
ZM 38245 Previous records?
mm VEC BE n Range of % CBL
Condylobasal length . 477 100 — —
Rostrum length . 258 54,1 2. 52,3—-55,6
Rostrum basal width : 136 28,5 12 27,6—31,0
Rostrum width 60 mm ant. to antorbital
notches : 116 24,3 11 22,2-26,5
Rostrum width at middle 97 20,3 11 17,3—25,2
Premaxillae, width at same point 46 9.6 7 8,7-14,25
Tip of snout to blowhole . 320 67,1 7 68,7-70,4
Tip of snout to pterygoid 300 62,9 5 62,5-66,5
Preorbital width ‘ 245 51,4 11 50,5-54,4
Postorbital width 263 55,1 12 53,6-57,6
Orbital width 249 52.2 7 51,3-54,6
Blowhole, width at 61 12,8 Z 11,9-16,6
Zygomatic breadth 263 55:1 11 54,0-57,6
Greatest width of premanxillaries 101 Fi? 12 19,9-23,6
Width of braincase across parietals 197 41,3 10 37,7-44,0
Length of upper toothrow L 186 39,0 11 36,6-40,7
R : 184 38,6 11 35,3-40,9
Hinder edge of upper toothrow L 184 38,6 5 38,6-41,0
to tip of premaxillae R 185 38,8 5 37,5—41,2
Length of lower toothrow L 164 34,4 8 34,6-38,6
Re. 163 34,2 8 33,8-39,2
Hinder edge of lower tooth L 170 35,6 5 35,6-38,2
row to tip of mandible R 170 35,6 5 34,4-38,5
Mandible length 384 80,5 9 79,4-81,8
Coronoid height 95 19,9 9 17,7—20,5
Length of symphysis . 38 8,0 7 6,7— 9,2
Post-temporal length 97? 20,3 10 17,1-21,0
Post-temporal height 62? 13,0 10 11,0-15,3
Width at 2 rostrum length 2 153 5 14,7-16,9
Cranial height : 154 323 6 28,5—40,4
Cranial length, internal 154 3233 5 27,8-33,9°
Tooth count? RU 22 14 21-26
LU 21 14 20-25
RL 2D Lg, 22-25
BE 22 13 22-25

1 From Bryden et al. (19772), Dawbin ef al. (1970), and Van Bree & Cadenat (1968).

2 Left side only.
* Count of alveoli.

3 Plus one outlying value of 18,7 per cent (Bryden et al. 1977a).

44 ANNALS OF THE SOUTH AFRICAN MUSEUM

Nearly all the skull dimensions (expressed as percentages of CBL) of
the Hout Bay animal fall within the range previously recorded for ‘adult’
P. electra. One exception is the measurement tip of snout to blowhole, which
appears shorter than any previously recorded. Comparison of the outline
of the nares with figures provided by Dawbin et al. (1970) and by Van Bree
& Cadenat (1968), however, suggest that either the Hout Bay specimen was
atypical, or that the anterior margin of the nares was damaged. The dimensions
of the tooth rows in general are close to or below the lower limits of the pre-
viously reported ranges, but the number of teeth (= alveoli) present is also at
or just above the lower limit recorded for other adults.

Many of the teeth showed extensive wear at the tip, particularly in the
lower jaw, so that the occlusal surfaces were flattened. The maximum diameter
of the five largest teeth in the upper and lower jaws (measured with dial calipers)
averaged 6,3 and 6,6 mm respectively, while three of the relatively unworn
teeth (all from the upper jaw) had overall dimensions (length x maximum
diameter) of 8,8 x 5,8, 7,7 x 5,9 and.7,6 x 5,8 mm. The tecthien mare cer.a
are, therefore, somewhat smaller than those of F. attenuata, where the five
largest teeth in upper and lower jaws of two specimens averaged 21,2 x 6,8
mm (upper), 23,2 x 7,4mm (lower), and 19,2 x 6,4mm (upper), 22 0 x 8,0
mm (lower) (Nishiwaki et al. 1965).

The vertebral column of the Hout Bay specimen was composed of eighty-
one vertebrae: previous vertebral counts for P. electra have been eighty-one
(Bryden et al. 1977a, Goodwin 1945) and eighty-two (Nakajima & Nishiwaki
1965). Comparison of counts of vertebrae in-different regions of the column
with published data is impossible when the criteria used for distinguishing the
regions are not specified (as indicated by De Smet 1977). Adopting De Smet’s
nomenclature, the vertebral formula of the Hout Bay specimen was as follows:
Cy = 4)-+ 3, Th.v. = 12, 1.7h.f =2, X = 18, Y = 307 oe

All epiphyses were fused to their centra, confirming the status of the
animal as adult.

Vertebrae numbers 46 and 47 (the 7th and 8th of the caudal series) were
partly fused together by their left transverse processes, clearly a pathological
condition.

There were 14 thoracic ribs on the right side and 13 on the left, 6 of which
on each side possessed a definite capitulum and tubercle. Each side of the 7th
thoracic vertebra, however, possessed a spur 14 to 17 mm long on the ventral
surface of the transverse process, while at similar positions on each side of the
8th thoracic vertebra there was a small protuberance. Similar structures were |
figured or described by Dawbin et al. (1970), also on the 7th and 8th thoracic
vertebrae, and described as rib neck vestiges: in an animal from Australia these
structures were present on the 8th, 9th and 10th thoracic vertebrae (Bryden
et al. 1977a.) A ‘spur’ on the 7th thoracic was also described by Nakajima &
Nishiwaki (1965). Vestigial catapophyses were present on the 6th to 11th
thoracic vertebrae: Bryden et al. (1977a) found them on the 7th to 10th vertebrae.

FIRST MELON-HEADED WHALE FROM SOUTH AFRICA 45

The 5th rib on the left side of the thorax bore evidence of a healed fracture
at a point about 40 per cent of its length from the capitulum.

Unlike F. attenuata, where the flippers normally contain four carpal
bones (Best 1970), the Hiratsuka specimen of P. electra had five carpals in
both flippers (Nakajima & Nishiwaki 1965). As determined by radiography,
the Hout Bay specimen had five carpals in the right flipper and six in the left,
the sixth being a small, almost circular element in contact with the cuneiform,
hamate and fourth metacarpal bones.

The phalangeal formula was

mon. 11: 8, 1: 6, 1V 23, V 32
Rishtl’=3, 11: 8, Il: 6, 1V :4, V :2

The minute terminal phalanges on the 2nd to 5th digits shown in Nakajima
& Nishiwaki’s (1965) X-rays were (apart from RIV) absent. There was no
indication of the bilateral asymmetry reported by Bryden et al. (1977a).

Despite Nakajima & Nishiwaki’s (1965) contention that the phalangeal
formula of P. electra resembled that of F. attenuata, there is some indication
that the relative numbers of phalanges in digits II and III differ. In 21 flippers
of F. attenuata examined by Nishiwaki et al. (1965), 76 per cent had a difference
of only one phalange between digits II and III, and 24 per cent a difference of
two phalanges. In both the Hiratsuka and Hout Bay specimens of P. electra,
there was a difference of two phalanges between these digits, while Bryden
et al. (1977a) gave the range for P. electra as 8-9 phalanges for digit II and 6-7
for digit III. This apparent difference between P. electra and F. attenuata may
account for the difference in flipper shape mentioned above.

The sternum consisted of four elements, of which the most posterior was
very small, as in the specimen examined by Nakajima & Nishiwaki (1965),
but unlike their example all four elements were unfused. There were ten pairs
of sternal ribs, as opposed to the nine found by Nakajima & Nishiwaki (1965).

DISCUSSION

Since Perrin (1976) summarized the known records of P. electra, and
illustrated their distribution, there have been several additional published
records. Caldwell et al. (1976) documented four specific records from the island
of St Vincent in the southern Caribbean. Bryden et al. (1977a) described two
specimens from Queensland and one from the Queensland—New South Wales
border of Australia. A mass stranding of fifty-three individuals on Moreton
Island, Queensland, was later described by Bryden et al. (1977b), and these
authors also listed two previously unpublished records for the Australian region
—an immature male stranded at Tweed Heads, New South Wales, in February
1967, and a sighting of a group of 100 off Stadbroke Island, Queensland, in
May 1975. Miyazaki & Wada (1978) mentioned an animal of this species
collected at sea in the western tropical Pacific. Perrin (1976) also omitted from
his figure the record from Derby, Western Australia listed by Dawbin et al.
(1970).

46 ANNALS OF THE SOUTH AFRICAN MUSEUM

The nearest published records of the species to South Africa are an animal
harpooned in mid-Atlantic at 03°03’N 24°40°W (Goodwin 1945) and a skeleton
from the central Indian Ocean from Gan Island, Addu Atoll, Maldive Islands,
at about 00°30’S 73°20’E (Dawbin et al. 1970). In addition, however, there
are six skulls (one with an incomplete skeleton) of this species in the British
Museum (Natural History) that were collected from stranded animals on the
south side of Aldabra Atoll (09°20’S 46°25’E) near a place called Dune Jean
Louis in September 1974. The catalogue numbers are 1980.147 to 1980.152
(M. C. Sheldrick in litt. 14 January 1980). The Hout Bay animal, therefore,
represents not only the first record for South Africa but also the first record
for the South Atlantic, and a major apparent range extension.

P. electra is usually considered to occur in tropical (Rice 1977) or tropical
and subtropical waters (Van Bree & Cadenat 1968). Under this assumption
the South African record, at 34°03’S and at the southern end of the cold
Benguela Current system, may represent an animal at the probable extreme
end of its range.

ACKNOWLEDGEMENTS

We should like to thank J. J. Moolman for reporting the stranding and
providing us with assistance at the harbour, R. W. Weeks and P.-J. Mace for
technical assistance, and S. X. Kannemeyer and M. Bogarde for preparation
of the skeleton. The squid beaks were identified by M. J. Imber (New Zealand
Wildlife Service, Wellington), the otoliths by G. J. B. Ross and A. Batchelor
(Port Elizabeth Museum), and the parasites by A. Verster (Veterinary Research
Institute, Onderstepoort). We are grateful to M. C. Sheldrick (British Museum,
Natural History) for allowing us to include the unpublished records from
Aldabra. For permission to examine and publish data on a specimen in his
collection, we thank the Director of the South African Museum. Permission
for publication was given by the Director of Sea Fisheries.

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6. SYSTEMATIC papers must conform to the International code of zoological nomenclature
(particularly Articles 22 and 51).

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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.

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nickles, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above exampie:
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 shou!d 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°51’S 25°39’E), collected by A. Smith, 15 January 1973.

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

7. SPECIAL HOUSE RULES

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

e.g. ‘... the Figure depicting C. namacolus ...’; *. . . in C. namacolus (Fig. 10)...’

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene

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.

Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.

PETER B. BEST
&
PETER D. SHAUGHNESSY

FIRST RECORD OF THE MELON-HEADED WHALE
PEPONOCEPHALA ELECTRA
FROM SOUTH AFRICA

VOLUME 83 PART 4 FEBRUARY 1981 ISSN 0303-2515

507.6%

AFRICAN
MUSEUM

OF THE SOUTH

CAPE TOWN

INSTRUCTIONS TO AUTHORS

1. MATERIAL should be original and not published elsewhere, in whole or in part.
2. LAYOUT should be as follows:

(a) Centred masthead to consist of
Title: informative but concise, without abbreviations and not including the names of new genera or species
Author’s(s’) name(s)
Address(es) of author(s) (institution where work was carried out)
Number of illustrations (figures, enumerated maps and tables, in this order)
(b) Abstract of not more than 200 words, intelligible to the reader without reference to the text
(c) Table of contents giving hierarchy of headings and subheadings
(d) Introduction :
(e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents
(f) Summary, if paper is lengthy
(g) Acknowledgements
(h) References
(i) Abbreviations, where these are numerous

Footnotes should be avoided unless they are short and essential.

Only generic and specific names should be underlined to indicate italics; all other marking
up should be left to editor and publisher.

4. ILLUSTRATIONS should be reducible to a size not exceeding 12 « 18 cm (19 cm including
legend); the reduction or enlargement required should be indicated; originals larger than
35 x 47 cm should not be submitted; photographs should be rectangular in shape and final
size. A metric scale should appear with all illustrations, otherwise magnification or reduction
should be given in the legend; if the latter, then the final reduction or enlargement should be
taken into consideration.

The number of the figure should be lightly marked in pencil on the back of each illustration.

5. REFERENCES cited in text and synonymies should all be included in the list at the end of
the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable):

(a) Author’s name and year of publication given in text, e.g.:

‘Smith (1969) describes... .’

‘Smith (1969: 36, fig. 16) describes...”

‘As described (Smith 1969a, 19695; Jones 1971)’
‘As described (Haughton & Broom 1927)...’
‘As described (Haughton et a/. 1927)...’

Note: no comma separating name and year
Dagination indicated by colon, not p.
names of joint authors connected by ampersand
- et al. in text for more than two joint authors, but names of all authors given in list of references.

For books give title in italics, edition, volume number, place of publication, publisher.

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.

Konun, 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.

“

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 83 Band
February 1981 Februarie
Part 4 Deel

THE SOUTH AFRICAN MUSEUM’S
MEIRING NAUDE CRUISES
PARI 12
CRUSTACEA DECAPODA
OF THE 1977, 1978, 1979°CRUISES

BRIAN KENSLEY

Cape Town Kaapstad

The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material
becomes available

Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na gelang van die
beskikbaarheid van stof

Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad

OUT OF PRINT/UIT DRUK

1, 2023, 5-8); 31-2: 425,.8, t=p.i), 503, 5. aoe
6(1, t.-p.i.), 7114), 8, 9(1-2, 7), 10(1-3),

11(1-2, 5, 7, t.—p.i.), 15(4—-5), 24(2), 27, 31(1-3), 32(5), 33, 45(1)

Kopieregnavrae aan die Suid-Afrikaanse Museum

ISBN 0 86813 006 0

Printed in South Africa by In Suid-Afrika gedruk deur
The Rustica Press, Pty., Ltd., Die Rustica-pers, Edms., Bpk.,
Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES
PART 42

CRUSTACEA DECAPODA OF THE 1977, 1978, 1979 CRUISES
By
BRIAN KENSLEY

Smithsonian Institution, Washington, D.C.

(With 11 figures)

[MS. accepted 1 October 1980]

ABSTRACT

Ninety-five species of Macrura, Anomura, and Brachyura Decapoda from deep water off
the east coast of South Africa are recorded. Of the fifteen new records for the area, Odonto-
zona spinosissima, Uroptychus edwardi, and Paralomis roeleveldae are described as new, while
the second record of Sergia inequalis Burkenroad is noted.

CONTENTS

PAGE
Introduction . : - , 49
Species list. : : , : 51
Station data. : : : 61
Systematic discussion . ; 62
Acknowledgements . : : fi!
References . : 2 ‘ : 77

INTRODUCTION

As a continuation of the reports on the South African Museum’s Meiring
Naude cruises on the east coast of South Africa, the present paper deals with
the Decapoda taken during the three cruises of 1977, 1978 and 1979. The
decapods of the two earlier cruises have already been reported (Kensley 1977a,
19775).

The area of the continental shelf investigated during all these cruises
stretches from Durban in the north to the Transkei coast (Fig. 1), in depths
ranging from 100 to 2 800 m. Although abbreviated station data are provided
for the stations mentioned in this paper, fuller information on these cruises
may be obtained from Louw (1980).

Abbreviations used throughout: SAM—South African Museum catalogue
number; SM— Meiring Naude station number; CL—carapace length; CW—
carapace width; RL—rostral length; ovig.—ovigerous; juv.—juvenile.

Ann. S. Afr. Mus. 83 (4), 1981: 49-78, 11 figs.

ANNALS OF THE SOUTH AFRICAN MUSEUM

137 4 140

262
spel Sas
250089 8516 2116 0210

e181

°
2uhS he e212

225
229

228,e226 220
231—§ °230 285; 218
227 © $221

°22
159 ve

159-60

162 161
ee

52
(geet ee m5
le 147, X53

118 Si3qlu9*

139 e
154-5

Fig. 1. Map showing localities of collecting stations.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES

SPECIES LIST

SUBORDER PENAEIDEA
Family Aristeidae

Subfamily Aristeinae
Aristaeomorpha foliacea (Risso)

Plesiopenaeus edwardsianus (Johnson) .
Plesiopenaeus nitidus Barnard
Subfamily Benthesicyminae
Bentheogennema intermedia (Bate)

* Bentheogennema pasithea (De Man)

Benthesicymus investigatoris Alcock & Anderson
Gennadas bouvieri Kemp.

Gennadas capensis Calman

Gennadas gilchristi Calman

* New record

SM Station

no.

Pane ee b.

N
MON MN

—y

ovig. 92 29

|
| [Rss

|
WwW
Ww

|
N Ww

juy.

52 ANNALS OF THE SOUTH AFRICAN MUSEUM

SM Station
no. 66 ovig. 22 99 juy.

Gennadas incertus (Balss) : : ; é ; = 126 — =

Gennadas kempi Stebbing ; , : saa oe al39

Gennadas parvus Bate. rae sh) een Sep Cem = leh

Gennadas propinquus Rathbun ; ; ; Aare a) i)

Gennadas scutatus Bouvier . : : : P . 119

|
|

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES

Gennadas tinayrei Bouvier

Family Solenoceridae
Haliporoides triarthrus Stebbing
* Hymenopenaeus halli Bruce .
Family Sergestidae

Petalidium foliaceum Bate.
*Petalidium obesum (Kr¢yer) .

Sergestes arcticus Kr¢yer

Sergestes armatus Kr¢yer

* New record

SM Station

no.

Pe ee ee eats gal Pah

ged

[element le |

ovig. 22 ~ 29

|
eae eS

[ese

Aelia te ale | al Sale ee Pease i

54 ANNALS OF THE SOUTH AFRICAN MUSEUM

SM Station
no. 6d ovig.22 29 juy.

Sergestes atlanticus H. Milne Edwards oo ee ca ge AS

ee ee ee remo Pe el jne |

Sergestes curvatus Crosnier & Forest . tories «: 133

|
el

| a

Sergestes disjunctus Burkenroad . <a ee eel

|
os

ieatel

Sergestes orientalis Hansen . 4 ; ; é) tee oS

—

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES

Sergestes sargassi Ortmann

Sergestes pectinatus Sund

Sergia creber (Burkenroad)
*Sergia gardineri (Kemp)
Sergia grandis (Sund)

* Sergia inequalis (Burkenroad)
Sergia laminatus (Burkenroad)

Sergia potens (Burkenroad)
Sergia prehensilis (Bate) .

* New record

SM Station

no.

alr a iieloe ee pees aeabapolanieen cen tS

ovig. 29 99

Pues lode sla SI Sa atl tee allealp iearleerlte lesa aslo ese tre ras ade ala eeae|

56 ANNALS OF THE SOUTH AFRICAN MUSEUM

SM Station |
no. 33 ovig.22 99 juy.

Sergia regalis (Gordon) : : : . : « 189

Sergia scintillans (Burkenroad) . . se wee «dS

ral

—

|
—

LOL [> meNrFRORe =
Lil | || | |

to
o
<4

N
© Oo
Oo—
—

| |
Re

Sergiaitalismani (Barnard). . 225.2 3) 2) 3. 128 — =

j——
Na
nS
[teal
|

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES

SUBORDER STENOPODIDEA
Family Stenopodidae
*Odontozona spinosissima sp. Nov.

SUBORDER CARIDEA
Family Oplophoridae
Acanthephyra armata A. Milne Edwards

* Acanthephyra curtirostris Wood-Mason & Alcock .

Acanthephyra eximia Smith

Acanthephyra pelagica (Risso)

* Acanthephyra prionota Foxton

Acanthephyra quadrispinosa Kemp

* New record

SM Station

no.

250

Se | Sey ese Se eal ee ht i Tt

ovig. 22
— 1
— 11
= 1
= 1
—_ 2
= 4
— 1
— 1
= 2
a 3

1 4
2 1
— 2
= 1
—_ 2
— 1
= 1
= 1
= 1
== 1
= 7
= 1
= 2
23 11
2 2
2 2
i 2
a 2
= 1
2 1
6 “
1 ie
0 0
1 ae

WwW
SS ele Aneel he beaseh |S ae el

—

juy.

Pasa

58 ANNALS OF THE SOUTH AFRICAN MUSEUM

Acanthephyra stylorostrata (Bate) .

Hymenodora gracilis Smith
* Meningodora miccyla (Chace)

Meningodora mollis Smith

* Meningodora vesca (Smith) .
Notostomus auriculatus Barnard

Notostomus elegans A. Milne Edwards

Notostomus gibbosus A. Milne Edwards
Oplophorus gracilirostris A. Milne Edwards
Oplophorus spinicauda A. Milne Edwards

Oplophorus typus H. Milne Edwards
*Systellaspis cristata (Faxon)

Systellaspis debilis (A. Milne Edwards)

* New record

SM Station
no.

218
220
221
223

| [eae eA, ae re Cohan ie

ora a a ede ee ee cee ema mee ela ie ep lela) pa

1
6
1

Reale deat: 7 ee aly eae ae eal

—_

66 ovig. 22 2

as law ees

ieee mimes de Sie ae ar ae a) me ee eae se | P|

—

oe eo

juy.

(ees tomas et en ae tees aa ee aie |

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES

Family Stylodactylidae

Stylodactylus stebbingi Hayashi & Miyake .

Family Pasiphaeidae

Eupasiphae gilesii Wood-Mason & Alcock .

Leptochela robusta Stimpson .
Parapasiphae sulcatifrons Smith

Pasiphaea meiringnaudei Kensley .

Pasiphaea sivado (Risso)

Family Pandalidae
Heterocarpus dorsalis Bate

Heterocar pus laevigatus Bate .

Heterocarpus tricarinatus Alcock & Anderson

Parapandalus richardi (Coutiére)

Plesionika longirostris (Borradaile)

Plesionika martia (A. Milne Edwards) .

Family Crangonidae

Metacrangon jacqueti bellmarleyi ae

Pontophilus sculptus (Bell)

SM Station

no.

alae leer weet aigiiay tl a

ovig. 22 2°
3 12
1 1
2 1
1 1
1 ee
_ 1
—_ 1
ae 1
_ 1
= D
— D
=e 1
= D
Lo 1
as 1
a 1
1 ce
1 ie
1 piel
Bae 1
ae 5
1 1
an 1
1 |
D D
Bue 1
se 1
1 jes

[aaa

eee a ea = al = |

60 ANNALS OF THE SOUTH AFRICAN MUSEUM

SM Station |
no. 33 ovig.22 99 juy.

SUBORDER PALINURA
Family Polychelidae

Stereomastis sculpta (Smith) . ; : : : - 234 — 1 ans a

Family Palinuridae

Projasus parkeri (Stebbing) . : 3 ‘ ; ». BIL 1 ~ = —_
235 1 — 1 1
237 5) — 2 4

SUBORDER ANOMURA

Family Axiidae

Calocaris alcocki McArdle . ; : : ; . 150 — _ 1 —

Family Lithodidae .

*Paralomis roeleveldae sp. nov. . ; ; , . 21 — — 1 —

Family Galatheidae

Munida sanctipauli Henderson . . . . . 237 1 — 1 —

Munida sp. : : ; : ; : : : = M29 — 1 — —

Family Chirostylidae

*Uroptychus edwardi sp. nov. : : : . 134 — 1 — —

Uroptychus nitidus (A. Milne Edwards) Pee res es 7/ 1 — 1 —
121.) 4 16 —

Uroptychus simiae Kensley : ‘ ‘ 3 ; 235 1 — — 1

SUBORDER BRACHYURA

Family Dromiidae

Pseudodromia spinosissima Kensley : ‘ . 163 1 aw” eal —
233 1 — — —

Family Cymonomidae

Corycodus disjunctipes (Stebbing) . ; ; 5 1 232 1 — 1 —

Cymonomus trifurcus Stebbing . ; : 5 . 129 — — 1 —
162 lie — — —
232 — — 3 —
250 if — — —
226 1 — — —

Family Homolidae

* Homolochunia valdiviae Doflein. . . . . 237 1 — — —

Family Dorippidae

Ethusa sp. . ; , : : ; ; : 2 ~ 121 1 — — —

Family Calappidae

Mursia armata De Haan. 5 ; 4 : : , 1239 1 — — —

Family Majidae | | |

Inachus guentheri (Miers) ane? i AWN Rae 3 GkSS 1 1 a

Macropodia formosa Rathbun ee : 3 £1 239. 2 gall — — —

Pleistacantha moseleyi (Miers) . : : : . 234 2 — — —

Family Hymenosomatidae - |

Hymenosoma orbiculare Desmarest . . . . 180 — — — 1

* New record

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES

SM Station
no. reNe4
Family Atelecyclidae
*Trachycarcinus glaucus Alcock & Anderson . pits eal 1
Family Geryonidae
Geryon sp. 121 —
233 2
Family Goneplacidae
Goneplax rhomboides (Linnaeus)... : meee 1
Litocheira kingsleyi (Miers) . : : : : eS =
PRY 2
Pilumnoplax heterochir (Studer) . . . . . 226 2
232 1
237 1
STATION DATA
B — Bongo net
BIT — Beam trawl
HD — Heavy dredge
RMT — Rectangular midwater traw!
SM Station
no. Position Depth (m)
63 27°10'S 33°14’E 140
88 215183259 297
96 28°14’S 32°49’E 465
99 28°23'S 32°41’E 920
119 30°14’S 31°13’E 750
121 30°32 S30°52E 900-625
123 30°33’S 30°48’E 690
126 30°39’S 30°5S9’E 464
129 30°53’S 30°31’E 850
131 30°43’S 30°40’E 780
132 30°45’S 30°42’E 830
133 31°O1/S 30°26 212
134 31°00’S 30°27’E 900
138 2072178 215 E 830
139 30°22'8 31°16’E 250
140 31°14’S 30°20’E 1120
144 31°26’S 30°06’E 212
145 31°30’S 30°04’E 1129
148 207178 31°25°E 750
150 30°14’S 31°25’E 1000
153 S001 S 31-2875 664
154 30°24’S 31°32 E 500
1157 30°05'S' 31S7E 750
159 32°40’S 28°50’E 690
160 32°45’S 28°47'E 583
162 32°55 S928. 3B 630
163 33°04’S 28°06’E 90
167 33105 2847 E 1091
168 33°14’S 28°18’E 816
170 33°10’S 28°14’E 708

* New record

ovig. 22 99

juy.

oe ae am

62 ANNALS OF THE SOUTH AFRICAN MUSEUM

SM Station

no. Position Depth (m) Gear
171 ark: Bo oe 33°16’S 28°13’E 792 RMT
173 : i. ae : 33°25’S 27°54’E 683 RMT
174 : Aan cle 337197S| 27-528 760 HD
WE) es sees Ss ae 33°25’S 27°49’E pa\|2 B
179 wat Me ‘ : 33°30’S 27°22’E 80 HD
180 o> Le ae : 33°29’S 27°21’E 80 HD
182 : : : . 33°38’S 27°49’E i SHl7/ RMT
183 ‘an ® : 33°48’S 27°47'E 474 RMT
184 4 ane : 33°39'8 27 11 E 86 HD
185 : : . : 33°39’S 27°11’E 90 HD
186 oy 8S ar 33°48’S 27°27'E 583 RMT
187 Bee G; : E 83-59 27 2c 982 RMT
189 ; ae: ; 34°04’S 27°10’E 212 B
190 ee : : 34°06’S 27°08’E 658 RMT
191 ; : fen. 34°11’S 27°08’E 542 RMT
194 : oa 4 31°34’S 30°20’E 2166 RMT
195 eS pets ae : 31°33’S 30°18’E 1050 RMT
197 : : : : 31°35’S 30°11’E 150 RMT
199 : : : : 31°33’S 30°08’E 250 RMT
201 utp ee : : 31°40’S 30°03’E 1333 RMT
203 : : : : 31°42’S 30°03’E 1750 RMT
204. ge : : 31°45’S 30°04’E pA WZ B
205 : s 5 ; 31°51’S 30°01’E 585 RMT
208 ; Oar : 31°53’S 29°48’E 1320 RMT
209 : : : 2 31°55’S 29°57’E 1260 RMT
211 5) eh : 32°00’S 29°50’E 415 RMT
214 Pea Fela : 32°15’S 29°36’E 1390 RMT
218 ne ied eee ; 32°30’S 29°13’E 916 RMT
220 pe Ym Stee 32°31’S 29°11’E 1416 RMT
221 ~~ : : 32,348 29 15°E 1170 RMT
223 es : ; 32°34’S 29°13’E 670 RMT
224 ee. <3 32°33’S 29°09’E 663 RMT
226 cate : : 32°28’S 28°58’E 710 HD
227 : : : . 32°32’S 28°55’E 790 BT
232: : ; i : 32°14’S 29°10’E 620 HD
233 soe ee : 32°15’S 29°09’E 580 BT
234 é : : : 32°15’S 29°09’E 520 BT
237 : hats ; 32°15’S 29°09’E 650 BT
239 wor : ‘ 32°14’S 29°00’E 90 BT
250 g ; 4 é 31°59’S 29°22’E 200 HD
254. 2 : ; ‘ 31°42’S 29°40’E 860 BT

SYSTEMATIC DISCUSSION
Family Aristeidae
Bentheogennema pasithea (De Man)
Bentheogennema pasithea: Crosnier, 1978: 31, figs 13c—d, 14d. [Full synonymy. ]
Previous records

Formosa; Caroline and Gilbert Islands; Indonesia; India; Somalia;
Seychelle Islands; Nosi Be, Madagascar.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 63

Material
1 ¢ CL (incl. rostrum) 12,5 mm SM 224 32°33’S 29°09’E 600-663 m.

Family Sergestidae
Petalidium obesum (Kr¢yer)
Fig. 2

Sergestes obesus Kréyer, 1859: 257, 279, pl. 4 (fig. 10a—f).

Sergestes sanguineus Chun, 1889: 538 (mastigopus larva).

Petalidium obesum: Hansen, 1896: 968; 1903: 56; 1922: 190, pl. 11 (figs 3-4). Burkenroad,
1937: 324. Wasmer, 1974: 165.

Petalidium foliaceum non Bate, Illig, 1927: 282, figs 1-5.

Previous records
Off Cape Verde, Canary Islands, Azores Islands (Hansen); off Liideritz;
Cape Point (Illig).

Remarks

Burkenroad (1937) noted that his new species P. suspiriosum differed from
other species of the genus in the gill structure, in having two arthrobranchs
above pereopod 4, instead of one as in P. obesum, or none as in P. foliaceum (see
Wasmer 1974: 165). Hansen (1922: 193), in discussing the gill formula of P.
obesum, mentioned that there was frequently a rudimentary gill above pereopod

Fig. 2. Petalidium obesum. A. Petasma. B. Outer antennular flagellum 3.

64 ANNALS OF THE SOUTH AFRICAN MUSEUM

4, and that its absence was probably due to mutilation along with bad preser-
vation. The thirty-one specimens in the present collection agree with P. foliaceum
in lacking a gill above pereopod 4.

The petasma agrees well with Hansen’s figures, although slight differences
are noted: the longer lobe of the lobus terminalis has only a single terminal hook
(three in Hansen’s description), while the shorter lobe of the lobus armatus
carries three (not two) hooks in the present material. Hansen’s figure of the
telsonic apex does not show the two distolateral spines seen in the Indian Ocean
material.

The rostrum is variable, in fact, it shows the same range of variation as
[lig (1927, fig. 1) illustrates for P. foliaceum. As Illig’s figure of the petasma
more closely resembles that of P. obesum (lacking the two distinctive stumpy
lobus armatus lobes of P. foliaceum Bate), there is strong reason to believe that
Illig’s specimens from the South Atlantic were P. obesum.

As the integument of Petalidium is very delicate, carapace lengths are diffi-
cult to measure; however, the average carapace length of four males of P. obesum
is 7,1 mm, while the two males of P. foliaceum have carapace lengths of 12,0 mm
each.

Sergia inequalis (Burkenroad)
Fig. 3

Sergestes inequalis Burkenroad, 1940: 51.
Sergestes (Sergia) inequalis: Yaldwyn, 1957: 9.

Description
Male

Rostrum with relatively slender apical spine, lacking dorsal denticle,
anterior margin oblique. Carapace with cervical sulcus becoming obsolete dor-
sally; postcervical sulcus distinct across dorsum; suprabranchial ridge strong.
Cornea reaching beyond midpoint of basal antennular peduncle segment. Latter
subequal in length to second segment; third segment somewhat shorter. Scapho-
cerite reaching to midpoint of third antennular peduncle segment, with twelve
small opaque-spot photophores. Maxilliped 3 slender, slightly longer than
pereopod |. Outer uropodal ramus with spine on outer margin at about distal
third, with twelve opaque-spot photophores near inner margin. Telson apically
acute.

Petasma: processus uncifer with small apical spine; lobus armatus short,
tapering, extended laterally, with eleven hooks; lobus accessorius subequal in
length to lobus armatus, but broader, with about twenty hooks; processus ven-
tralis broad, outer margin sinuous, apically acute, reaching to lobus accessorius
and lower lobus connectens lobe; lobus connectens lower lobe slender, extending
laterally, curved downwards, with ten to twelve terminal hooks, upper lobe short,
extended upwards; lobus terminalis directed laterally, short, squat, with three

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES 65

Fig. 3. Sergia inequalis. A. Carapace in lateral view. B. Scaphocerite. C. Outer uropodal

ramus. D. Anterior and posterior view of petasma. l.a.—lobus armatus, l.acc.—lobus acces-

sorius, l.c.—lobus connectens, |.i.—lobus inermis, |.t.—lobus terminalus, p.u.— processus
uncifer, p.v.—processus ventralis.

terminal hooks; lobus inermis extending beyond lobus terminalis, short, distally
rounded.
Previous records - 3

Dana Expedition station 3768 7°33’S 115°22’E 810m, off Sunda Islands,
Java Sea.

66 ANNALS OF THE SOUTH AFRICAN MUSEUM

Material

SAM-A16810 1 3 CL (incl. rostrum) 11,5 mm SM 194 31°34’S 30°20’E
1150-2166 m.

Remarks

Burkenroad (1940) described S. inequalis from a single male, but provided
no figures. The species has not been recorded since. From Burkenroad’s descrip-
tion of the petasma, there can be little doubt that the present male is the same
species.

Family Stenopodidae
Odontozona spinosissima sp. nov.

Figs 4-5
Description

Female

Translucent integument bearing numerous flattened spines. Carapace with
strong cervical, postcervical, and hepatic grooves; rostrum compressed, with
twelve dorsal teeth, two ventrodistal teeth, and strong ventrolateral ridge running
into orbital margin posteriorly. Orbital spine set back from margin, larger than
surrounding spines; strong antennal spine set slightly back from margin;
pterygostomian spine marginal; forwardly-directed carapace spines posterior to
cervical groove arranged in more or less vertical cinctures. Pleonal segments
dorsolaterally bearing numerous closely packed ridges and grooves; pleonite 3
largest, dorsally smoothly convex; pleura of pleonites 1 and 2 ventrally rounded,
pleuron | about one-third width of pleuron 2, bearing transverse ridges and
grooves; pleuron 4 ventrally truncate, bearing spines and ridges, small marginal
tooth posteroventrally, two posterolateral teeth; pleuron 5 with stronger postero-
ventral tooth plus three smaller posterolateral marginal teeth, bearing spines
laterally; pleuron 6 ventrally truncate, with small posteroventral denticle, bearing
spines laterally. Telson (apex damaged) with at least one pair of lateral spines;
two strong, rounded dorsal ridges present.

Eye reaching to about midlength of rostrum; mediodorsal surface of eye-
stalk armed with spines, those overlapping cornea relatively elongate. Basal
segment of antennular peduncle with flattened apically acute lobe on outer
margin. Scaphocerite with spines on outer margin, elongate setae on inner;
basal peduncular segment with one ventral and two large dorsolateral marginal
spines plus numerous smaller scattered spines; second and third segments with
scattered ventral spines. Mandibular palp 3-segmented, first segment short,
second and third segments subequal, latter with numerous setae. Maxilla | with
slender palp; distal lobe with nine spines and several setae on straight inner
margin, proximal lobe broadly rounded, setose. Maxilla 2 with broad scapho-
gnathite, slender palp, four narrow endites. Maxilliped 1 with broad bilobed
epipod, slender exopod, 3-segmented endopod. Maxilliped 2 exopod with
peduncle equal in length to flagellum; endopod merus largest segment, equal in

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES

Fig. 4. Odontozona spinosissima. A. Holotype in lateral view. B. Anterior carapace. C. Pleo-
nite 6 and uropodal base. D. Telson (damaged) and left uropod.

68 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 5. Odontozona spinosissima. A. Mandible. B. Maxilla 1. C. Maxilla 2. D. Maxilliped 1.
E. Maxilliped 2. F. Pereopod 5 dactylus.

length to dactylus and carpus together. Maxilliped 3 with elongate exopod;
elongate setae on inner margins of five distal endopodal segments. Epipodites
present on pereopods |—4. Pereopod | chela three-quarters length of carpus, equal
in length to merus. Pereopod 4 elongate, slender, propodus half length of carpus
consisting of six subsegments, with fine spinules on posterior margin; carpus of
eight subsegments; merus about two-thirds length of carpus. Pereopod 5 dactylus
biunguiculate; propodus of six subsegments; carpus of nine subsegments;
merus about two-thirds length of carpus. Pleopod 1 uniramous, peduncle very
short, ramus elongate-lanceolate, with setose margins. Uropodal basis with
strong distal tooth and several smaller marginal teeth plus several surface spines ;
outer ramus broader than inner, with eight teeth on outer margin, distally
broadly rounded; dorsal surface with two strong rounded ridges, bearing scat-
tered spines; inner ramus distally narrower than outer ramus, with four spines

THE SOUTH AFRICAN MUSEUM’S MEIRING NAULDE CRUISFS 69

on outer proximal margin, single rounded ridge dorsally, with scattered dorsal
spines.

Material
Holotype SAM-A16811 12 CL (incl. rostrum) 7,0 mm RL 2,4 mm
SM 250 31°59’S 29°22’E 150-200 m.

Remarks

Although only a single damaged specimen is available, this species is so
markedly distinctive as to deserve description.

The genus Odontozona Holthuis is characterized by the possession of a
compressed body, cinctures of anteriorly-directed spines on the carapace, maxil-
liped 3 possessing a large exopod, but lacking external spinules on the ischium,
biunguiculate dactyli on pereopods 4 and 5, and a uropodal endopod having two
dorsal ridges (Holthuis 1946: 5).

O. spinosissima bears a strong resemblance to O. sculpticaudata Holthuis,
described from a single ovigerous female from Sape Straits, east of Soembawa,
Indonesia, especially in the abdominal sculpturing. From examination of Holt-
huis’s type and from the description, these two species differ in several easily-
observed features, summarized in the following table.

O. sculpticaudata O. spinosissima
Rostral dentition . . 5/2 22
Pleon sculpture . . Fewgroovesandridges Many grooves and ridges
memonts — . . Bluntanterior tooth Rounded
present
Riconites: . | . # £Ifansverse catia No transverse carina
present
Pleura4&5 . . . Anteriorand posterior Anterior tooth absent
tooth present posterior tooth present
Uropodalrami. . . Lackingdorsalspines | Numerous dorsal spines
present
Outer uropodal ramus . 6marginal teeth 8 marginal teeth

Etymology
The specific name derives from the extremely spinose condition of the cara-
pace and pleon.

Family Chirostylidae
Uroptychus edwardi sp. nov.
Figs 6-7
Description
Female

Carapace middorsal length (excluding rostrum) almost three-quarters
greatest carapace width; dorsally smooth, gently convex, widest across branchial

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 6. Uroptychus edwardi. Holotype in dorsal view.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES Jel

Fig. 7. Uroptychus edwardi. A. Anterior sternum. B. Three basal segments of maxilliped 3
endopod. C. Antennal peduncle. D. Basal antennular segment. E. Chela. F. Dactylus and
propodus of ambulatory pereopod.

regions; anterior margins somewhat sinuous between spiciform rostrum and
anterolateral spines; posterodorsal margin concave. Ventrolateral carapace plate
ending anteriorly in short spine. Sternum with smoothly even median concavity,
with tiny median slit. Sternites of maxilliped 3 and pereopod | laterally rounded.

fp ANNALS OF THE SOUTH AFRICAN MUSEUM

Basal antennular segment with unarmed rounded distal lobe. Antennal
peduncle segments unarmed, acicle not reaching end of second segment. Cheli-
peds slender, five and a half times length of carapace (including rostrum);
dactylus about half length of propodal palm; distal half of finger and thumb
distinctly narrowed, grooved on inner faces, with outer cutting edges finely
denticulate; proximal half of dactylus bearing strong, finely denticulate process
on cutting margin, fitting closely against and distal to similar process on fixed
finger; carpus four-fifths length of propodus (including fixed finger), two distal
spines present, four or five low tubercles medioventrally; merus about two-
thirds length of carpus, with strong distodorsal spine. Propodi of ambulatory
pereopods slightly curved, with dense band of setae on ventral margin; dactyli
curved, with row of conical spines and numerous fine setae on ventral margin.

Material

Holotype SAM-A16033 1 ovigerous 9 CL (incl. rostrum) 7,0 mm
CW 8,0 mm SM 134 31°00’S 30°27’E 900 m.

Remarks

- Although only a single female of this species is available, it is sufficiently
distinct to warrant description. |
U. edwardi belongs to that group of species of Uroptychus possessing a
carapace wider than long, and lacking dorsal spination. The spiciform rostrum
and anterolateral spines distinguish U. edwardi from all other species of this
group, including U. siraji Tirmizi, U. onychodactylus Tirmizi, U. foulisi Kensley,
U. suluensis Van Dam, U. setosidigitalis Baba, U. scambus Benedict, and U.
glyphodactylus MacGilchrist. The two latter species most closely resemble the
present species, especially in the short but spiciform anterolateral carapace
spines, but both species possess broadly triangular rostra.

Etymology

Station SM 134, at which U. edwardi was captured, lies on the continental
shelf off Port Edward, hence the specific name.

Family Lithodidae
Paralomis roeleveldae sp. nov.
Figs 8-9
Description
Female

Carapace (including rostrum) slightly longer than wide, covered with short,
rounded tubercles of varying sizes; regions well defined. Gastric region strongly
convex, with large acute tubercle at middorsal point and two smaller tubercles
in posterior part; margin with two or three strong spines; cardiac region defined
by grooves; branchial region with large acute tubercle at about midpoint,

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES

Fig. 8. Paralomis roeleveldae. A. Holotype in dorsal view. B. Holotype, abdomen.

74 ANNALS OF THE SOUTH AFRICAN MUSEUM

anterior margin with two strong spines; two large contiguous spinose tubercles
at posterolateral angle, single large tubercle between posterolateral angle and
midpoint of posterior margin. Rostrum of strong inferior median spine with
three small median spines dorsal to median spine, and two strong dorsolateral
spines with a pair of small anterior spines at base. Abdomen with second somite
a single plate covered with conical tubercles. Median and lateral plates of somites
3-5 bearing rounded-flattened and smaller conical tubercles. Lateral plates of
left side bearing short spinose tubercles. On right side, lateral plate of somite 3
bearing small marginal plate at distal angle. Somite 4 with two marginal plates,
each divided marginally and carrying fine spines. Somite 5 similar to somite 4.
Somite 6 rectangular, terminal somite short, with two terminal spines. Median
plates 4 and 5 each with deep transverse groove proximally.

Eyestalks with single relatively strong terminal spine extending beyond
cornea and several tiny spines dorsally. Antennules unarmed. Basal antennal
peduncle segment with single distal spine; second segment with laterodistal spine
and three or four smaller spines at base; acicle with five strong spines (including
terminal spine) on outer margin, several smaller spines dorsally and on inner
margin. Maxilliped 3 with small spine on inner distal margin of second endopod
segment (ischium); outer surface of merus granular.

Left cheliped slightly shorter and less robust than right, spination and granu-
lation similar; spines of upper distal region of merus becoming stronger than

‘ef Ag
Oe on oa
| Kens fk ‘See

Fig. 9. Paralomis roeleveldae. A. Basal antennal segments. B. Rostrum (lateral spine broken)
and eyestalk. C. Carapace tubercles enlarged.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES a3

spinose-granulations of outer face, with several elongate spines on inner and
dorsal surfaces ; fingers of chela subequal to palm in length; propodi with several
elongate spines on inner margin. Right chela with two or three rounded proximal
cusps on cutting edges of finger and thumb, tips subacute, corneous; left chela
lacking cusps, tips subacute, corneous. Ambulatory pereopods 2-4 with strong
spines on dorsal margins of carpi and meri, surfaces granulate-spinose, lower
margins with strong spines, dactyli unarmed except for single proximal tubercle.

Material

Holotype SAM-A16045 12 CL (incl. rostrum) 50,0 mm CW 45,0 mm
SM 121 30°32’S 30°52’E 625-900 m.

Remarks

The present species appears to be most closely related to Paralomis investi-
gatoris Alcock & Anderson, 1899, from deep water off the coast of Travancore,
India. This resemblance lies especially in the overall carapace shape, similarity
of the carapace and abdominal tuberculation, and pereopodal armature. Several
differences separate the two species, however, including the lack of stronger
spinose tubercles on the carapace and marginal abdominal spines in the Indian
species, and the lack of basal rostral spines and the relatively more slender cheli-
peds in P. roeleveldae. Alcock & Anderson (1899) mention the similarity of their
species to P. aspera Faxon from the Pacific coast of Panama. Although Faxon’s
species resembles P. roeleveldae in the general shape and carapace armature, it
lacks longer carapace tubercles, elongate marginal spines, the inferior rostrum
is multidentate, and the pereopods lack elongate spines.

Paralomis seagranti Eldredge (1976) from Guam, although superficially
similar to the present species, possesses a relatively more elongate carapace,
elongate setae on the appendages, relatively short marginal carapace spines,
fewer antennal acicle spines, shorter spines on the setose ambulatory pereopods,
while lacking a large gastric spine and spines on the posterior carapace margin.

This is the first record of the genus from the South-western Indian Ocean.

Etymology

The species is named for Martina Roeleveld of the South African Museum,
in thanks for her help during the Meiring Naude cruises.

Family Atelecyclidae
Trachycarcinus glaucus Alcock & Anderson
Figs 10-11

Trachycarcinus glaucus Alcock & Anderson, 1899: 8. Alcock, 1899: 59, pl. 2 (fig. 2). Alcock &
MacGilchrist, 1905: pl. 76 (figs 1-2). Guinot & Sakai, 1970: 203.

Previous records
Off Travancore coast, southern India, 860 m.

76 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. i0. Trachycarcinus glaucus. Male in dorsal view.

Fig. 11. Trachycarcinus glaucus. A. Pleopod 1 3. B. Pleopod 2 3.

THE SOUTH AFRICAN MUSEUM’S MEIRING NAUDE CRUISES Fa

Material

SAM-A16018 1 3g CL (nel. rostrum) 23,7 mm CW 22,0 mm SM 121
625-900 m.

Remarks

The largest male from Travancore measured CL 18,5 mm, CW 14,5 mm.
The present male is thus probably the largest known.

ACKNOWLEDGEMENTS

My sincere thanks are due to Captain G. Foulis and the crew of the R.V.
Meiring Naude and to my scientific colleagues on the cruises; to the Trustees and
Director of the South African Museum, Cape Town, for making the decapod
material available to me; Dr D. Platvoet of the Zoologisch Museum, Amsterdam,
for the loan of type material of Odontozona; Carolyn Bartlett Gast (Smithsonian
Institution) for the beautiful figures of Odontozona spinosissima; Messrs Michael
Carpenter (Smithsonian Institution) and Sidney Kannemeyer (South African
Museum) for assistance with the photographs; and-to Drs F. A. Chace, Jr., and
R. B. Manning of the Department of Invertebrate Zoology, Smithsonian Insti-
tution, for critically reading and commenting on the manuscript.

REFERENCES

AtLcock, A. 1899. An account of the deep-sea Brachyura collected by the Indian Marine Survey
Ship ‘Investigator’. Calcutta: Trustees of the Indian Museum.

Autcock, A. & ANDERSON, A. R. S. 1899. Natural history notes from H.M. Royal Indian
Marine Survey Ship ‘Investigator’, Commander T. H. Heming, R.N., commanding.
Series III, No. 2. An account of the deep-sea Crustacea dredged during the surveying-
season of 1897-98. Ann. Mag. nat. Hist. (7) 3: 1-27.

Atcock, A. & MAcGiLcurisT, A. C. 1905. Illustrations of the Zoology of the Royal Indian
Marine Survey Ship ‘Investigator’, under the Command of Captain T. H. Heming, R.N.
Til. Zool. ‘Investigator’ 11, plates 68-76.

BURKENROAD, M. D. 1937. The Templeton Crocker Expedition. 12. Sergestidae (Crustacea
Decapoda) from the Lower Californian Region, with descriptions of two new species
and some remarks on the Organs of Pesta in Sergestes. Zoologica, N.Y. 22: 315-329.

BURKENROAD, M. D. 1940. Preliminary descriptions of twenty-one new species of pelagic
Penaeidea (Crustacea Decapoda) from the Danish Oceanographical Expeditions. Ann.
Mag. nat. Hist. (11) 6: 35-54.

CHuNn, C. 1889. Bericht iiber eine nach den Canarischen Inseln im Winter 1887-88 ausgefihrte
Reise. Sber. preuss. Akad. Wiss. 1889: 519-553.

CrosNigER, A. 1978. Crustacés Décapodes Peneides Aristeidae (Benthesicyminae, Aristeinae,
Solenocerinae). Faune Madagascar 46: 1-197.

ELDREDGE, L. G. 1976. Two new species of lithodid (Anomura, Paguridea, Lithodidae) crabs
from Guam. Micronesica 12: 309-315.

GuINnoT, D. & SAKAI, T. 1970. Un nouveau Trachycarcinus, T. elegans sp. nov. (Crustacea
Decapoda Brachyura). Bull. Mus. natn. Hist. nat., Paris (2) 42: 201-205.

HANSEN, H. J. 1896. On the development and the species of the Crustaceans of the genus
Sergestes. Proc. zool. Soc. Lond. 1896: 936-970

HANSEN, H. J. 1903. The Crustaceans of the genera Peralidium and Sergestes from the ‘Chal-
lenger’, with an account of luminous organs. Proc. zool. Soc. Lond. 1903: 52-79.

78 ANNALS OF THE SOUTH AFRICAN MUSEUM

HANSEN, H. J. 1922. Crustacés Décapodes (Sergestidés) provenant des campagnes des Yachts
Hirondelle et Princesse-Alice (1885-1915). Res. Camp. sci. Monaco 64: 1-232.

Ho.tuuts, L. B. 1946. The Decapoda Macrura of the Snellius Expedition. I. Temminckia 7:
1-178.

Ittic, G. 1927. Die Sergestiden der Deutschen Tiefsee-Expedition, 3. Natantia. Wiss. Ergebn.
dt. Tiefsee-Exped. ‘Valdivia’ 23: 279-354.

KeENSLEY, B. 1977a. The South African Museum’s Meiring Naude Cruises. Part 2. Crustacea,
Decapoda, Anomura and Brachyura. Ann. S. Afr. Mus. 72: 161-188.

KENSLEY, B. 19775. The South African Museum’s Meiring Naude Cruises. Part 5. Crustacea,
Decapoda, Reptantia and Natantia. Ann. S. Afr. Mus. 74: 13-44.

Kr@yer, H. 1859. Forsog til en monographisk fremstelling af kraebsdyrslaegten Sergestes.
K. danske Vidensk. Selsk. Skr. 4: 217-304.

Louw, E. 1980. The South African Museum’s Meiring Naude Cruises. Part 10. Station data
1977, 1978, 1979. Ann. S. Afr. Mus. 81: 187-205.

WaASMER, R. A. 1974. A description of Petalidium suspiriosum Burkenroad, 1937 (Decapoda,
Natantia). Crustaceana 27: 159-169.

YALDWYN, J. C. 1957. Deep-water Crustacea of the genus Sergestes (Decapoda Natantia)
from Cook Strait, New Zealand. Zoology Publs Vict. Univ. Coll. 22: 1-27.

6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature
(particularly Articles 22 and 51).

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15SA
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

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°51’S 25°39’E), collected by A. ‘Smith, 15 January 1973.

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

7. SPECIAL HOUSE RULES

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

e.g. ‘... the Figure depicting C. namacolus...; ‘...in C. namacolus (Fig. 10)...’

’ (b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene

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.

Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.

BRIAN KENSLEY

THE SOUTH AFRICAN MUSEUM’S
MEIRING NAUDE CRUISES

PART 12

CRUSTACEA DECAPODA

OF THE 1977, 1978, 1979 Gis

VOLUME 83 PART 5 FEBRUARY 1981
ob

OF THE SOUTH AFRICAN
MUSEUM

CAPE TOWN

INSTRUCTIONS TO AUTHORS

1. MATERIAL should be original and not published elsewhere, in whole or in part.
2. LAYOUT should be as follows:

(a) Centred masthead to consist of
Title: informative but concise, without abbreviations and not including the names of new genera or species
Author’s(s’) name(s)
Address(es) of author(s) (institution where work was carried out)
Number of illustrations (figures, enumerated maps and tables, in this order)
(b) Abstract of not more than 200 words, intelligible to the reader without reference to the text
(c) Table of contents giving hierarchy of headings and subheadings
(d) Introduction z
(e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents
(f) Summary, if paper is lengthy
(g) Acknowledgements
(h) References
(i) Abbreviations, where these are numerous

Footnotes should be avoided unless they are short and essential.

Only generic and specific names should be underlined to indicate italics; all other marking
up should be left to editor and publisher.

4. ILLUSTRATIONS should be reducible to a size not exceeding 12 « 18 cm (19 cm including
legend); the reduction or enlargement required should be indicated; originals larger than ©
35 x 47 cm should not be submitted; photographs should be rectangular in shape and final
size. A metric scale should appear with all illustrations, otherwise magnification or reduction
should be given in the legend; if the latter, then the final reduction or enlargement should be
taken into consideration.

The number of the figure should be lightly marked in pencil on the back of each illustration.

5. REFERENCES cited in text and synonymies should all be included in the list at the end of
the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable):

(a) Author’s name and year of publication given in text, e.g.:

‘Smith (1969) describes... .’

‘Smith (1969: 36, fig. 16) describes .. .’

‘As described (Smith 1969a, 19695; Jones 1971)’
‘As described (Haughton & Broom 1927)...’
‘As described (Haughton et al. 1927)...’

For books give title in italics, edition, volume number, place of publication, publisher.

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.

Konan, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

(continued inside back cover)

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fre PRESHWATER AMPHIPODA (CRUSTACEA) OF
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CHARLES L. GRIFFITHS

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THE FRESHWATER AMPHIPODA (CRUSTACEA) OF
SOUTH AND SOUTH WEST AFRICA

CHARLES L. GRIFFITHS
Zoology Department, University of Cape Town

(With 8 figures)
[MS. accepted 9 September 1980]

ABSTRACT

Sixteen species and one variety of freshwater Amphipoda are recognized, of which two,
Paramelita flexa sp. nov. and Sternophysinx alca sp. nov., are described as new. The fauna is
composed of three taxonomically and geographically distinct elements—a single species of
Ingolfiellidae recorded from South West Africa, a group of three Sternophysinx species which
occur in the Transvaal, and twelve species and one variety of Paramelita which are restricted to
the south-western Cape Province.

CONTENTS
PAGE
Introduction Se) AS ce ee oe : are 79
Systematics . : : : 2 é é é : 81
Family Banehadae : : . : 81
Pseudocrangonyx—Sternophysinx family eroup : 91
Family Ingolfiellidae . : : : ° . 96
Acknowledgements . : : : . . : . 96
References . - ig Cie Ne cae hy ig 96
INTRODUCTION

The freshwater amphipod fauna of South Africa was last reviewed by
Barnard (1927), who recorded a total of eleven species and one variety, all but
one of which were assigned to the genus Gammarus (=Paramelita). The taxo-
nomic positions of all these species have subsequently been revised, and four
further valid species and one invalid species have been described from the south-
ern African region. Considerable collections of unidentified material have also
accumulated. The time thus appears opportune for an updated review of the
fauna and of the distribution records of the various species. In the course of
such an analysis the collections of the South African Museum, Cape Town and
the Albany Museum, Grahamstown, as well as material sent by A. J. Cannone
of the University of the Witwatersrand have been examined. The material
includes a number of new locality records as well as two new species, which are
described below. The opportunity is also taken to provide keys to the known
species of Paramelita and Sternophysinx, and to illustrate the taxonomically
important features of each species.

Ann. S. Afr. Mus. 83 (5), 1981: 79-97, 8 figs.

ANNALS OF THE SOUTH AFRICAN MUSEUM

HOTTE arora
HOLLAND
MTS.

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im ©)
m
vu
m
Z
Zz
Uy)
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vy

. aurantius

» auricularius
. barnardi

. Capensis
Ccrassicornis
granulicornis
kogelensis
nigroculus
seticornis
spinicornis
tulbaghensis
flexa sp. nov.

U0 0 0-0 00 0070 00

Mx vozode0cn<s

Fig. 1. Map of the south-western Cape Province showing distribution records of Paramelita
species.

FRESHWATER AMPHIPODA OF SOUTH AND SOUTH WEST AFRICA 81

Only truly freshwater and subterranean species are included in this analysis,
estuarine forms having been treated by Griffiths (1976). The freshwater fauna is
clearly divided into three elements. Of these the ingolfiellid Leleupiella is known
only from northern South West Africa, Sternophysinx (three species) is recorded
only from the Transvaal, and Paramelita (twelve species and one variety) is
restricted to the south-western Cape.

In accordance with modern practice, the pereiopods are numbered 3-7, so
that they correspond to the pereon segments on which they occur. It should be
noted that, while this system is used by Holsinger & Straskraba (1973) and
Thurston (1973), authors such as Methuen (191la, 1911b) and Barnard (1916,
1927, 1966) number the pereiopods 1-5.

SYSTEMATICS
KEY TO THE GENERA OF SOUTHERN AFRICAN FRESHWATER AMPHIPODA
1. Body vermiform, coxae minute, widely separated, pleopods reduced to

SAEZ UCIOSS J ode SEASONED re ree cert eRe Sree et Leleupiella (p. 96)
Body laterally compressed, coxae well developed, contiguous or overlap-
Pimp lcOpods Taree, sDITAMOUS. 46's oscis ocd aon adn de bee ome aS 2
2. Coxae not overlapping, pereon segments 2-7 with bladder-like medioventral
PRR SSCS Vleet bership ones Sea apa Men en ara egcgh ets Sternophysinx (p. 91)
Coxae overlapping, pereon segments bearing accessory gills but lacking
Hace mOneniidl PLOCESSES. «. 5 oye os cs che ves cpa een ent Paramelita (p. 81)

Superfamily CRANGONYCTOIDEA Bousfield, 1973
Family Paramelitidae Bousfield, 1977
Paramelita Schellenberg, 1926

The species described by Barnard (1916, 1927) were transferred from Gam-
marus to Paramelita by Schellenberg (1937). In a revision of the family Gammar-
idae, Bousfield (1977) subsequently placed this genus in his new family
Paramelitidae.

Paramelita spp are restricted to fully freshwater habitats, both in surface
streams and in caves. All known records are from the south-western Cape
Province, South Africa. The recorded distribution patterns within the genus are
given in Figure 1.

KEY TO SPECIES OF PARAMELITA

feevestconspicuous, black (Fig. GA—B). el ee cn cw dee cles we dtuaielee isle dew ne ole) s 2
Eyes small, white, invisible im preserved material (Fig. 4A)...... 2.0.5... .2506.005-.. 3
2 Posterior margins of antenna 2 and pereiopods 2-7 densely setose posteriorly, especially
IMM Ne OMRCONY) eigen te Ries icc te a one. asecey shoves. Goce, ante! S tanaiere tne nigroculus var. persetosus
Posterior margins of antenna 2 and pereiopods 3—7 moderately setose, lacking setal brushes
CEN GIB) 5s, dc 6 a Sitonccy Ce EEE OC en Mite RE Fon Santee Cah ast tear a ae nigroculus
3 Peduncle of antenna 2 ¢ greatly enlarged and/or with articles 3 or 4 posterodistally lobed
Roig OOM UO EOS Nes) ee cg Mega cue eke cha tafe egaie ki aidlte, 010 6 GIRPEEA peu, Susohclew Taare odni 4

Peduncle of antenna 2 3 not enlarged, articles 3 and 4 not lobed or toothed (Figs 4A, 2D). .8

82 ANNALS OF THE SOUTH AFRICAN MUSEUM

4 Article 3 of antenna 2 3 posterodistally lobed (Figs 3A, 5A)..........cccccccececcecee 5

Article 3 of antenna 2.¢: not lobed... i io..34.0. Sos soe aeselose os ota hee. oe 6

5 Antenna 2 ¢ linear, coxa 4 quadrangular, pereiopod 3 chelate in adult § (Fig. 3A-C)....

auricularius

Antenna 2 3, article 5 attached at right angles to 4, coxa 4 posteriorly excavate, pereiopod

3-h normal (Fig. SA, G)iesacis cnda nats aad eae ee an oe flexa sp. nov.

6 Antenna 2 3, article 4 with a strong posterodistal tooth, article 5 attached normally to 4

(Figs 3D). 2.5 ocecsk betes aged Cumemn es hss cess Shek se spinicornis

Antenna 2 3, article 4 not toothed, article 5 bent at right angles to 4 (Fig. 3E).......... i

7 Coxa 4 not posteriorly excavate, dactyls of pereiopods 3-7 each with a single spinule

(Fig. 3FHG) ic ess be es a ee BA crassicornis

Coxa 4 distinctly excavate posteriorly, dactyls of pereiopods 3 and 4 with two spinules, of 5

with three spinules and of 6 and 7 with four spinules (Fig. 3H-I)............ tulbaghensis

8 Palm of gnathopod 2 distinctly oblique (Fig. 2A). . > 2s. >....2 see 9

Palm of gnathopod 2 transverse or slightly oblique (shorter than hind margin) (Fig. 2I-J)

9 Coxa 4 strongly excavate posteriorly, uropod 3 strongly setose (Fig. 4A-B)...... capensis

Coxa 4 shallowly excavate posteriorly, uropod 3 spinose, not setose (Fig. 2B—C). . barnardi

10 Coxa 4 distinctly excavate posteriorly (Fig. 2B)......... «0-12 een ee 11

Coxa 4 quadrate, lacking posterior excavation (Fig. 2H)..............0.0c0ececeeess 12

11 Peduncle of antenna 2 densely setose posteriorly, article 2 of outer ramus of uropod 3

obsolete (Fig. 2D, F) i ice ses ee ve elec a4 sa Be CRE OR eee seticornis
Peduncle of antenna 2 not strongly setose, outer ramus of uropod 3 with distinct second .

joint (Piss 2G)o. seeks o's ese oie dae oes oad oot ele alels 0b) ORIG) ee kogelensis

12 Article 2 of gnathopod 2 with a group of spines on posterior margin, palm strongly convex,

with distinet defining tooth (Big: 21)... 3.25 os 42. oe se ee granulicornis

Article 2 of gnathopod 2 not spinose, palm moderately convex, lacking defining tooth

(Bigs 20) 256 oes oa be ban 0b 5 Gab Awe wip bdo wid Reon ee aurantius

Paramelita aurantius (K. H. Barnard, 1927)
Fig. 2J
Gammarus aurantius Barnard, 1927: 173-174; pl. 10, figs 6, 16.

Remarks

One of a closely related group of species comprising P. aurantius, P. granu-
licornis, P. kogelensis, and P. seticornis. Distinguished by the quadrate coxa 4,
sparsely setose antenna 2, and absence of a defining tooth and of spines on
gnathopod 2.

Distribution
Hottentots Holland Mountains, south-western Cape.
Paramelita auricularius (K. H. Barnard, 1916)
Fig. 3A—C
Gammarus auricularius Barnard, 1916: 209-210, pl. 27, figs 26-68; 1927, 169-170.

Remarks

The ear-like lobe on article 3 of antenna 2 3 distinguishes this species from
all others except P. flexa sp. nov., from which it differs in the shape of coxa 4,

FRESHWATER AMPHIPODA OF SOUTH AND SOUTH WEST AFRICA 83

Fig. 2. A-C. Paramelita barnardi, female, 9 mm. A. Gnathopod 2. B. Coxa 4. C. Uropod 3.
D-F. Paramelita seticornis, male, 5 mm. D. Head and antennae. E. Coxa 4. F. Uropod 3.
G. Paramelita kogelensis, male, 6 mm. Uropod 3. H-I. Paramelita granulicornis, male, 6 mm.
H. Coxa 4. I. Gnathopod 2. J. Paramelita aurantius, male, 7 mm. Gnathopod 2.

ANNALS OF THE SOUTH AFRICAN MUSEUM

7 | i,
f
|
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D
E
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Fig. 3. A. Paramelita auricularius, male, 5,5 mm. Head and antennae. B—C. Male, 6 mm.

B. Pereiopod 3. C. Coxa 4. D. Paramelita spinicornis, male, 8 mm. Head and antennae.

E-G. Paramelita crassicornis, male, 8 mm. E. Head and antennae. F. Coxa 4. G. Articles

6 and 7 of pereiopod 7. H-I. Paramelita tulbaghensis, male, 7 mm. H. Coxa 4. I Articles
6 and 7 of pereiopod 7.

FRESHWATER AMPHIPODA OF SOUTH AND SOUTH WEST AFRICA 85

non-flexed antenna 2, and in the extraordinary subchelate condition of pereiopod
3 in the adult ¢.

Distribution
Top of Table Mountain only.

Paramelita barnardi Thurston, 1973
Fig. 2A—C
Paramelita barnardi Thurston, 1973: 159-168, figs 1-3.

Remarks

Similar to P. capensis but distinguished by the weakly excavate coxa 4 and
spinose (not setose) uropod 3.

Distribution

Known only from the type locality, a cave above Kalk Bay on the Cape
Peninsula.

Paramelita capensis (K. H. Barnard, 1916)
Fig. 4
Gammarus capensis Barnard, 1916: 203-205, pl. 27, figs 20-22; 1927: 169.
Paramelita ctenodactyla Schellenberg, 1926: 367, fig. 57.

Remarks

The degree of setation of the pleon segments and of pereiopods 5-7 is very
variable, ranging from the condition shown in Figure 4A to one in which the
dorsal surface of the pleon and the anterior margins of articles 2-6 of pereiopods
5—7 are densely clothed in setae. Antenna 2 may be as long as and sturdier than
antenna 1. A wide variety of forms may be collected from the same stream,
suggesting that the various forms are not of any taxonomic significance.

C. capensis may be recognized by its large size (15-25 mm) at maturity,
unmodified antennae, oblique palm of gnathopod 2, deeply excavate coxa 4,
and setose uropod 3.

Distribution
Widely distributed from Clanwilliam in the north to Bredasdorp in the east.

Paramelita crassicornis (K. H. Barnard, 1916)
Fig. 3E-G
Gammarus crassicornis Barnard, 1916: 207-209, pl. 27, figs 24-25.
Remarks

The unusual form of antenna 2 ¢, in which the peduncle is enlarged and
bent between articles 4 and 5, is found only in this species, P. flexa sp. nov. and
P. tulbaghensis. P. crassicornis is recognized by the quadrate coxa 4, the single

86 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 4. Paramelita capensis, male, 18 mm. A. Lateral aspect. B. Uropod 3. C. Telson.

spinule on article 7 of pereiopods 3-7 and the absence of a posterodistal lobe
on article 3 of antenna 2 J.

Distribution

Northern and western slopes of Table Mountain.

Paramelita flexa sp. nov.
Fig. 5
Description (of male, 7 mm)

Head slightly shorter than pereon segments | and 2 together, anterolateral
lobes deep, rounded-truncate, eyes invisible in alcohol. Antenna 1 as long as
body, flagellum 1,5 times length of peduncle, 26-articulate, accessory flagellum
5-articulate. Antenna 2 somewhat shorter but stouter than 1, article 3 strongly

FRESHWATER AMPHIPODA OF SOUTH AND SOUTH WEST AFRICA 87

Fig. 5. Paramelita flexa sp. nov., male, 7 mm. A. Head and antennae. B. Mandible. C. Max-
illa 1. D. Maxilla 2. E. Maxilliped. F. Gnathopod 2. G. Coxa 4 and pereiopod 4. H. Pereio-
pod 7. I. Uropod 3. J. Telson.

88 ANNALS OF THE SOUTH AFRICAN MUSEUM

lobed posterodistally, article 4 three times length of 3, curved ventrally, article 5
bent almost at right angles to 4, flagellum 16-articulate. Cutting-edge of mandible
5-toothed, lacinia mobilis (left side) with four teeth, spine row of nine strongly
pectinate spines, molar strongly triturative, palp articles 2 and 3 subequal, distal
half of 3 lined with dense row of short setae. Inner plate of maxilla 1 with four
plumose setae, outer plate bearing ten stout toothed spines, palp exceeding
outer plate, with five terminal spines and three subterminal setae. Inner plate of
maxilla 2 a little shorter and narrower than outer plate, both strongly setose
terminally. Inner plate of maxilliped with three short blunt spines and fifteen
plumose setae, outer plate bearing twelve medial spine-teeth and seven terminal
pectinate setae, palp densely setose medially.

Pereon segments dorsally smooth, coxae 1-3 slightly deeper than corre-
sponding segments, rounded-quadrate, coxa 4 distinctly excavate posteriorly,
5 and 6 bilobed, 7 rounded-quadrate. Gnathopods | and 2 of similar structure
but 2 slightly the larger, article 6 slightly longer than wide, palm convex, slightly
oblique, defined by three short stout spines, dactyl as long as palm. Pereiopods
3 and 4 of similar structure, article 4 slightly produced anterodistally, 5 with
five posterior spines, 6 with six pairs of spines posteriorly, 7 with four spinules
posteriorly. Pereiopods 5-6, articles 4, 5 and 6 subequal, dactyl with seven
spinules anteriorly. Pereiopod 7 similar to 5 and 6 but dactyl with nine spinules.

Pleon segments 1-3 each with a few setae along posterodorsal margin, first
pleonal epimeron rounded, 2 and 3 quadrate, each with long plumose setae just
above distal margin, posterior margin with short setae in minute notches. Pleon
segments 4-6 each with a few dorsal setae, uropod | extending slightly beyond 2,
rami equal, inner ramus of uropod 2 slightly the shorter, uropod 3 exceeding 2
by half length of outer ramus, peduncle as broad as long, inner ramus short,
half length of peduncle, terminating in a short spine and one seta, outer ramus
twice length of peduncle, three groups of spines on each margin, apex strongly
spinose, concealing minute second article. Telson slightly longer than broad,
cleft almost to base, each lobe with one large subapical spine between two long
setae, dorsal margin with two long and two short setae, plus one pair of minute
setae on lateral margin.

Holotype
Albany Museum MISC 52B, male, 7 mm.

Type locality

Palmiet River (34° 09’S 19°01’E), beneath bridge on main Elgin—Grabouw
road, south-western Cape, 20 October 1952.

Material

Two juveniles from the same sample as the type specimen and an ovigerous
female and four further juveniles collected by the author on 7 December 1979
from the same locality (SAM-A16776).

FRESHWATER AMPHIPODA OF SOUTH AND SOUTH WEST AFRICA 89

Etymology

From the Latin flexus (bending), an allusion to the flexion of antenna 2
between articles 4 and 5.
Relationships

The posterodistal lobe on article 3 of antenna 2 of this species is similar to
that found in P. auricularius. Antenna 2 in P. auricularius is, however, linear,
whereas in P. flexa sp. nov. article 5 is attached at right angles to article 4, in a
manner similar to that occurring in P. crassicornis and P. tulbaghensis. P. auricu-
larius may also be distinguished from P. flexa sp. nov. in the form of coxa 4 and
pereiopod 3 as well as by its distribution, which is limited to Table Mountain.

Paramelita granulicornis (K. H. Barnard, 1927)
Fig. 2H-I

Gammarus granulicornis Barnard, 1927: 175-177, pl. 10, figs 10-11, 20.
Remarks

The second gnathopod of this species, with its spinose article 2 and convex,
transverse palm, defined by an acute tooth, is diagnostic. The square coxa 4
and spinose posterior margin of article 2 of pereiopod 4 also aid in identification.
Distribution

Hottentots Holland Mountains.

Paramelita kogelensis (K. H. Barnard, 1927)
Fig. 2G
Gammarus kogelensis Barnard, 1927: 172-173, pl. 10, figs 9, 21.
Remarks

Closely related to P. seticornis but differs in the degree of setation of the
peduncle of antenna 2 (both species have the flagellum setose) and in details
of the spination of the pereiopods and of uropod 3.

Distribution
Hottentots Holland Mountains.

Paramelita nigroculus (K. H. Barnard, 1916)
Fig. 6
Gammarus nigroculus Barnard, 1916: 206-207, pl. 27, fig. 23; 1927: 168-169.
Remarks

The dark eyes distinguish the species, which is otherwise similar to P.
capensis. Although a variety based on degree of setation (var. persetosus) 1s
recognized, the degree of setation is very variable, even within individual
samples.

90 ANNALS OF THE SOUTH AFRICAN MUSEUM

"i Hi [Ze

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QQ

SF,
G _—
Ga =,
S. 87
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BR AH
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A
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a
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a
yo
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Fig. 6. A. Paramelita nigroculus var. persetosus, male, 13 mm. Head and antennae. B-—C. Para-
melita nigroculus, male, 12 mm. B. Head and antennae. C. Coxa 4 (both sides).

Distribution

The most common and widely distributed Paramelita species, occurring
from Tulbagh and the Cedarberg in the north to Swellendam in the east.

Paramelita seticornis (K. H. Barnard, 1927)
Fig. 2D-F
Gammarus seticornis Barnard, 1927: 171-172, pl. 10, figs 7, 17.
Remarks

A small form, mature at 5 mm, distinguished from the allied P. kogelensis
by the form of antenna 2 3.

Distribution
Hottentots Holland Mountains.

FRESHWATER AMPHIPODA OF SOUTH AND SOUTH WEST AFRICA 91

Paramelita spinicornis (K. H. Barnard, 1927)
Fig. 3D
Gammarus spinicornis Barnard, 1927: 174-175, pl. 10, figs 8, 18-19.

Remarks
The form of antenna 2 ¢ in this species is characteristic.

Distribution
Hottentots Holland Mountains eastwards to Swellendam.

Paramelita tulbaghensis (K. H. Barnard, 1927)
Fig. 3H-I
Gammarus tulbaghensis Barnard, 1927: 170-171, pl. 10, figs 5, 15.

Remarks

The enlarged and bent peduncle of antenna 2 ¢ is similar to that of P.
crassicornis, but coxa 4 is excavate posteriorly and there are more spinules
on the dactyls of the pereiopods (two in pereiopods 3 and 4, three in pereiopod 5,
and four in pereiopods 6 and 7). Lacks the lobe on article 3 of antenna 2 g
found in P. flexa sp. nov.

Distribution

Known only from the type locality in the Sneeuwgat Valley, Tulbagh,
south-western Cape.

Pseudocrangonyx—Sternophysinx family group (Bousfield, 1977)
Sternophysinx Holsinger & Straskraba, 1973

Originally erected to accommodate Crangonyx robertsi and two new species
from the Transvaal, Sternophysinx was transferred from the Gammaridae to a
new Superfamily Bogidielloidea by Bousfield (1977) and again to the Super-
family Crangonyctoidea by Bousfield (1978). The genus has not been recorded
outside the Transvaal and is hence geographically isolated from Paramelita.
Morphologically Sternophysinx may readily te distinguished from Paramelita
by the smaller, non-overlapping coxae and distinctive bladder-like sternal
processes on pereon segments 2-7.

KEY TO SPECIES OF STERNOPHYSINX

1 Article 2 of pereiopods 5—7 widened, with distinct posterodistal lobe (Fig. 7A-B)........ Z
Article 2 of pereiopods 5—7 narrowly tapering, without posterodistal lobe (Fig. 7C—D)... .3

2 Articles 6, or 5 and 6 of pereiopods 5-7 bearing long threadlike setae posteriorly, dactyls
pete SETI CMOE SPINGsy (EIS) GIAD: 6c. Saki oS onde aie. ate MER een es ea ee Riess filaris
Articles 5 and 6 of pereiopods 5-7 lacking thread-like setae, dactyls with single anterior
FERRE CRSE, TNO Ue NS 2 Oe pee eee ee ne ee a SR oT tn robertsi

3 Palm of gnathopods 1 and 2 evenly convex, body length about 5 mm at maturity........

; transvaalensis
Palm of gnathopods 1 and 2 excavate (Fig. 8H, I), body length at maturity about 10-12 mm
alca sp. nov.

92 ANNALS OF THE SOUTH AFRICAN MUSEUM

Sternophysinx alca sp. nov.
Figs 7D, 8
Description (of female, 15 mm)

Antenna 1 about half length of body, flagellum 22-articulate, accessory
flagellum uni-articulate, about as long as first segment of primary flagellum.
Antenna 2 somewhat shorter than 1, gland cone prominent, flagellum 9-articu-
late. Mandible with large triturative molar, spine row of eight spines, palp well
developed, articles 2 and 3 subequal. Palp of maxilla 1 bi-articulate, bearing
seven terminal spines, outer plate with six serrate spines apically, inner plate
with two terminal plumose setae. Both inner and outer plates of maxilla 2
strongly setose apically, inner plate with two long subterminal plumose setae.
Inner plate of maxilliped armed with three short thick spines, outer plate with
thirteen medial and terminal spines, palp 4-segmented.

Pereon dorsally smooth. Coxa 1 slightly produced anteroventrally, coxae
2-4 quadrate, 4 not strongly excavate posteriorly, 5 and 6 bilobed, 7 semi-
circular. Gnathopod 1 short but stout, palm defined by large pair of spines,

Fig. 7. Pereiopod 7 in Sternophysinx species. A. S. filaris, male, 8 mm. B. S. robertsi, male,
8mm. C. S. transvaalensis, male, 5 mm. D. S. alca, female, 13 mm.

FRESHWATER AMPHIPODA OF SOUTH AND SOUTH WEST AFRICA 93

posterior margin bearing three further pairs of large spines, palmar margin lined
by short setae, with semicircular excision and small tooth at its midpoint,
dactyl exceeding palm, nail long, bearing small secondary process at its base.
Gnathopod 2 as | but article 2 slender, more elongate and without anterior
setae. Pereiopods 3 and 4 with article 2 slightly expanded and with long slender
setae equally spaced along posterior margin. Pereiopod 5 considerably shorter
than 6 or 7, article 2 not expanded, lacking posterodistal lobe, dactyl with two
spines. Pereiopods 6 and 7 similar to 5 but much longer, extending to end of
uropods, dactyl with single spine. Pereon segments 2—7 each with bladder-like ©
median sternal process.

Pleon segments 1—4 each with few small setae along posterodorsal margin.
Pleonal epimera rounded-quadrate, two small setules at posterodistal corner of
each, posterior margins almost straight. Uropod 1 extending slightly beyond 2,
outer ramus slightly the shorter, both rami dorsally and terminally spinose.
Uropod 3 considerably exceeding 1 and 2, outer ramus with five groups of setae
along ventral and four along dorsal margin, inner ramus minute, rounded,
bearing single terminal seta. Telson slightly longer than broad, apically emargi-
nate, each lobe with four large terminal spines.

Holotype
SAM-A16775, female, 13 mm.

Type locality

Peppercorn’s Cave, Makapansgat, Transvaal (24°09’S 29°12’E) collected
from a small clear pool, June 1979, by A. J. Cannone of the Zoology Depart-
ment, University of the Witwatersrand.

Material

Twenty-five individuals from Peppercorn’s Cave and from Ficus Cave,
Makapansgat.

Etymology

Named for Dr Al Cannone, who discovered this species and allowed the
author to describe it.

Relationships

The excavation and tooth on the palm of gnathopods | and 2 readily
identify this species. S. alca sp. nov. is also distinguished from S. robertsi and
S. filaris by the narrowly tapering article 2 of pereiopods 5—7, and from S.
transvaalensis by its larger body size and by details of spination of the pereiopods.
The species is found in the same pools as S. robertsi.

94 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 8. Sternophysinx alca sp. nov., female, 15 mm. A. Lateral aspect. B. Accessory flagellum.
C. Mandible. D. Lower lip. E. Maxilla 1. F. Maxilla 2. G. Maxilliped. H-I. Gnathopods
1,2. J. Uropod 3. K. Telson.

FRESHWATER AMPHIPODA OF SOUTH AND SOUTH WEST AFRICA 95

Sternophysinx filaris Holsinger & Straskraba, 1973
Fig. 7A

? Eucrangonyx robertsi (partim.) Methuen, 1911a: 948-957, pls 49-51.
Sternophysinx filaris Holsinger & Straskraba, 1973: 75-76, figs 2-3.

Remarks

Distinguished by the thread-like setae along the posterior margins of
articles 5 and 6 of pereiopods 5 and 6 and of article 6 of pereiopod 7. The material
reported from Irene by Methuen (191la) should probably be allocated to S.
filaris, rather than S. robertsi, the latter being positively known only from the
Makapan Caves area some 200 km to the north.

Distribution
Sterkfontein Caves, and (?) Irene, Transvaal.
Sternophysinx robertsi (Methuen, 1911)
Fig. 7B

Eucrangonyx robertsi Methuen, 1911a: 948-957, pls 49-51; 19116: 96-101.
Barnard, 1927: 141, 209.
[Non]|Crangonyx robertsi: Barnard, 1949: 523-525 (=Sternophysinx transvaalensis).

Remarks

The expanded article 2 of pereiopods 5-7 distinguishes this species from
S. transvaalensis and S. alca sp. nov., while the posterior pereiopods are shorter
than those of S. filaris, lack thread-like seta on articles 5 and 6 and have only a
single spine on article 7. The material identified as S. robertsi by Barnard (1949)
—SAM-A8372—has been re-examined and transferred to S. transvaalensis.

Distribution
Makapan Caves and springs in the vicinity.
Sternophysinx transvaalensis Holsinger & Straskraba, 1973
Fig. 7C
Crangonyx robertsi (non Methuen, 1911a): Barnard, 1949: 523-525.
Sternophysinx transvaalensis Holsinger & Straskraba, 1973: 76-79, figs. 4—5.
Remarks

The small size, smoothly convex palm of gnathopods 1 and 2, unexpanded
article 2 of pereiopods 5-7, and absence of long setae on these appendages serve
to identify this species. The material identified as Crangonyx robertsi by Barnard
(1949) has been re-examined and the structure of pereiopod 7 (Fig. 7C) clearly
identifies it as belonging to S. transvaalensis.

Distribution

Surface streams in the northern Drakensberg region.

96 ANNALS OF THE SOUTH AFRICAN MUSEUM

Suborder INGOLFIELLIDEA Hansen, 1903

Family Ingolfiellidae Hansen, 1903
Leleupiella Karaman, 1959
Leleupiella eggerti Ruffo, 1964

Leleupiella eggerti Ruffo, 1964: 1019-1031, figs 1-2.
Ingolfiella opisthodorus Barnard, 1966: 189-197, figs 1-2.

Remarks

First described by Ruffo (1964) from Tsumeb, this species was subsequently
redescribed as new in a posthumous paper by Barnard (1966). No further
material has been recorded. L. eggerti is the only ingolfiellid recorded from
South or South West Africa and is notable for its large size (up to 25 mm). The
elongate body form, minute coxae and carpochelate gnathopods readily dis-
tinguish the Ingolfiellidae from members of the Crangonyctoidea.

Distribution

Recorded from boreholes near Tsumeb, South West Africa.

ACKNOWLEDGEMENTS

My thanks to Dr A. J. Cannone, who initiated this project by bringing to
my attention the material described herein as Stenophysinx alca sp. nov.
Additional material was kindly lent by the South African Museum and the
Albany Museum, Grahamstown. Mrs Leonora Freeland typed the manuscript.

REFERENCES

BARNARD, K. H. 1916. Contributions to the crustacean fauna of South Africa. 5. The Amphi-
poda. Ann. S. Afr. Mus. 15: 105-302.

BARNARD, K. H. 1927. A study of the freshwater isopodan and amphipodan Crustacea of
South Africa. Trans. R. Soc. S. Afr. 14: 139-215.

BARNARD, K. H. 1949. A fresh-water amphipod from Mont-aux-Sources. Ann. Natal Mus.
5 23—52 5.
BARNARD, K. H. 1966. The occurrence of the genus Ingolfiella (Crustacea, Amphipoda) in
South Africa, with description of a new species. Ann. Mag. nat. Hist. 13: 189-197.
BOUSFIELD, E. L. 1973. Shallow-water gammaridean Amphipoda of New England. New York:
Cornell University Press.

BOUSFIELD, E. L. 1977. A new look at the systematics of gammaroidean amphipods of the
world. Crustaceana, Suppl. 4: 282-316.

BOUSFIELD, E. L. 1978. A revised classification and phylogeny of amphipod crustaceans.
Trans. R. Soc. Can. (1V) 16: 343-390.

GRIFFITHS, C. L. 1976. Guide to the benthic marine amphipods of southern Africa. Cape Town:
South African Museum.

HOLsINGER, J. R. & STRASKRABA, M. 1973. A new genus and two new species of subterranean
amphipod crustaceans (Gammaridae) from South Africa. Ann. Spéléol. 28: 69-79.

METHUEN, P. A. 1911a. Onan amphipod from the Transvaal. Proc. zool. Soc. Lond. 2: 948-957.

METHUEN, P. A. 19115. Transvaal Crustacea. Part 2. Notice of a freshwater amphipod from
South Africa. Ann. Transv. Mus. 3: 96-101.

FRESHWATER AMPHIPODA OF SOUTH AND SOUTH WEST AFRICA 97

Rurro, S. 1964. Studi sui Crostacei Anfipodi. 58. Un nuovo Ingolfiellide delle acque sotter-
ranee dell’ Africa di Sud Ovest. Boll. Zool. 31: 1019-1034.

SCHELLENBERG, A. 1926. Die Gammariden der Deutschen Siidpolar-Expedition 1901-1903.
Dt. Siidpol.-Exped. 18: 235-414.

SCHELLENBERG, A. 1937. Kritische Bemerkungen zur Systematik der Siisswassergammariden.
Zool. Jb. 69: 469-516.

TuHursTon, M. H. 1973. A new species of Paramelita (Crustacea: Amphipoda) from South
Africa. Ann. S. Afr. Mus. 62: 159-168.

—

i *e ~ oie : a -

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* *
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i
u =
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:
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6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).

Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15SA
Nucula (Leda) bicuspidata hepa 1845: 37.
Leda plicifera A. Adams, : 50.
Laeda bicuspidata Hanley, ies: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

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°51’S 25°39’E), collected by A. ‘Smith, 15 January 1973.

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

7. SPECIAL HOUSE RULES

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

e.g. ‘... the Figure depicting C. namacolus ...’; ‘. . . in C. namacolus (Fig. 10)...’

' (b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
oe oa Toit but. A. L. du Toit; Von Huene but F. von Huene

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.

Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.

CHARLES L. GRIFFITHS

THE FRESHWATER AMPHIPODA (CRUSTACEA) OF
SOUTH AND SOUTH WEST AFRICA

|| VOLUME 83 PART 6 FEBRUARY 1981 | ISSN 0303-2515

Ba 7.0%

"OF THE SOUTH AFRICAN >
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| CAPE TOWN.

INSTRUCTIONS TO AUTHORS

1. MATERIAL should be original and not published elsewhere, in whole or in part.
2. LAYOUT should be as follows:

(a) Centred masthead to consist of
Title: informative but concise, without abbreviations and not including the names of new genera or species
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5. REFERENCES cited in text and synonymies should all be included in the list at the end of
the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable):

(a) Author’s name and eo of publication given in text, e.g.:

‘Smith (1969) describes .

‘Smith (1969: 36, fig. 16) eee

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For books give title in italics, edition, volume number, place of publication, publisher.

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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. gen. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Gouus (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.

(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 83 _ Band
February 1981 Februarie
Part’ '=.6) \" Deel

THE GENERA DICYNODON AND DIICTODON
AND THEIR BEARING ON THE CLASSIFICATION
OF THE DICYNODONTIA (REPTILIA, THERAPSIDA)
By

MICHAEL A. CLUVER
&
NICHOLAS HOTTON III

Cape Town Kaapstad

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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

THE GENERA DICYNODON AND DIICTODON AND THEIR
BEARING ON THE CLASSIFICATION OF THE DICYNODONTIA

(REPTILIA, THERAPSIDA)
By
MICHAEL A. CLUVER
South African Museum, Cape Town
&

NICHOLAS HorTToN III
Smithsonian Institution, Washington, D.C.

(With 30 figures and 1 table)

[MS. accepted 7 October 1980]

ABSTRACT

The Late Permian dicynodont genera Dicynodon, Oudenodon, Diictodon and Kingoria are
diagnosed on the basis of skull details seen in the primary type material. Many of the species
previously included under Dicynodon have been referred to other genera, but no attempt at
assessing the validity of these types at the species level has been made. Diictodon has the greatest
stratigraphical range among the four genera, occurring throughout the three zones of the Lower
Beaufort of the South African Karoo, while the characteristic features of Dicynodon can be
identified in modified form in a number of Triassic genera. Dicynodon and Oudenodon appear
to be more closely related to each other than to Diictodon. Kingoria, with several unusual
features in the palate and lower jaw, holds the least in common with other Late Permian
genera. Recognition of the characters distinguishing the four genera from each other should
aid future studies on the systematics and evolution of this important therapsid group.

CONTENTS

PAGE
Introduction . ; : : ; : : : : ; 99
General skull morphology - . : : : : é : : 101
The genus Dicynodon . : ; : : ; : : : : 106
MieRt CRS TOHACHOUONL: Ao rc. ree S LOOMS ee yl Os it7
The genus Diictodon be aa lcd UL le Mas a Bae eh Bi, £22
The genus Kingoria . : 136
Interrelationships of Doesoree Oipe tee Deecte and Reneoee 141
Acknowledgements . 3 ; E : ‘ ; 5 : : : 143
References § , : : : : : : ; , : ; 143
Abbreviations . ‘ , : 3 : ‘ ; : ; : : 145

INTRODUCTION

The infra-order Dicynodontia represents a Late Permian and Triassic
radiation of herbivorous therapsids. Its first appearance in the lowermost
Tapinocephalus Zone of the South African Karoo’s Beaufort Group coincides

Ann. S. Afr. Mus. 83 (6), 1981: 99-146, 30 figs, 1 table.

100 ANNALS OF THE SOUTH AFRICAN MUSEUM

with that of the earliest known South African land vertebrates. Although dicyno-
donts, mainly of Triassic age, are now known from east Africa, India, China,
Antarctica, Europe, and North and South America, the most complete record
of the group is the South African one against which the other more restricted
occurrences must be compared.

The evolution of the Dicynodontia, as far as can be traced in features of
the skull morphology, chiefly involved numerous modifications of a basic inno-
vation of the feeding process, and in terms of general skull structure the earliest
and latest known representatives of the infra-order are essentially similar. Chief
among the departures from the primitive condition are the tendency for replace-
ment of the upper and lower dentitions by horny beaks, the establishment of a
sliding quadrate-articular jaw joint, and the lateral migration of a portion of the
external jaw adductor musculature to a new site of origin on the lateral surface
of the temporal arch.

The dicynodont skull is consequently a highly distinctive one, and from the
time of its original description by Owen (1845a) has frequently been described
and figured in its many variations. However, the great abundance of specimens
collected from numerous Beaufort localities, and the preference for detailed skull
roof characters shown in the taxonomic work of earlier palaeontologists, com-
bined to create a large number of poorly diagnosed dicynodont ‘species’ divided
among a number of ‘genera’ of doubtful validity. In the bibliographic work of
Haughton & Brink (1954), which takes into at least partial account revisions by
Van Hoepen (1934) and Toerien (1953), no fewer than 111 South African species
are assigned to the catch-all genus Dicynodon. Since the bulk of these dicyno-
donts is known from the Permian part of the Beaufort succession, it is among
these forms that the taxonomic confusion is at its worst.

It has long been recognized that this state of affairs has served to obscure
the potential importance of the Dicynodontia in a variety of fields of study.
Attempts at rectifying the situation have resulted in a number of detailed studies
on selected dicynodont groups; the cranial morphology of Placerias (Camp &
Welles 1956), Kingoria (Cox 1959), Daptocephalus (Ewer 1961), Lystrosaurus
(Cluver 1971), and Oudenodon (Keyser 1975) is now well known and these genera
serve as standards of comparison with other forms.

Following the work of Toerien (1953), more attention has been paid to
details of the palate and lower jaw (e.g. Cluver 1970, 1974a, 1974b, 1975;
Keyser 1975). Cluver (1970) recognized previously undescribed cranial features
in specimens of Permian age identified as Dicynodon testudirostris. Subsequent
comparisons have shown that these features are not present in the type specimen
(BMNH 36233) of Dicynodon lacerticeps, upon which Owen’s (1845a) original
description of Dicynodon was based. Rather, they are found in the type specimen
(AMNH 5308) of Diictodon galeops Broom, 1913. It is now clear that the genus
Diictodon is valid and distinct from Dicynodon, has greater stratigraphical and
geographical ranges than previously suspected, and is abundant in certain Upper
Permian Karoo localities (Cluver & Hotton 1979). Furthermore, comparisons

THE GENERA DICYNODON AND DIICTODON 101

based on type material confirm the validity of the genera Oudenodon and
Kingoria, as revised and defined by Keyser (1975) and Cox (1959) respectively.
In the following sections, expanded diagnoses of these two genera are included
to allow full comparison with Dicynodon and Diictodon. Diagnoses of these four
important dicynodont genera help to remove much of the uncertainty surround-
ing the status of the type specimens currently assigned to Dicynodon. Locality
and stratigraphic information with respect to type specimens has been taken
from Haughton & Brink (1954) and Kitching (1970, 1977). The latter author’s
division of the Permian part of the Beaufort group into Tapinocephalus, Ciste-
cephalus, and Daptocephalus zones is followed.

GENERAL SKULL MORPHOLOGY

The highly distinctive skull structure of dicynodonts (Figs 1-9) is modelled
around a jaw mechanism unique among tetrapods. The preorbital portion of the
skull is short, and the temporal region expanded to accommodate a greatly
enlarged temporal musculature. The temporal muscles arose medially from a
broad, flat process of the postorbital that covers much of the parietal in most
forms.

The highest areas of origin are marked by bony parietal ridges which in
some dicynodonts meet in the midline to form a longitudinal crest behind the
parietal foramen. Lateral portions of the temporal muscles arose from a flattened
zygoma formed chiefly by the squamosal, and posterior portions from a laterally
expanded process of the squamosal that slants downward and forward with
respect to the zygoma.

The quadrate is inserted into the distal end of the ventral process of the
squamosal and is covered laterally by the quadratojugal (Figs 3-4). In lateral
aspect the squamosal process appears to be suspended from the back of the
skull, and its function as a support for the quadrate and an origin for jaw
muscles is most accurately reflected by the term suspensorium.

The lateral expansions of the squamosals give a plate-like configuration to
the otherwise rather massive occiput (Fig. 8). The bones shared between the
occiput and the short, stout basis cranii tended to fuse very early in life.

The jaws (Figs 3, 7-9) are short, stout, beak-like and generally toothless.
In the upper jaw the premaxillae are fused in the midline and firmly sutured to
the maxillae and more posterior parts of the palate. In the lower jaw the dentaries
are fused at the symphysis, which is remarkably robust and deep. Coronoid
bones are lacking and prominent coronoid processes are seldom developed on
the dentaries. In life, the temporal musculature slanted sharply forward from
its origins in the temporal fossa to insert on the coronoid region and on the
lateral face of the dentary, about half-way between the articulation of the lower
jaw and the tip of the symphyseal beak. In some forms the lateral part of the
insertion, which is bounded dorsally by the coronoid margin of the dentary and
ventrally by the mandibular fenestra, is expanded laterally into a shelf-like
structure (lateral shelf, Crompton & Hotton 1967) (Figs 3, 8). Deeper portions

102 ANNALS OF THE SOUTH AFRICAN MUSEUM

Sq.

[ss
3cm

Fig. 1. Dicynodon sp. SAM-—B88. Skull in dorsal view.

of the temporal musculature inserted inside the jaw ramus, in and around the
adductor (Meckelian) fossa (Cluver 1974a, 1975).

The articulation of the lower jaw is unique in that both bearing surfaces
are essentially convex (Figs 3, 7, 9), that of the articular being about twice as
long as that of the quadrate. This can be construed as clear evidence of longi-
tudinal sliding of the lower jaw during the bite, and the angle at which muscular
force was applied shows that the motion during which the bite was most effective
was one of retraction.

The toothless margins of the premaxillae and maxillae are usually sharp-
edged. Except in Endothiodon and its closest allies, the middle of the maxilla is

THE GENERA DICYNODON AND DIICTODON 103

Fig. 2. Dicynodon sp. SAM-—B88. Stereophotograph of skull in dorsal view. Scale: 5 cm.

thickened and produced downward as a caniniform process. Because of its thick-
ness the caniniform process is triangular in cross-section presenting lateral,
medial and posterior surfaces. In those forms in which it is present, the tusk
erupts from the ventromedial surface of this process. In some forms, small non-
caniniform teeth erupt from the flat medial surface of the maxilla (and in some
cases from the premaxilla as well). A secondary palate, analogous to that which
is found in cynodont mammal-like reptiles and mammals, is present and consists
chiefly of a broad, plate-like posterior expansion of the fused premaxillae.

A finely-punctate sculpture is developed on the anterior and dorsal surfaces
of the dentary symphysis, on the external surfaces of the premaxillae and pala-
tines (in most forms), and on most external surfaces of the maxillae. It may be
assumed. that during life these surfaces were covered by horn because of their
resemblance to bone surfaces that are covered by horn in living turtles and birds.

_ The three-sided caniniform process exhibits sculpture on its medial and
lateral surfaces, but its posterior surface is smooth. The large masticatory
muscles passed just behind the posterior surface, the smoothness of which
suggests a place of attachment for a large and muscular Mundplatte, which in
life lay in front of the temporal musculature in the corner of the mouth (Cluver
1975). The posterior surfaces of the caniniform processes thus provide useful

104 ANNALS OF THE SOUTH AFRICAN MUSEUM

3cm

Fig. 3. Dicynodon sp. SAM-B88. Skull and lower jaw in lateral view.

landmarks for the corners of the mouth, and the mouth opening can be delimited
posteriorly by a line drawn between the medial sides of the backs of the canini-
form processes at the general level of the ventral margins of the premaxillae. The
posterior margin of the mouth, defined here in functional terms, corresponds
closely to that defined by Cruickshank (1968) on more exclusively anatomical
grounds, i.e. the front of the choanae.

As confirmed by Agnew (1959), the skull is akinetic, and premaxillae,
maxillae, palatines, pterygoids and ectopterygoids are firmly knit in the palate.
In addition, the pterygoids are so closely and complexly sutured to the basis

THE GENERA DICYNODON AND DIICTODON 105

cranii a little behind their midlength (Olson 1944) (Figs 5-6) that, except in
serial section, they appear fused to it.

The blade-like palatine rami of the pterygoids are elongate and oriented
anteroposteriorly; there are no transverse pterygoid flanges. Between them the
palatine rami enclose a deep, rather broad vault, into the front of which the
choanae open. Anteriorly this vault, which may be termed the interpterygoid
fossa, is partially roofed by the vomers and dorsal wings of the palatines (Fig. 5).
The vomers enter the interpterygoid fossa as a single midline structure arising
from the back of the premaxilla between the choanae, and bifurcate posteriorly
at a variable distance behind the secondary palate. This bifurcation forms the
anterior margin of the variably narrow fusiform or ovoid interpterygoid vacuity,
which pierces the roof of the interpterygoid fossa. Through the interpterygoid
vacuity the ventral edge of the slender parasphenoid rostrum may be seen in
well-preserved and carefully prepared skulls. Only the interpterygoid vacuity,
so restricted, is homologous with the phylogenetically ancient interpterygoid
vacuity of less specialized reptiles and anthrocosaurian amphibians, and pre-
sumably was inherited with little change from such ancestors. The interpterygoid
fossa, which has come to surround the interpterygoid vacuity, is a new entity
formed in dicynodonts by rearrangement of the palatine rami of the pterygoids

Fig. 4. Dicynodon sp. SAM-B88. Stereophotograph of skull in lateral view. Scale: 5 cm.

106 ANNALS OF THE SOUTH AFRICAN MUSEUM

lab. fos.

3cm

Fig. 5. Dicynodon sp. SAM-—B88. Skull in ventral view.

and the formation of a secondary palate. The fossa is as distinctive of dicyno-
donts as the structure and function of their peculiar jaw articulation, to which
its development is probably closely related.

THE GENUS DICYNODON (Figs 1-11)

Dicynodon Owen
Diagnosis

Medium-sized to large dicynodonts (average skull length 100 mm to over
400 mm), single pair of maxillary tusks in upper jaw, lower jaw edentulous. Post-
orbitals tend to cover parietals behind parietal foramen. Septomaxilla merges

THE GENERA DICYNODON AND DIICTODON 107

smoothly with outer surface of snout, does not meet lacrimal. Low boss formed
over external nares by nasals. Caniniform process of maxilla arises as ventral
extension of palatal rim. Palatal rim sharp-edged, uninterrupted by notch.
Palatal portion of palatine large, makes short contact with premaxilla. Vomers
form long, narrow septum in interpterygoid fossa, interpterygoid vacuity short.
Ectopterygoid small, displaced laterally. Labial fossa present between maxilla,
palatine and jugal. Pterygoid makes short contact with maxilla. Basioccipital
tubera separated by intertuberal ridge. Fused dentaries carry narrow dentary
tables .dorsal edge of dentary carries deep sulcus behind dentary tables. Rear of
dentary extended dorsally to form weak coronoid process. Mandibular fenestra
large, bounded dorsally by lateral dentary shelf.

Dicynodon lacerticeps Owen

Dicynodon lacerticeps Owen, 1845a: 59, pls 3-4,

Type specimen
Skull and lower jaw, BMNH 36233.

Locality

Tarka prolongation of the Winterberg, Cape Province.

Fig. 6. Dicynodon sp. SAM-—B88. Stereophotograph of skull in ventral view. Scale: 5 cm.

108 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 7. Dicynodon sp. SAM-—B88. Stereophotograph of lower jaw in lateral view. Scale: 5 cm.

Horizon

Daptocephalus Zone.
Diagnosis
As for genus.

Remarks

Owen published two accounts of Dicynodon lacerticeps in 1845, both based
on a lecture delivered by him before the Geological Society of London on 8
January of that year. The paper which appeared in the Transactions of the
Geological Society (Owen 1845a) with illustrations of the fossil specimens is
regarded as having priority here; in a subsequent paper in the same journal
Owen (1856: 233) refers to this description as ‘my former account’ of the new
forms. The paper which appeared in the Proceedings of the Geological Society
(Owen 18455) is a shortened version of the Transactions article.

One reason for the uncertainty surrounding the status of the many described
species of Dicynodon is the poor preservation of the type specimen of Dicynodon
lacerticeps itself (Figs 10-11). The specimen is a fairly complete skull and lower
jaw, but much of the surface bone is damaged and sutures are subsequently
difficult or impossible to trace. However, recent preparation has exposed several

109

THE GENERA DICYNODON AND DIICTODON

boc.

3cm

Po
noice

gee te ee
. .

1ew.

dorsal vi

jaw in

Fig. 8. Dicynodon sp. SAM-B88. A. Occipital view of skull. B. Lower

110 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 9. Dicynodon sp. SAM-B88. Stereophotograph of lower jaw in dorsal view. Scale: 5 cm.

key areas of the skull and enough can now be seen to distinguish it from other
dicynodont genera founded on better type material.

The dorsal skull surface is gently curved from the parietal region to the tip
of the snout, and in lateral view the skull appears relatively deep. The postorbitals
approach each other very closely behind the parietal foramen, but may be
separated by a thin parietal crest. A large, leaf-like preparietal and a clear post-
frontal are present, as well as a relatively small prefrontal. The prefrontal may
be separated from the lacrimal on the side of the snout by a narrow posterior
process of the nasal, but poor preservation does not allow confirmation of this
point.

The sharp-edged palatal rim is continued on to the anterior part of the
caniniform process without interruption, and in lateral view forms a continuous
arc. A pair of powerful maxillary tusks are present, as are two anterior pre-
maxillary ridges on the palatal surface of the premaxilla. Matrix obscures the
area where the more posterior, median ridge is normally found in dicynodonts.

The basioccipital tubera are joined by a raised area of the basicranium, the
intertuberal ridge, as seen in Lystrosaurus (Cluver 1971). The deepest part of the
ridge is on the basioccipital, behind the basioccipital-basisphenoid suture. Only
the rear of what appears to have been a wide interpterygoid vacuity is preserved.
The palatines are large, but their anterior palatal portions are partly obscured
and their relationships with the maxillae and premaxillae cannot be determined.

In the poorly preserved lower jaw, dentary tables are present and the dorsal
edge of the dentary appears to be excavated in the form of a longitudinal sulcus.

THE GENERA DICYNODON AND DIICTODON 111
pm.

3cm

Fig. 10. Dicynodon lacerticeps BMNH 36233. Type specimen. Skull in dorsal view, partially
reconstructed and with distortion corrected.

In spite of the poor condition of the type specimen, the combination of
characters which can be determined in it is sufficient to permit referral of
additional, fully preserved specimens (Figs 1-9) to Dicynodon lacerticeps and to
allow the formulation of a full generic diagnosis. The species listed below may
be retained in the genus Dicynodon on the basis of type material; an analysis
of the validity of these species is beyond the scope of this generic-level revision,
but future work will undoubtedly result in the establishment of a smaller number
of valid species.

Dicynodon testudiceps Owen
Dicynodon testudiceps Owen, 1845a: 71, pls 5-6.
Type specimen
Anterior part of skull, BMNH 47051.

ial ea ANNALS OF THE SOUTH AFRICAN MUSEUM

icc

3cm

Fig. 11. Dicynodon lacerticeps BMNH 36233. Type specimen. Partial ventral view of skull,
distortion corrected.

Locality
Fort Beaufort, Cape Province.
Horizon

Cistecephalus Zone.

Dicynodon leoniceps Owen
Dicynodon leoniceps Owen, 1876: 32, pls 24-26.
Type specimen
Skull, BMNH 47047.
Locality
Gats River, Graaff-Reinet district, Cape Province.
Horizon
Daptocephalus Zone.
Van Hoepen (1934) proposed this species as the type of a new genus,

Daptocephalus. Comparison of the type specimen with the type of Dicynodon
lacerticeps shows that large size is the only feature that can be used to distinguish

THE GENERA DICYNODON AND DIICTODON 113

Daptocephalus from Dicynodon. The species leoniceps, which may prove to be a
valid one, is accordingly reassigned to Dicynodon.

Dicynodon lissops Broom

Dicynodon lissops Broom, 1913: 450, fig. 11.
Type specimen

Skull, AMNH 5508.
Locality

Wilgerbosch, Graaff-Reinet district, Cape Province.
Horizon

Daptocephalus Zone.

Dicynodon leontops Broom

Dicynodon leontops Broom, 1913: 451, fig. 12.
Type specimen

Skull AMNH 5582.
Locality

Bethulie, Orange Free State.
Horizon

Daptocephalus Zone.

Dicynodon alticeps Broom & Haughton

Dicynodon alticeps Broom & Haughton, 1913: 37, pl. 7.
Type specimen

Skull, SAM-—2347.
Locality

Dunedin, Beaufort West district, Cape Province.
Horizon

Cistecephalus Zone.

Dicynodon watsoni Broom

Dicynodon watsoni Broom, 1921: 653, fig. 32.
Type specimen

Skull, SAM-7849.
Locality

East of New Bethesda, Graaff-Reinet district, Cape Province.
Horizon

Daptocephalus Zone.

114 ANNALS OF THE SOUTH AFRICAN MUSEUM

Dicynodon gilli Broom
Dicynodon gilli Broom, 1932: 176, fig. 60.
Type specimen
Skull, SAM-—4008.
Locality
Watervlei, Graaff-Reinet district, Cape Province.

Horizon

Cistecephalus Zone.

Dicynodon venteri Broom
Dicynodon venteri Broom, 19356: 69, fig. 10.
Type specimen
Skull, TM 199.
Locality
New Bethesda, Graaff-Reinet district, Cape Province.

Horizon

Daptocephalus Zone.

Dicynodon validus Broom
Dicynodon validus Broom, 19356: 70, fig. 11.
Type specimen
Skull, TM 252.

Locality
Leeukloof, Beaufort West district, Cape Province.

Horizon
Cistecephalus Zone.

Dicynodon microdon Broom

Dicynodon microdon Broom, 1936: 376, figs 22-23.
Type specimen

Skull, TM 267.
Locality

Bethesda Road, Graaff-Reinet district, Cape Province.
Horizon

Daptocephalus Zone.

THE GENERA DICYNODON AND DIICTODON

Dicynodon luckhoffi Broom
Dicynodon luckhoffi Broom, 1937: 306, fig. 6.
Type specimen
Skull, SAM-—K1219

Locality
Zuurplaats, Graaff-Reinet district, Cape Province.

Horizon
Daptocephalus Zone.

Dicynodon macrodon Broom
Dicynodon macrodon Broom, 1940a: 81, fig. 1OA—B.
Type specimen
Skull, RC 22.

Locality
Grootfontein, Murraysburg district, Cape Province.

Horizon
Cistecephalus Zone.

Dicynodon cadlei Brooin
Dicynodon cadlei Broom, 1940a: 82, fig. 11.
Type specimen
Skull, RC 23.

Locality
Hoeksplaas, Murraysburg district, Cape Province.

Horizon
Daptocephalus Zone.

Dicynodon calverleyi Broom
Dicynodon calverleyi Broom, 194065: 179, fig. 21.
Type specimen
skull, RC 39.

Locality

Klipfontein, Graaff-Reinet district, Cape Province.
Horizon

Cistecephalus Zone.

1M)

116 ANNALS OF THE SOUTH AFRICAN MUSEUM

Dicynodon grahami Broom
Dicynodon grahami Broom, 19406: 180, fig. 22.
Type specimen
Skull, RC 39.

Locality
Klipfontein, Graaff-Reinet district, Cape Province.

Horizon
Daptocephalus Zone.

Dicynodon trigonocephalus Broom
Dicynodon trigonocephalus Broom 1940b: 182, fig. 24.
Type specimen

Skull, RC 38.

Locality
Klipfontein, Graaff-Reinet district, Cape Province.

Horizon
Cistecephalus Zone.

Dicynodon aetorhamphus Broom
Dicynodon aetorhamphus Broom, 1948: 605, figs 23A, 24A.
Type specimen
Skull, RC 85.
Locality

Hoeksplaas, Murraysburg district, Cape Province.

Horizon

Cistecephalus Zone.

Dicynodon leontocephalus Broom
Dicynodon leontocephalus Broom, 1950: 246, fig. 1.
Type specimen
Skull, RC 96.
Locality
Springfontein (Springfield), Middelburg district, Cape Province.
Horizon

Daptocephalus Zone.

THE GENERA DICYNODON AND DIICTODON 117

Dicynodon clarencei Broom
Dicynodon clarencei Broom, 1950: 247, fig. 2.

Type specimen
Skull, RC 77.

Locality
Hanover, Cape Province.

Horizon
Cistecephalus Zone.

THE GENUS OUDENODON (Figs 12-17)
Oudenodon Owen

Diagnosis

Medium-sized to large dicynodonts (skull length ranging from 100 mm to
over 300 mm), teeth lacking in both upper and lower jaws. Postorbitals well
separated on skull roof by parietals. Septomaxilla recessed within external naris,
lacrimal may extend forward above maxilla to posterior margin of naris. Nasal
forms boss over naris. Maxilla carries weak caniniform process, with sharp-
edged posterior crest. Palatal portion of palatine large, meeting posterior border
of premaxilla. YVomers form short septum in anterior part of interpterygoid
fossa, interpterygoid vacuity long and narrow. Basipterygoid region constricted.
Ectopterygoid large, pterygoid does not contact maxilla. Dentaries carry
narrow dentary tables, dorsal edge of dentary carries deep sulcus. Coronoid
process weak or absent. Weak lateral dentary shelf above large mandibular
fenestra.

Oudenodon baini Owen

Oudenodon baini Owen, 1860: 46, pl. 1 (fig. 1).
Type specimen

Skull lacking lower jaw, BMNH 36232.
Locality

Near Fort Beaufort, Cape Province.
Horizon

Cistecephalus Zone.
Diagnosis

As for genus.
Discussion

The almost complete type skull (Figs 12-15) has been fully prepared and
detailed comparisons with other dicynodont taxa are possible. The most dis-

118 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 12. Oudenodon baini BMNH 36232. Type specimen. Skull in dorsal view.

tinctive features in the palate are the long, narrow interpterygoid vacuity, the
deep and relatively narrow secondary palate, the sharp maxillary crest behind
the caniniform process, and the absence of maxillary tusks. These characteristics,
taken together with the wide parietal exposure in the skull roof, are sufficient to
permit referral of more fully preserved specimens to Oudenodon for inclusion
of lower jaw features in the generic diagnosis.

In lateral view (Figs 15-16) the skull of Oudenodon is not as deep as that of
Dicynodon, and the anterior surface of the short snout lies almost at right angles
to the flat surface of the dorsal skull roof. The palatal rim in front of the canini-
form process is blunt, but the usual pair of anterior premaxillary ridges is

2xele:

lat. pal. f +4 oi 7H

THE GENERA DICYNODON AND DIICTODON 119

st. boc. €0¢. op

Fig. 13. Oudenodon baini BMNH 36232. Type specimen. A. Skull in ventral view. B. Occipital

view.

120 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 14. Oudenodon baini BMNH 36232. Type specimen. Stereophotograph of palate. Scale:.
8 cm.

Fig. 15. Oudenodon baini BMNH 36232. Type Specimen. Skull in lateral view.

THE GENERA DICYNODON AND DIICTODON 121

po.

orb.

Fig. 16. Oudenodon sp. SAM-—6045. Skull and jaw in lateral view.

present. In general shape and in the possession of dentary tables and a dorsal
dentary sulcus, the lower jaw is essentially similar to that of Dicynodon.

- Keyser (1975) has reviewed the species of Oudenodon and lists all type
material fully ; it is unnecessary to repeat the information here. Keyser recognizes
only three species, these being Oudenodon baini Owen, Oudenodon grandis
(Haughton), and Oudenodon luangwaensis (Boonstra) and suggests that the large
number of other dicynodont types that can be assigned to the genus are synonyms
of Oudenodon baini.

122 ANNALS OF THE SOUTH AFRICAN MUSEUM

ref.l.

3 cm

Fig. 17. Oudenodon sp. SAM-6045. Lower jaw in dorsal view.

THE GENUS DUCTODON

Diictodon Broom
Diagnosis

Medium-sized dicynodonts (average skull length 110 mm), jaws either
lacking teeth altogether, or bearing a single pair of maxillary tusks. Postorbitals
tend to cover parietals behind pineal foramen. Septomaxilla recessed within
external naris, maxilla rises high in side of snout to meet the nasal. Nasal forms
boss over external naris. Maxilla carries prominent caniniform process, clearly
demarcated from anterior maxillary palatal rim. Sharp anterior edge of canini-
form process set medially to anterior palatal rim. Palatal portion of palatine
small, does not meet premaxilla. Vomers form short septum in interpterygoid

——

THE GENERA DICYNODON AND DIICTODON 123

—— ns

3cm

Fig. 18. Diictodon galeops AMNH 5308. Type specimen. Skull in dorsal view.

fossa. Interpterygoid vacuity long. Ectopterygoid large, separating pterygoid
from maxilla. Fused dentaries carry wide dorsal dentary tables, with high medial
borders. Rear of dentary table extended medial to level of inner surface of jaw
ramus. Dorsal edge of dentary rounded behind dentary table, no coronoid
process present. Mandibular fenestra large, no expanded lateral dentary shelf
for insertion of adductor musculature.

Diictodon galeops Broom
Diictodon galeops Broom, 1913: 453, fig. 15.
Type specimen
Skull lacking lower jaw, AMNH 5308.

124 ANNALS OF THE SOUTH AFRICAN MUSEUM

Locality
‘Slachter’s Nek’, Somerset East district, Cape Province.

Horizon

Upper Permian, Cistecephalus Zone.

Diagnosis
Preparietal bone large, surrounding parietal foramen. Postfrontals absent,

caniniform processes and maxillary tusks small. Parietals partially exposed
between postorbitals behind parietal foramen.

Remarks

The type skull (Figs 18-21) is well preserved but lacks the lower jaw,
quadrates and stapes. Small tusks are present. Distinctive features of the speci-
men are the inflated preparietal bone, which surrounds the parietal opening, and
the absence of postfrontals on the skull roof. No septomaxillae can be seen, and
it is likely that these bones became disassociated from the skull, as the stapes
and quadrates evidently did, prior to fossilization.

The postorbitals approach each other behind the pineal opening but do not
cover the parietals fully. The maxilla rises high in the side of the snout and meets
the nasal so that the lacrimal is confined to the orbital border. In ventral view
it can be seen that the base of the caniniform process is offset medially to the
palatal rim in the form of an anteriorly facing blade, separated from the palatal
rim by a clear notch (Figs 20-21). Besides the usual single median and double

Fig. 19. Diictodon galeops AMNH 5308. Type specimen. Skull in lateral view.

THE GENERA DICYNODON AND DIICTODON 125

:
&

Fig. 20. Diictodon galeops AMNH 5308. Type specimen. Skull in ventral view.

anterior palatal ridges, there is a more lateral ridge on each side of the secondary
palate, medial to the caniniform process. The palatal portion of the palatine is
small, and does not meet the premaxilla. The long, wide interpterygoidal vacuity
extends far forward to where the vomers unite and descend to meet the posterior
spine of the premaxilla. The lateral pterygoidal borders of the interpterygoid
fossa are strongly constructed, but there is no pterygoid-maxilla contact.

Diictodon feliceps (Owen)
Dicynodon feliceps Owen, 1876: 45, pl. 43.

Type specimen
Skull, BMNH 47052.

126 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 21. Diictodon galeops AMNH 5308. Type specimen. Stereophotograph of palate. Scale:
5 cm.

Locality
Fort Beaufort, Cape Province.

Horizon

Cistecephalus Zone.

Diagnosis

Postorbitals cover parietals behind parietal foramen, parietal foramen not
surrounded by preparietal. Narrow postfrontal exposed on dorsal skull roof.
Caniniform process large, maxillary tusks present or absent.

Remarks

The species feliceps is the earliest described form which shows the charac-
teristics of Diictodon as diagnosed above, but examination of dicynodont type
material shows that a number of other described species of Dicynodon, including

THE GENERA DICYNODON AND DIICTODON 27

3.¢m
Fig. 22. Diictodon testudirostris SAM-—10086. Skull in dorsal view.

D. testudirostris (see Cluver 1970), closely resemble the species feliceps and can
be referred to the genus Diictodon (Figs 22-26).

An analysis of the validity of these species is outside the scope of the present
generic level revision and, as in the case of Dicynodon and Oudenodon, the list
below is compiled only on the basis of type material which can be included under
Diictodon. Future investigations may well result in a smaller number of recog-
nizable species.

Diictodon jouberti (Broom)
Dicynodon jouberti, Broom, 1905: 331.

Type specimen
Skull, SAM-695.

28 ANNALS OF THE SOUTH AFRICAN MUSEUM

3cm

Fig. 23. Diictodon testudirostris SAM-10086. A. Skull in ventral view. B. Occipital view.

THE GENERA DICYNODON AND DIICTODON 129

Locality
Gouph Tract, or Koup, Beaufort West district, Cape Province.

Horizon
Tapinocephalus Zone.

Diictodon psittacops (Broom)
Dicynodon psittacops Broom, 1912: 869, pl. 92.
Type specimen

Skull and skeleton, AMNH 5534.
Locality

Beaufort West district, Cape Province.

art.

SS SSS SSS |
3cm

Fig. 24. Diictodon testudirostris SAM-—10086. Skull and lower jaw in lateral view.

130 ANNALS OF THE SOUTH AFRICAN MUSEUM

Horizon
Cistecephalus Zone.

Diictodon ictidops (Broom)
Dicynodon ictidops Broom, 1913: 466, figs 5-6.
Type specimen
Skull, AMNH 5510.
Locality
Beaufort West commonage, Cape Province.

Horizon

Cistecephalus Zone.

Diictodon palustris (Broom)
Emydorhynchus palustris Broom, 1913: 456, fig. 19.
Type specimen
Skull, AMNH 5512.
Locality
New Bethesda, Graaff-Reinet district, Cape Province.

Horizon
Daptocephalus Zone.

Remarks

The type specimen, although poorly preserved, displays a sufficient number
of characters to warrant its referral to Diictodon, which has page priority.
Additional preparation of the type (TM 241) of Emydorhynchus formosus
Broom, 1935a (mislaid at present, E.S. Vrba 1979 pers. comm.), will be needed
to determine if this species should also be included in Diictodon.

Diictodon testudirostris (Broom & Haughton)
Dicynodon testudirostris Broom & Haughton, 1913: 36, pl. 7.
Type specimen
Skull, SAM-2354.
Locality
Dunedin, Beaufort West district, Cape Province.
Horizon

Cistecephalus Zone.

THE GENERA DICYNODON AND DIICTODON 131

add. fos.

art.

3cm

Fig. 25. Diictodon testudirostris SAM-—10086. Lower jaw in dorsal view.

Fig. 26. Diictodon testudirostris SAM-—10078. Stereophotograph of occlusal surfaces of upper
and lower jaws. Scale: 5 cm.

132 ANNALS OF THE SOUTH AFRICAN MUSEUM

Diictodon pygmaeus (Broom & Haughton)
Dicynodon pygmaeus Broom & Haughton, 1917: 123, fig. 23.
Type specimen
Skull, SAM-2664.

Locality
Dunedin, Beaufort West district, Cape Province.

Horizon
Cistecephalus Zone.

Diictodon sollasi (Broom)

Dicynodon sollasi Broom, 1921: 648, figs 28-29.
Type specimen

Skull, SAM-7420.
Locality

Biesjiespoort, Victoria West district, Cape Province.
Horizon

Cistecephalus Zone.

Diictodon macrorhynchus (Broom)
Dicynodon macrorhynchus Broom, 1921: 657, fig. 36.
Type specimen
Skull, BMNH R.4954.

Locality
New Bethesda, Graaff-Reinet district, Cape Province.
Horizon

Daptocephalus Zone.

Diictodon haughtonianus (Huene)
Dicynodon haughtonianus Huene, 1931: 186, fig. 25.
Type specimen
Skull, Institiit fir Geologie und Palaontologie, University of Tiibingen.
Locality
Blaauwkrans, Prince Albert district, Cape Province.
Horizon

Tapinocephalus Zone.

THE GENERA DICYNODON AND DIICTODON

Diictodon rubidgei (Broom)
Dicynodon rubidgei Broom, 1932: 189, fig. 62F.
Type specimen
Skull, BMNH 47081.

Locality
Graaff-Reinet district, Cape Province.

Horizon
Cistecephalus Zone.

Diictodon nanus (Broom)
Diictodon nanus, Broom, 1936: 379, fig. 25A.
Type specimen

Skull, TM 268.
Locality

Houd Constant, Graaff-Reinet district, Cape Province.

Horizon
Daptocephalus Zone.

Diictodon grimbeeki (Broom)
Dicynodon grimbeeki Broom, 1935a: 7, figs 6—7.
Type specimen
Skull, TM 253.
Locality

Leeukloof, Beaufort West district, Cape Province.

Horizon
Cistecephalus Zone.

Diictodon huenei (Broili & Schréder)

Dicynodon huenei Broili & Schroder, 1937: 118, figs 1-4.
Oudenodon huenei (Broili & Schréder) Toerien, 1953: 97.

Type specimen

Skull, University of Munich, 1934 vii 46.

Locality
La-de-da, Beaufort West district, Cape Province.

133

134 ANNALS OF THE SOUTH AFRICAN MUSEUM

Horizon

Tapinocephalus Zone.

Diictodon broomi (Broili & Schréder)
Dicynodon broomi Broili & Schroder, 1937: 132, figs 5-13.
Type specimen

Skull, University of Munich, 1934 viii 47a.

Locality
La-de-da, Beaufort West district, Cape Province.

Horizon

Tapinocephalus Zone.

Diictodon grossarthi (Broili & Schréder)
Dicynodon grossarthi Broili & Schroder, 1937: 150, figs 14-18.
Type specimen

Skull, University of Munich, 1934 viii 48.

Locality
La-de-da, Beaufort West district, Cape Province.

Horizon

Tapinocephalus Zone.

Diictodon whitsonae (Broom)

Dicynodon annae Broom, 19406: 181, fig. 23.
Dicynodon whitsonae (Broom) Toerien, 1954: 937.

Type specimen
Skull, RC 42.
Locality

Wellwood, Graaff-Reinet district, Cape Province.

Horizon

Cistecephalus Zone.

Diictodon pseudojouberti (Boonstra)
Dicynodon pseudojouberti Boonstra, 1948: 60.

Type specimen
Skull, SAM-774.

THE GENERA DICYNODON AND DIICTODON 35

Locality
Prince Albert Road, Cape Province.
Horizon

Tapinocephalus Zone.

Diictodon vanderhorsti (Toerien)
Dicynodon vanderhorsti Toerien, 1953: 91, fig. 60.
Type specimen
Skull, BPI 175.
Locality
Antjiesfontein, Prince Albert district, Cape Province.

Horizon

Tapinocephalus Zone.

Diictodon antjiesfonteinensis (Toerien)
Dicynodon antjiesfonteinensis Toerien, 1953: 93, figs 61-62.
Type specimen
Skull, BPI 219.
Locality
Antjiesfontein, Prince Albert district, Cape Province.

Horizon
Tapinocephalus Zone.

Diictodon tienshanensis (Sun)
Dicynodon tienshanensis Sun, 1973: 56.
Type specimen
Skull, in Institute of Vertebrate Palaeontology and Palaeoanthropology,
Peking.
Locality
‘Turfan Basin’, Sinkiang Province, People’s Republic of China.

Horizon
_ Lower Ko-Ko-Ya Group, Upper Permian.

Remarks

Cluver & Hotton (1979) pointed out that the specimen mentioned by Yuan
& Young (1934) and described as Dicynodon tienshanensis by Sun (1973) can be
referred to the genus Diictodon. The specimen in question is to date the only
Diictodon recorded outside South Africa.

136 ANNALS OF THE SOUTH AFRICAN MUSEUM

THE GENUS KINGORIA (Figs 27-30)
Kingoria Cox

Diagnosis

Medium-sized dicynodonts (average skull length 160 mm), jaws lacking teeth
altogether or bearing a single pair of maxillary tusks. Parietals exposed between
postorbitals behind parietal foramen. Septomaxilla recessed within opening
of naris, maxilla rises high in side of snout to meet nasal. Low boss formed by
nasal. Maxilla carries prominent caniniform process, palatal rim continued with-
out interruption on to anterior blade of caniniform process. Rear of caniniform
process extended as keel to level of ectopterygoid. Palatal portion of palatine very
small, but making contact with greatly expanded premaxilla. Vomers form short
septum in interpterygoid fossa, interpterygoid vacuity long and narrow. Ecto-
pterygoid large, separating pterygoid from maxilla. Basipterygoid region con-
stricted. Fused dentaries taper to form rounded anterior tip of lower jaw, no
dentary tables present. Dorsal edge of dentary narrow, lateral dentary shelf widely
expanded. Coronoid process weak or absent. Mandibular fenestra reduced or
absent. Angular forms sharp ventral keel behind reflected lamina.

Kingoria nowacki (Huene)

Dicynodon nowacki von Huene, 1942: 156, fig. 2.
Kingoria nowacki (von Huene) Cox, 1959: 321.
Type specimen

Skull, Institiit fir Geologie und Paldontologie, University of Tubingen,
K-12.
Locality

Kingori, Tanzania.

Horizon

Kawinga Formation (Charig 1963).
Diagnosis
As for genus.

Discussion

A diagnosis of Kingoria, which has been fully described and characterized
by Cox (1959), is provided only in the interest of completeness. Cox suggested
that Dicynodon galecephalus Broom was related to or congeneric with Kingoria,
but detailed examination of dicynodont type material shows that this is only
one of a number of species previously included under Dicynodon which may be
assigned to Kingoria. A potential problem of priority and nomenclature exists
in that, on the basis of the poorly preserved type, Dicynodontoides parringtoni
Broom, 19406, appears to be related to or congeneric with Kingoria. In the

THE GENERA DICYNODON AND DIICTODON 137

Ey
3cm

Fig. 27. Kingoria nowacki. Skull in dorsal view (after Cox 1959).

interests of stability, it is proposed that at this stage Dicynodontoides be retained
as a separate genus, distinct from but related to Kingoria, until preparation of
the type specimen (RC 45) allows a full comparison with the type of Kingoria
nowacki to be made.

Kombuisia frerensis Hotton, 1974, from the lower Triassic Cynognathus
zone, is the only Triassic form so far described that can be related to the upper
Permian Kingoria.

Kingoria recurvidens (Owen)
Dicynodon recurvidens Owen, 1876: 46, pl. 69.

Type specimen
Skull, BMNH 40709.

138 ANNALS OF THE SOUTH AFRICAN MUSEUM

3cm

Fig. 28. Kingoria nowacki. Skull and lower jaw in lateral view (after Cox 1959).

Locality
Fort Beaufort, Cape Province.

Horizon

Cistecephalus Zone.

Kingoria gracilis (Broom)
Oudenodon gracilis Broom, 1901: 162.
Type specimen
Skull, SAM-590.
Locality
Pearston, Cape Province.
Horizon

Cistecephalus Zone.

THE GENERA DICYNODON AND DIICTODON

pm.

Fig. 29. Kingoria nowacki. Skull in ventral view (after Cox 1959).

Kingoria grahami (Broom)
Dicynodon grahami Broom, 19406: 180, fig. 22.

Type specimen
Skull, RC 40.

Locality

St Olives, Graaff-Reinet district, Cape Province.
Horizon

Daptocephalus Zone

139

140 ANNALS OF THE SOUTH AFRICAN MUSEUM

Kingoria howardi (Broom)
Dicynodon howardi Broom, 1948: 604, fig. 22A.
Type specimen

Skull, RC 83.

Locality
Riversdale, Graaff-Reinet district, Cape Province.

Horizon
Cistecephalus Zone.

Kingoria duvenhagei (Broom)
Dicynodon duvenhagei Broom, 1948: 607, fig. 25.
Type specimen

Skull, RC 64.

Locality
Doornkloof, Graaff-Reinet district, Cape Province.

Horizon
Daptocephalus Zone.

Zam art.

Fig. 30. Kingoria nowacki. Lower jaw in dorsal view (after Cox 1959).

THE GENERA DICYNODON AND DIICTODON 141

Kingoria galecephala (Broom & Robinson)

Dicynodon galecephalus Broom & Robinson, 1948: 404.
Kingoria galecephala (Broom & Robinson) Cox, 1959: 324.

Type specimen
Skull, RC 97.

Locality
Ferndale, Graaff-Reinet district, Cape Province.

Horizon
Cistecephalus Zone.

INTERRELATIONSHIPS OF DICYNODON, OUDENODON,
DITCTODON, AND KINGORIA

Recognition of four central Permian dicynodont genera on the basis of the
diagnoses given above has brought a clear picture of dicynodont interrelation-
ships a step closer. While a full review of dicynodonts based on diagnoses of all
genera cannot be attempted here, comparisons between Dicynodon, Oudenodon,
Diictodon, and Kingoria nevertheless suggest closer relationships among some
genera than among others (see Table 1).

In terms of lower jaw and palatal morphology, Dicynodon and Oudenodon
resemble each other closely. It is unlikely that these similarities, involving
dentary tables, a dorsal dentary sulcus, a deeply vaulted secondary palate with
high palatal rim, and a large palatal development of the palatine, would have
arisen as a consequence of convergence, and the differences which are seen in the
palatal structure of the two genera are very likely the result of permanent loss of
the maxillary tusk in Oudenodon and accompanying modification of masticatory
function.

Diictodon resembles Dicynodon and Oudenodon in the presence of dentary
tables, but the structure of the dorsal edge of the dentary as well as the palatal
rim, caniniform process, and palatine sets the genus well apart from the others.

On the basis of palatal and lower jaw structure, Kingoria holds only a
remote relationship with the other genera. The lower jaw, with flared lateral
dentary shelves, blunt anterior tip, and highly reduced mandibular fenestra, is
unique and, taken in conjunction with palatal structure, suggests an origin of the
genus well separated from that of Dicynodon, Oudenodon or Diictodon.

Of the four genera discussed above, Diictodon has the greatest stratigraphic
range. Specimens that can be referred to the genus are common in collections
from the Tapinocephalus Zone, and are extremely abundant in certain Ciste-
cephalus Zone localities. In addition, a specimen of Diictodon has been identified
from the Upper Permian of Tienshan, China (Cluver & Hotton, 1979). The
genus appears to persist to the very top of the Daptocephalus Zone, but has not
been recorded from the Triassic Karoo formations. Dicynodon, Oudenodon, and

142 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 1
Distribution of character states

Dicynodon Oudenodon Diictodon Kingoria

postcaniniform crest . 5 : ; x

palatal rim continuous. By oe Xx < x
palatal rim notched . : ; : mK

broad inter-temporal region . . x x
narrow inter-temporal region . ; Xx x

tusks present . : : ; : x

tusks absent AURA ee x

tusks absent or present . : ; x Xs
dorsal sulcus in dentary . : ; x S<

dentary tables. ; . ‘ : x x x

dentary tables absent. : : : an
weak dentary shelf . ira ; x x <

wide dentary.shelf .. +...) % ; x
septomaxilla recessed : : : XK x x
septomaxilla exposed. , ; é x

large palatine. : : : : x x

small palatine . : : : E x x

Kingoria have not been identified from Tapinocephalus Zone localities, but all
three are relatively common in the Daptocephalus Zone. Oudenodon appears
to be better represented in Cistecephalus Zone localities than are Dicynodon or
Kingoria, but whereas Oudenodon did not survive the Permian—Triassic
transition, both Kingoria and Dicynodon can claim relationships with Triassic
genera. Kombuisia frerensis (Hotton, 1974) is clearly a specialized relative of
Kingoria, while the features that set Dicynodon apart from the other Permian
genera are seen in modified form in Lystrosaurus and Kannemeyeria (Cluver
1971), as well as in many of the later, non-South African genera for which good
descriptions exist (see Keyser & Cruickshank 1979). Future revision of the group
of which Dicynodon is an early and primitive member will very likely show it to
be geographically and stratigraphically the most wide-ranging dicynodont taxon.

Further speculation on the relationships of Dicynodon, Oudenodon, Diicto-
don and Kingoria would be fruitless before the status of other Permian dicyno-
donts has been clarified, in particular the pristerodontid genera and the primitive

THE GENERA DICYNODON AND DIICTODON 143

forms from the lowermost Tapinocephalus zone of the Beaufort series. However,
recognition of the distinctive morphological features characterizing the four
genera as set out above should aid materially in revealing what will undoubtedly
prove to be a highly complex phylogeny of dicynodonts.

ACKNOWLEDGEMENTS

M. A. Cluver is indebted to the South African Council for Scientific and
Industrial Research and the Trustees of the South African Museum for travel
grants enabling him to study the collections of the British Museum (Natural
History), London; American Museum of Natural History, New York; and the
Smithsonian Institution, Washington, D.C.

N. Hotton is indebted to National Science Foundation Grants Nos G14707
and GB1647, and Smithsonian Research Foundation Grants Nos 3319 and
413615, enabling him to visit and undertake fieldwork in South Africa. He
wishes especially to acknowledge the generous help and encouragement of
Dr J. W. Kitching, of the Bernard Price Institute for Palaeontological Research,
University of the Witwatersrand, and also the cooperation of Dr A. S. Brink,
formerly of that institution.

Dr A. J. Charig, British Museum (Natural History), London, and Dr E. S.
Gaffney, American Museum of Natural History, New York, are thanked for the
loan of important type material in their care. The stereophotographs of Oudeno-
don baini (Fig. 14) and Diictodon galeops (Fig. 21) were taken by Messrs T.
Parminter (British Museum (Natural History), London) and C. Tarka (American
Museum of Natural History, New York) respectively. Other stereophotographs
were taken by Mr N. J. Eden (South African Museum). Preparation of the
South African Museum material illustrated in the text was largely the work of
Mrs I. M. Chesselet, then of the Museum’s Department of Palaeontology.

REFERENCES

AGNEW, J. D. 1959. Cranio-osteological studies in Dicynodon grimbeeki with special reference
to the sphenethmoid region and cranial kinesis. Palaeont. afr. 6: 77-107.

BoonstrA, L. D. 1948. On the anomodont reptiles from the Tapinocephalus zone of the Karroo
System. In: ROYAL SociETY OF SOUTH AFRICA. Robert Broom Commemorative volume:
57-64. Cape Town: Royal Society of South Africa.

Broiwi, F. & SCHRODER, J. 1937. Beobachtungen an Wirbeltieren der Karrooformation.
XXVIII. Uber einige neue Anomodontier aus der Tapinocephalus Zone. S. B. bayer.
Akad. Wiss. 1937: 118-168.

Broom, R. 1901. On the structure and affinities of Udenodon. Proc. zool. Soc. Lond. 1901:
162-190.

Broom, R. 1905. Notice of some new fossil reptiles from the Karroo Beds of South Africa.
Rec. Albany Mus. 1: 331-337.

Broom, R. 1912. On some new fossil reptiles from the Permian and Triassic beds of South
Africa. Proc. zool. Soc. Lond. 1912: 859-876.

Broom, R. 1913. On some new genera and species of dicynodont reptiles, with notes on a few
others. Bull. Am. Mus. nat. Hist. 32: 441-457.

Broom, R. 1921. On some new genera and species of anomodont reptiles from the Karroo
beds of South Africa. Proc. zool. Soc. Lond. 1921: 647-674.

144 ANNALS OF THE SOUTH AFRICAN MUSEUM

Broom, R. 1932. The mammal-like reptiles of South Africa. London: Witherby.

Broom, R. 1935a. A new genus and some new species of mammal-like reptiles. Ann. Transy.
Mus. 18: 1-12.

Broom, R. 19355. On some new genera and species of Karroo fossil reptiles Ann. Transy. Mus.
18: 55-72.

Broom, R. 1936. On some new genera and species of Karroo fossil reptiles with notes on some
others. Ann. Transy. Mus. 18: 349-386.

Broom, R. 1937. A further contribution to our knowledge of the fossil reptiles of the Karroo.
Proc. zool. Soc. Lond. 1937: 299-318.

Broom, R. 1940a. Some new Karroo reptiles from the Graaff-Reinet District. Ann. Transy.
Mus. 20: 71-87.

Broom, R. 1940b. On some new genera and species of fossil reptiles from the Karroo Beds of
Graaff-Reinet. Ann. Transy. Mus. 20: 157-192.

Broom, R. 1948. A contribution to our knowledge of the vertebrates of the Karroo Beds of
South Africa. Trans. R. Soc. Edinb. 61: 577-929.

Broom, R. 1950. Three new species of anomodonts from the Rubidge collection. Ann. Transy.
Mus. 21: 246-250.

Broom, R. & HAUGHTON, S. H. 1913. On the two new species of Dicynodon. Ann. S. Afr. Mus.
12: 36-39.

Broom, R. & HAUGHTON, S. H. 1917. Some new species of Anomodontia (Reptilia). Ann. S.
Afr. Mus. 12: 119-125.

Broom, R. & ROBINSON, J. T. 1948. Some new fossil reptiles from the Karroo Beds of South
Africa. Proc. zool. Soc. Lond. 118: 392-407.

Camp, C. L. & WELLES, S. P. 1956. Triassic dicynodont reptiles. Part I. The North American
genus Placerias. Mem. Univ. Calif. 13: 255-304.

CHARIG, A. J. 1963. Stratigraphical nomenclature in the Songea Series of Tanganyika. Rec.
geol. Surv. Tanganyika 10: 47-53.

CLuverR, M. A. 1970. The palate and mandible in some specimens of Dicynodon testudirostris
Broom & Haughton (Reptilia, Therapsida). Ann. S. Afr. Mus. 56: 133-153.

CLuver, M. A. 1971. The cranial morphology of the dicynodont genus Lystrosaurus. Ann. S.
Afr. Mus. 56: 155-274.

CLuver, M. A. 1974a. The skull and mandible of a new cistecephalid dicynodont. Ann. S. Afr.
Mus. 64: 137-155.

CLuver, M. A. 19746. The cranial morphology of the Lower Triassic dicynodont Myosaurus
gracilis. Ann. S. Afr. Mus. 66: 35-54.

CLuver, M. A. 1975. A new dicynodont from the Tapinocephalus Zone (Karoo System, Beau-
fort Series) of South Africa, with evidence of the jaw adductor musculature. Ann. S. Afr.
Mus. 67: 7-23.

CLuver, M. A. & Hotton, N. 1979. The dicynodont genus Diictodon (Reptilia, Therapsida)
and its significance. Proc. IV International Gondwana Symposium, 1977, Calcutta: 176-183.

Cox, C. B. 1959. On the anatomy of a new dicynodont genus, with evidence of the position of
the tympanum. Proc. zool. Soc. Lond. 132: 321-367.

CromPTon, A. W. & HoTTON, N. 1967. Functional morphology of the masticatory apparatus
of two dicynodonts (Reptilia, Therapsida). Postilla 109: 1-51.

CRUICKSHANK, A. R. I. 1968. A comparison of the palates of Permian and Triassic dicynodonts.
Palaeont. afr. 11: 23-31.

HAUGHTON, S. H. & Brink, A. S. 1954. A bibliographic list of Reptilia from the Karroo beds
of Africa. Palaeont. afr. 2: 1-187.

Horton, N. 1974. A new dicynodont (Reptilia, Therapsida) from Cynognathus zone deposits
of South Africa. Ann. S. Afr. Mus. 64: 157-165.

HUuENE, F. Von. 1931. Beitrag zur Kenntnis der Fauna der siidafrikanischen Karrooformation.
Geol. palaeont. Abh. (N.F.) 18: 158-228.

HUENE, F. Von. 1942. Die Anomodontier des Ruhuhu-Gebietes in der Tuibinger Sammlung.
Palaeontographica 94: 154-184.

Keyser, A. W. 1975. A re-evaluation of the cranial morphology and systematics of some
tuskless Anomodontia. Mem. geol. Surv. Rep. S. Afr. 67: 1-110.

Keyser, A. W. & CRUICKSHANK, A. R. I. 1979. The origins and classification of Triassic
dicynodonts. Trans. geol. Soc. S. Afr. 82: 81-108.

THE GENERA DICYNODON AND DIICTODON 145

Ewer, R. F. 1961. The anatomy of the anomodont Daptocephalus leoniceps (Owen). Proc.
zool. Soc. Lond. 136: 375-402.

KITCHING, J. W. 1970. A short review of the Beaufort zoning in South Africa. In: JUGS 2nd
Symposium Gondwana Stratigraphy and Palaeontology: 309-312. Cape Town, Johannes-
burg: CSIR.

KitcuINnG, J. W. 1977. The distribution of the Karoo vertebrate fauna. Mem. Bernard Price
Inst. palaeont. Res. 1: 1-131.

Oxson, E. C. 1944. The origin of mammals based on the cranial morphology of the therapsid
suborders. Spec. Pap. geol. Soc. Am. 55: 1-136.

OwEN, R. 1845a. Report on the reptilian fossils of South Africa. Part 1. Description of certain
fossil crania, discovered by A. G. Bain in sandstone rocks at the south eastern extremity
of Africa, referable to an extinct genus of Reptilia (Dicynodon), and indicative of a new
tribe or sub-order of Sauria. Trans. geol. Soc. Lond. 7: 59-84.

OwEN, R. 18455. Description of certain fossil crania, discovered by A. G. Bain, Esq., in sand-
stone rocks at the south-eastern extremity of Africa, referable to different species of an
extinct genus of Reptilia (Dicynodon), and indicative of a new tribe or suborder of Sauria.
Proc. geol. Soc. 4: 500-504.

OwEN, R. 1856. Report on the reptilian fossils of South Africa. Part II. Description of the
skull of a large species of Dicynodon (D. tigriceps, Ow.), transmitted from South Africa
by A. G. Bain, Esq. Trans. geol. Soc. Lond. 7: 233-240.

OweEN, R. 1860. On some reptilian fossils from South Africa. Q. J/. geol. Soc. Lond. 16: 49-63.

OwEN, R. 1876. Descriptive and illustrated catalogue of the fossil Reptilia of South Africa in
the collection of the British Museum. London: British Museum (Natural History).

Sun, A. L. 1973. A new species of Dicynodon from Sinkiang: Vert. Palasiatica 11: 52-58.

TOERIEN, M. J. 1953. The evolution of the palate in South African Anomodontia and its
classificatory significance. Palaeont. afr. 1: 49-117.

TOERIEN, M. J. 1954. Lystrosaurus primitivus, sp. nov. and the origin of the genus Lystrosaurus.
Ann. Mag. nat. Hist. (12) 7: 934-938.

VAN HoeEPEN, E. C. N. 1934. Oor die indeling van die Dicynodontidae na aanleiding van nuwe
vorme. Palaeont. Navors. nas. Mus. Bloemfontein 2: 67-101.

YuAn, P. L. & Youna, C. C. 1934. On the discovery of a new Dicynodon in Sinkiang. Bull.
geol. Soc. China 13: 563-573.

ABBREVIATIONS
add. fos. adductor fossa
ang. angular
art. articular
bas. basisphenoid
boc. basioccipital
bo.t. basioccipital tuber
den. dentary
den. s. dentary shelf
den. t. dentary table
ect. ectopterygoid
eoc. exoccipital
ept. epipterygoid
for. mag. foramen magnum
fr. frontal
icc. canal for internal carotid artery
ip. interparietal
ipt. vac. interpterygoid vacuity
itr. intertuberal ridge
jug. jugal
lab. fos. labial fossa
lac. lacrimal

lat. pal. f.

lateral palatal fenestra

146

ANNALS OF THE SOUTH AFRICAN MUSEUM

lateral shelf

maxilla

notch

opisthotic
orbitosphenoid
parietal

palatine
parasphenoid
postcaniniform crest
prefrontal
premaxilla
postorbital
postfrontal
preparietal

prootic

presphenoid
pterygoid
posttemporal fenestra
quadrate
quadratojugal
reflected lamina
surangular
septomaxilla
supraoccipital
squamosal

stapes

tusk

tabular

vomer

foramen for facial nerve

American Museum of Natural History, New York
British Museum (Natural History), London
Bernard Price Institute for Palaeontological

Research, Johannesburg

Rubidge Collection, Wellwood, Graaff-Reinet
South African Museum, Cape Town

Transvaal Museum, Pretoria

6. SYSTEMATIC papers must conform to the International code of zoological nomenclature
(particularly Articles 22 and 51).

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“not acceptable.

Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.

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MICHAEL A. CLUVER
&
NICHOLAS HOTTON III

THE GENERA DICYNODON AND DIICTODON
AND THEIR BEARING ON THE CLASSIFICATION
OF THE DICYNODONTIA (REPTILIA, THERAPSIDA)

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FiscHER, P.-H., DuvAL, 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.
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.
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ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 83 Band
August 1981 Augustus
Part 7 Deel

M
a S
Alig, >
vio nov M8

REVISION OF THE LATE VALANGINIAN
CEPHALOPODA FROM THE SUNDAYS RIVER
FORMATION OF SOUTH AFRICA, WITH
SPECIAL REFERENCE TO THE GENUS
OLCOSTEPHANUS

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Printed in South Africa by In Suid-Afrika gedruk deur
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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

REVISION OF THE LATE VALANGINIAN CEPHALOPODA
FROM THE SUNDAYS RIVER FORMATION OF SOUTH AFRICA,
WITH SPECIAL REFERENCE TO THE GENUS OLCOSTEPHANUS

MICHAEL R. COOPER

Queen Victoria Museum, Salisbury*

(With 205 figures)
[MS. accepted 8 August 1980]

ABSTRACT

The cephalopod fauna from the Sundays River Formation is revised and shown to com-
prise 14 species and varieties of Olcostephanus, 2 species each of Distoloceras, Bochianites
and Belemnopsis, and 1 species each of Neohoploceras, Eodesmoceras, Partschiceras, and
Eutrephoceras. Sexual dimorphism is recognized within Olcostephanus and the implications
and importance of this phenomenon are discussed at length. The fauna is considered of
latest Valanginian (O. baini Zone) age. The new genus Jeanthieuloyites is proposed for Roger-
sites quinquestriatus Besairie, and one new species of Olcostephanus, O. riccardii sp. nov.,
is described.

CONTENTS

PAGE
Introduction ; : : : ; ; eee 7h
Age of the Sundays River Formation 3 ; ‘ : ; ook Ail
Systematics . r : f : : : : é : 7 > 54
Genus Bee eiceran ; : : : : : f ; =e SASS
Genus Bochianites . ; ; g : 3 : 5 : - e156
Genus Olcostephanus . : : : ‘ : : 2) 6
Sexual dimorphism in ammonites : : ; : : eli
Sexual dimorphism in Olcostephanus . : : : ne Seely
The peristome in Olcostephanus . : c ‘ J Se We
Homoeomorphy in Olcostephanus . : 181
Description of the Sundays River species of Giconenkaaas 182
Genus Neohoploceras . : ‘ : : : ‘ : . 344
Genus Distoloceras : : : : : ; : j =) 346
Genus Eodesmoceras . i ‘ : , : ; ‘ Rees 53)
Genus Belemnopsis : : : ; ; ; : P s #355
Genus Eutrephoceras . : : : ; A ; 2 z- Soy
Summary . : : , : : é : : ; : <1), 358
Acknowledgements . : ‘ : ; ‘ ; ‘ : 5) 9360
References . E - ; ; : 3 ; : : ; = ool

INTRODUCTION

In the southern and south-eastern Cape, late Mesozoic sediments occur as
a widely scattered series of basins, representing basin infills in uneven terrain.
The bulk of these deposits are of non-marine origin, and it is only in the
Uitenhage basin (Fig. 1), where these sediments attain their fullest development,

* Present address: National Museum, Bulawayo

147

Ann. S. Afr. Mus, 83 (7), 1981: 147-366, 205 figs.

148 ANNALS OF THE SOUTH AFRICAN MUSEUM

SE tt A

Sundays River

ALGOA BAY

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PORT ELIZABETH

. Swartkops River

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Fig. 1. The geology of the Uitenhage Basin (after Geological Survey Map of 1962).

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KIRKWOOD FORMATION
Collecting sites

Faults

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REVISION OF LATE VALANGINIAN CEPHALOPODA 149

that the succession is complete. It was to the sediments of this area that Tate
(1867) first applied the name ‘Uitenhage Beds’.

Within the sediments of the Uitenhage Group (Winter 1973), a tripartite
subdivision has long been recognized which, using current lithostratigraphic
nomenclature, is:

3. Sundays River Formation
2. Kirkwood Formation
1. Enon Formation

With the break-up Gondwanaland, the depositional history of the area
was initially one of rapid deposition of fluviatile sandstones and torrential
conglomerates of the Enon Formation, followed by the more quiescent,
lagoonal to marginally marine environments of the Kirkwood Formation,
finally terminating in the transgressive marine deposits of the Sundays River
Formation. It is the rich invertebrate faunas of the latter unit that have provided
the main basis for the dating of the Uitenhage Group.

Fossils were first recorded from the Uitenhage Group by Hausmann (1837)
who described some molluscs from the Sundays River valley, including the
ammonites Ammonites spinosissimum and a ‘Hamites’ sp., the latter being
erroneously compared with the late Albian species H. intermedius J. Sowerby
and H. funatus Brongniart. He considered the fauna to be of Lower Cretaceous
age, a determination supported by Goldfuss (1837, 1840) who figured two of
Hausmann’s shells, whilst Krauss (1843, 1850) assigned a Neocomian age to
a small collection of bivalves from the Swartkops River.

In 1851 Portlock (1852) exhibited a collection of fossil plants and molluscs,
collected by Rubidge from the Sundays River valley, to which he assigned a
probable Jurassic age.

Bain (1856) doubtfully referred the Uitenhage fossils to the Lias, based on
the abundance of the supposedly Liassic form, ‘Gryphaea incurva’, actually a
misidentification of Aetostreon imbricatum (Krauss). In an appendix to Bain’s
paper, Sharpe described a new collection of fossils from the Sundays and
Swartkops Rivers, including the ammonites Ammonites atherstoni and A. baini.
An erroneous comparison of these two species with ‘Lower Oolite’ (Bajocian)
forms led Sharpe to conclude that the fauna was most closely related to those
from the ‘Lower and Middle Oolite’ (Bajocian—Bathonian) of Europe.

In 1857 Atherstone suggested a partly Jurassic and partly Cretaceous age
for the Uitenhage Group, although Wyley (1859) placed the Enon Formation
as low as the New Red Sandstone (Triassic) and correlated the Sundays River
Formation with the ‘Oolites’ (Middle Jurassic).

In his description of several new molluscs from the Uitenhage Group,
Tate (1867) considered the deposit to represent a condensed sequence spanning
the entire Jurassic period, with the possible exception of the ‘Upper Oolite’.
Amongst the new forms described were Ammonites subanceps, compared with
A. anceps Reinecke from the ‘Middle Oolite’ (a Callovian Reineckia), Hamites

150 ANNALS OF THE SOUTH AFRICAN MUSEUM

africanus, and the belemnite Belemnites africanus which was compared with the
Jurassic B. aucklandicus.

Stoliczka (1871), in his monograph of the Cretaceous Bivalvia of southern
India, suggested that some of the Uitenhage forms showed Cretaceous rather
than Jurassic affinities.

The bivalve Crassatella complicata Tate was assigned by Dames (1873) to
the genus Ptychomya and considered to indicate a Neocomian age, a suggestion
supported by a fragment of ammonite which Dames believed to be identical
with Ammonites astieri d’Orbigny. ; 4

Lycett (1879), in the concluding chapter of his monograph of the British
fossil trigoniids, expressed the opinion that some of the more characteristic of
the Uitenhage forms pointed decisively to a Cretaceous age.

In a critical review of Tate’s work, Neumayr (in Holub & Neumayr, 1882),
besides describing several new bivalves and figuring the holotype of Crioceras
spinosissimum (Hausmann) for the first time, dealt with the age of the Sundays
River faunas at some length. He considered Olcostephanus atherstoni (Sharpe),
O. baini (Sharpe), ‘Crioceras’ spinosissimum (Hausmann), “Trigonia’ ventricosa
Krauss, ‘T’. conocardiiformis Krauss, Crassatella complicata Tate, and ‘Exogyra@’
imbricata Krauss to be essentially Cretaceous in character. He also considered
Tate’s Ammonites subanceps to possibly represent merely the juvenile whorls
of Crioceras spinosissimum.

Griesbach (1880), Jones (1884), Moulle (1885), Futterer (1897), and
Molengraaf (1900) all assigned a Jurassic age to the Uitenhage Group, while
Giirich (1887), Schenk (1888), Molengraaf (1890), and Lemoine (1906), on the
other hand, suggested a Lower Cretaceous age.

Pavlow (in Pavlow & Lamplugh 1892) assigned both Ammonites atherstoni
and A. baini to the Neocomian subgenus Holcostephanus (Astieria), thereby
suggesting a Lower Cretaceous age for the Sundays River Formation. More-
over, he considered H. atherstoni (Sharpe) to be identical to H. psilostomus
(Neumayr & Uhlig) from northern Germany.

Newton (1896) published a complete list of the known Mollusca from the
Uitenhage Group and assigned them a Neocomian age, while Passarge (1904)
suggested an Upper Jurassic-Lower Cretaceous age.

Kitchin (1908), in his exhaustive study, gave a concise treatment of the
earlier literature and dealt with the age of the fauna at great length. He con-
sidered the Cephalopoda to carry the greatest weight in the assignment of an
age to the deposit, noting that ‘... the known species of Holcostephanus (sensu
stricto) are almost wholly, if not entirely, confined to strata of Upper Valan-
ginian and Lower Hauterivian age’ (p. 30). Having shown that Tate’s compari-
son of many of the Bivalvia with Jurassic forms was erroneous, Kitchin
(1908: 39) was led to conclude ‘...that no portion of the Uitenhage Series
represents a period of time earlier or later than the Neocomian. It must be said,
indeed, that the almost entire restriction of Holcostephanus sensu stricto
(= Astieria Auctorum), to the upper part of the Valanginian and the lower

REVISION OF LATE VALANGINIAN CEPHALOPODA 151

beds of the Hauterivian in Europe suggests much narrower limits, when we
consider how important a place is taken by members of this genus in character-
izing the cephalopod-fauna of the Uitenhage beds.’ Amongst the ammonites,
Kitchin (1908) described six new species—Bochianites glaber, Phylloceras
rogersi, Holcostephanus wilmanae, H. modderensis, H. rogersi, and H. uiten-
hagensis, as well as referring to a Belemnites sp. from Coega.

Wegener (1909), in a review of the subgenus Holcostephanus (Astieria) into
which he included all the Uitenhage forms, accepted the majority of these as
valid, with the exception of H. wilmanae which he considered to represent a
variety of H. (Astieria) psilostomus Neumayr & Uhlig.

In 1924 Spath erected the new genus Rogersites, type species R. modderensis
(Kitchin), to include all the Olcostephaninae from the Uitenhage Group with-
out, however, providing a formal diagnosis.

In a revision of the Uitenhage Cephalopoda in 1930, Spath had still not
formulated a generic diagnosis for Rogersites, but included into it all previously
described olcostephanids from the Uitenhage Group, as well as describing the
new species Rogersites kitchini, R. crassicostatus, R. sphaeroidalis, R. otoitoides,
Eodesmoceras haughtoni and the nautiloid Eutrephoceras uitenhagense.

AGE OF THE SUNDAYS RIVER FORMATION

Kitchin’s (1908) detailed study of the faunas from the Sundays River
Formation dispelled any doubt as to the presence of Jurassic forms, while
concluding that ‘.. . the Marine Beds represent not more than the strata at the
top of the Valanginian and the base of the Hauterivian’.

Besairie (1936) and Collignon (1962) have recorded rich olcostephanid
faunas from Madagascar which include Olcostephanus atherstoni (Sharpe),
O. baini (Sharpe), O. uhligi (Collignon) (= O. fascigerus Spath), and Distoloceras
spinosissimum (Hausmann) from the Valanginian of that island. Partschiceras
rogersi (Kitchin), the type of which was collected midway up the cliffs behind
Colchester, and thus high in the marine succession, is reported by Collignon
(1962) only from the Lower Valanginian, although Haughton (1963) mentions
its occurrence in strata of early Hauterivian age from the same island. Like
most phylloceratids, therefore, it is presumably a long-ranging species of little
use for detailed biostratigraphic correlation.

Bose (1923) and Imlay (1937, 1938, 1940, 1960) have recorded rich
olcostephanid faunas from the Upper Valanginian and Lower Hauterivian of
Mexico. Imlay (1937, 1938) considered the upper part of the Taraises Formation
to be of early Hauterivian age. His faunal list from these beds included
- Olcostephanus, Maderia, Mexicanoceras, Acanthodiscus, Leopoldia, Neocomites,
Distoloceras, Valanginites, Thurmannites (= Thurmanniceras), Bochianites,
Kilianella, and Dichotomites. The pyritic olcostephanid nuclei Maderia and
Mexicanoceras are endemic and of no value in biostratigraphic correlation.
According to Wright (in Arkell et al. 1957), Neocomites, Thurmanniceras,

152 ANNALS OF THE SOUTH AFRICAN MUSEUM

Valanginites, and possibly Kilianella are all restricted to Valanginian and older
strata. A somewhat older age is also suggested for the lower portion of the
Upper Member of the Taraises Formation by the faunal association Olco-
stephanus—Bochianites—Distoloceras, which characterizes the Upper Valanginian
of Speeton (England), Verdon (France) and Madagascar, as well as being
typical of the Uitenhage Formation. This is supported by the presence of
Kilianella mayranensis Imlay which closely resembles K. roubaudiana (d’ Orbigny),
the differences hardly warranting specific separation. It seems likely, therefore,
that Imlay’s fauna should be considered as having come from more than one
palaeontological zone, including Upper Valanginian strata.

A re-examination of the Valanginian stratotype led Barbier & Thieuloy
(1963) to subdivide the Valanginian into a lower Kilianella roubaudiana Zone
and an upper Saynoceras verrucosum Zone. The so-called ‘Astieriaschicht’, rich
in Olcostephanus atherstoni (Sharpe) (Barbier & Thieuloy 1963: 82), is placed at
the top of the Valanginian, pending detailed reinvestigation. In 1967, however,
Moullade & Thieuloy wrote ‘. . . il subsiste cependant toujours un large hiatus
non caractérisé par les Ammonites entre le Valanginien ‘moyen’ a Saynoceras
verrucosum et les termes ultimes du Valanginien supérieur’. The terminal
Valanginian was subdivided into a lower zone of Himantoceras trinodosum
(Thieuloy) and an upper zone of Sarasinella ambigua (Uhlig), while the base
of the Hauterivian was marked by the appearance of Acanthodiscus radiatus
(Brugiere).

Busnardo & Cotillon (1964) recorded a rich olcostephanid fauna from
Bas—Verdon in France, from calcareous marls at the very top of the Valanginian,
overlain by so-called Valanginian—Hauterivian passage beds, in turn succeeded
by Lower Hauterivian beds with Acanthodiscus radiatus, Olcostephanus filosus
(Baumberger) and O. aff. psilostomus (Neumayr & Uhlig). The Upper Valan-
ginian fauna included O. atherstoni (Sharpe).

In Argentina, strata with O. atherstoni (Sharpe) are apparently overlain
(Riccardi et al. 1971) by a Lower Hauterivian faunal assemblage with
Acanthodiscus cf. radiatus (Brugiére). Riccardi et al. (1971) believed their fauna
to show close agreement with the “Astieriaschicht’ of the Swiss Jura, but pre-
ferred tentatively to date the Argentinian Olcostephanus atherstoni assemblage
at ‘... late Valanginian to earliest Hauterivian, pending detailed biostratigraphic
revision on a world-wide scale’.

At Speeton (Spath 1924) there is an apparent hiatus, possibly only local,
during the Upper Valanginian—basal Hauterivian, and derived fossils of this
age occur in the black nodules of Division D,. Spath (1924: 86) considered
‘... the main development of Olcostephanus, of which A. astieri is the genotype,
is at the base of the Hauterivian and uppermost Valanginian’.

The top bed of the Middle Member of the Chichali Formation in the Trans
Indus Range of northern Pakistan has yielded a rich Olcostephanus fauna
(Spath 1939; Fatmi 1977) which leaves little doubt as to its contemporaneity
with the Sundays River faunas. Species in common include O. fascigerus

REVISION OF LATE VALANGINIAN CEPHALOPODA 153

Spath, O. baini baini (Tate), O. baini var. sphaeroidalis (Spath), O. rogersi
(Spath), and perhaps O. densicostatus (Wegner) and O. perinflatus (Matheron).

Imlay & Jones (1970) recorded Olcostephanus cf. O. atherstoni Baumberger
(non Sharpe) (=? O. baini (Sharpe)) from the zone of Buchia keyserlingi
(Lahusen) in Oregon, considered to be of Middle to Upper Valanginian age,
and associated with the genera Bochianites, Neocomites, Thurmanniceras,
Neocraspedites, and Polyptychites.

From Fernao Velosa, in northern Mozambique, Spath (1930) has recorded
Olcostephanus schenki (Oppel) (= O. baini (Sharpe)), and Haploceras (Neolis-
soceras) cf. grasianum (d’Orbigny). Strata of a similar age also crops out at
Mahiba Hill, to the west of Port Amelia, where Newton (1924) recorded frag-
ments of Lytoceras together with the belemnite Duvalia. From this same locality
Spath (1930: 134) also records a ‘.. . portion of the periphery of a Neocomitid
(Lyticoceras of the type of L. regalis (Bean) or Neocomites neocomiensis
(d’Orbigny) as figured by Sayn), and the impression of a fragment of the
Uitenhage Bochianites africanus are decisive and unmistakable’.

The coarse ribbing of the olcostephanids from the Uitenhage Group has
long been a distinctive feature. It is of interest to note, therefore, that where the
basal Hauterivian is best known (Debelmas & Thieuloy 1963), the associated
species of Olcostephanus are frequently finely and densely ribbed, with such
taxa as O. filosus (Baumberger) and O. sayni (Kilian) common. Olcostephanus
atherstoni (Sharpe) also occurs but, as will be shown, represents a stock-name
for a number of homoeomorphic macroconchs. The absence of such finely
ribbed forms, together with Acanthodiscus, from the Sundays River Formation
supports the suggestion that this unit should be considered of latest Valanginian
age only.

In order to help resolve the Valanginian—Hauterivian boundary problem,
it seems necessary to introduce a zone of Olcostephanus baini at the top of the
Valanginian (Collignon 1962). The following Valanginian—Lower Hauterivian
zonal scheme would appear to be of global significance:

Crioceratites duvali

Lower Hauterivian : i
pi { Acanthodiscus radiatus

Olcostephanus baini
Saynoceras verrucosum
Kilianella roubaudiana
Thurmanniceras thurmanni

Valanginian

The Uitenhage fauna may be assigned, therefore, to the assemblage zone
of O. baini. Besides the Uitenhage species described herein, other important
~ elements of the zone include Neocomites spp, Himantoceras spp, Neohoploceras
spp, Distoloceras spp, Sarasinella spp, Leopoldia spp, Bochianites spp, Olco-
stephanus spp, together with the less diagnostic Hemilytoceras liebigi (Oppel),
Haploceras’ (Neolissoceras) grasianum (dOrbigny), Phylloceras thetys
(d’Orbigny), P. serum (Oppel) and so on.

154 ANNALS OF THE SOUTH AFRICAN MUSEUM

SYSTEMATICS

A revision of the Uitenhage ammonite fauna, characterized by the abund-
ance of forms assigned to the genus Olcostephanus, some of which have attained
unusually large size, has long been overdue, especially in view of the recognition
of sexual dimorphism within many groups of the Perisphinctaceae, and more
recently in the genus Olcostephanus itself (Riccardi et al. 1971). Moreover, the
erection of numerous species (and genera) within the Olcostephaninae, based
upon subtle and generally insignificant differences with no regard for intra-
specific variation, has led not only to taxonomic chaos but also to difficulties
in interregional correlation.

The following abbreviations are used to indicate the source of the material:

AM Albany Museum, Grahamstown

BM British Museum (Natural History), London
LJE, AAS Geological Survey, Pretoria

MNHP Natural History Museum, Paris

OUM Oxford University Museum, Oxford

PEM Port Elizabeth Museum, Port Elizabeth

Fig. 2. Schematic diagram showing the measurements dis-
cussed in the text.

REVISION OF LATE VALANGINIAN CEPHALOPODA 155

The abbreviations for measurements given in the text are explained in
Figure 2 where D = diameter, H = height, Uo = outer umbilical diameter
(between umbilical bullae), Ui = inner umbilical diameter (between umbilical
seams), Wi = intercostal width. All measurements are given in millimetres,
and dimensions, as a percentage of the diameter, are included in parentheses.

Class CEPHALOPODA Cuvier, 1797
Subclass AMMONOIDEA Zittel, 1884
Order PHYLLOCERATIDA Arkell, 1950
Superfamily PHYLLOCERATACEAE Zittel, 1884
Family Phylloceratidae Zittel, 1884
Subfamily Phylloceratinae Zittel, 1884
Genus Partschiceras Fucini, 1920
Type species Ammonites partschi Stur, 1851 (non Klipstein 1843);
by original designation of Fucini, 1920
Discussion

This is a long-ranging genus, having been recorded from Lower Jurassic
(Sinemurian) to Upper Cretaceous (Maastrichtian). It differs from Phylloceras
in being more inflated with maximum width at mid-flank, and in commonly
developing ribs as well as lirae in maturity. Moreover, Phylloceras has triphyllic
saddles not diphyllic as in Partschiceras. Wiedmann (1962 has recently included
Phyllopachyceras Spath as a junior synonym in this genus.

Partschiceras rogersi (Kitchin, 1908)
Fig. 3

Phylloceras rogersi Kitchin, 1908: 179, pl. 8 (fig. 19, 19a—c). Spath, 1930: 140. Du Toit, 1954:
384. Haughton, 1963: 274.

Non Phylloceras rogersi Kitchin var. nov., Krenkel, 1910: 223, pl. 22 (fig. 9) (= P. krenkeli
Zwierzycki).

Phyllopachyceras rogersi (Kitchin) Collignon, 1962: 20, pl. 181 (figs 815-816).

Material

The holotype is the only example so far collected from the Sundays River
Formation and its present whereabouts is unknown.

Fig. 3. Partschiceras rogersi (Kitchin). The holotype, by
monotypy, with part of the suture (after Kitchin 1908). x 1.

156 ANNALS OF THE SOUTH AFRICAN MUSEUM

Holotype

By monotypy, the original of the specimen figured by Kitchin (1908: 179,
pl. 8 (fig. 19, 19a-c)), from the Sundays River Formation, mid-way up the cliffs
behind Colchester.

Diagnosis
An immature Partschiceras of late Valanginian age characterized by
extremely fine, dense lirae, with no sign of ribbing.

Description

A rather inflated, very involute form, with slightly convex flanks and
maximum width at mid-flank. Ornament comprises fine, prorsiradiate, flexuous
ribs. At a whorl height of 9 mm there are 12 ribs within a 2 mm distance along
the venter.

Discussion

Kitchin’s holotype, the whereabouts of which is not known, still remains
the only record of this species (and genus) from the Sundays River Formation.
Since this specimen appears to be immature and the coarse ribbing of this
genus appears typically in maturity, the validity of this species is in doubt.

Partschiceras infundibulum (d’Orbigny) appears to be more compressed,
whilst also possessing the coarse ribbing not seen in the Uitenhage example.

Phylloceras krenkeli Zwierzycki was established for the specimen figured
by Krenkel (1910: 223, pl. 22 (fig. 9)) as P. rogersi var. nov., and said to differ
from the Uitenhage species in being more compressed, with the greatest width
at the umbilical margin and not at mid-flank, having a different suture, and in
being more coarsely ribbed, with only eight ribs within a 2 mm distance.

Occurrence

The holotype was collected from mid-way up the cliffs behind Colchester,
and thus near the top of the marine succession.

Collignon (1962) has recorded this species from the Lower Valanginian
of Madagascar, while Haughton (1963) mentions its occurrence in beds of
Lower Hauterivian age from the same island. It would seem, therefore, that
P. rogersi, like many phylloceratids, is a relatively long-ranging species.

Suborder ANCYLOCERATINA Wiedmann, 1966
Superfamily ANCYLOCERATACEAE Meek, 1876
Family Bochianitidae Spath, 1922
Subfamily Bochianitinae Spath, 1922
Genus Bochianites Lory, 1898

Type species Baculites neocomiensis d’Orbigny, 1842;
by original designation of Lory, 1898.
Discussion

This is a relatively long-ranging genus, having been recorded from Tithonian
to Hauterivian strata. Janenschites was said to have a more denticulate suture

REVISION OF LATE VALANGINIAN CEPHALOPODA S57

than Bochianites with long elements, but similar ornament. It has, however,
recently (Wiedmann 1962) been included in the synonymy of Bochianites.

Bochianites glaber Kitchin, 1908
Fig. 4

Bochianites glaber Kitchin, 1908: 181, pl. 8 (figs 20-21). Hatch & Corstorphine, 1909: 303
fig. 75f. Spath, 1930: 155. Du Toit, 1954: 384. Klinger & Kennedy 1979: 17.

Material

The lectotype, SAM-4695, with some fragments crowded together in the
same block of matrix (SAM-—12736) from c. 2 km upstream from the Swartkops
bridge.

Holotype

By lectotype designation herein, the original of Bochianites glaber figured
by Kitchin (1908: 181, pl. 8 (fig. 20)) from the road below the railway cutting,
c. 2 km from Rawson bridge on the main line in the Swartkops River Valley.

Fig. 4. Bochianites glaber Kitchin. A. Ventral view of
the lectotype, SAM-4695, x 4. B-C. Ventral and
lateral views of the lectotype, x 3.

158 ANNALS OF THE SOUTH AFRICAN MUSEUM

Diagnosis

A small, very finely ribbed, juvenile Bochianites, with an elliptical whorl
section at the largest known growth stages.

Description

A small, slowly expanding Bochianites, with an initially circular whorl
section, becoming elliptical with age. Ornament comprises fine, prorsiradiate
lirae, arching across the siphonal line and straight or slightly convex adapically
across the dorsum.

Discussion

A number of fragmentary straight shafts within a single block of matrix
would seem to belong to this species, the fine ribbing only becoming visible
under a hand-lens. No constrictions were observed.

Baculites maldonadi Karsten (1856: 105, pl. 2 (fig. 2)) (Fig. 5) approaches

Fig. 5. Baculites maldonadi Karsten (after Karsten 1856). x 1.

the present species but as they represent different growth stages comparison is
difficult. The adapical portion of Karsten’s species shows the same thread-like
ribbing as B. glaber, but adorally it coarsens considerably to resemble
B. africanus.

Whilst there are a number of other Bochianites species with which B. glaber
might be compared, the immature nature and poor preservation of the South
African material make such comparisons unwarranted. Indeed, B. glaber might
even represent the earliest growth stages of Umgazaniceras thieuloyi Klinger
& Kennedy (1979: 12, figs 1-2, 3A—D, 4-5). As such it is perhaps best regarded
as a nomen dubium.

Occurrence

At present B. glaber is known only from the Sundays River Formation
and is thus of latest Valanginian age.

Bochianites africanus (Tate, 1867)
Figs 6-7

Hamites africanus Tate, 1867: 150, pl. 7 (fig. 5). Holub & Neumayr, 1882: 271.
Bochianites africanus (Tate) Kitchin, 1908: 225. Spath, 1930: 153, pl. 14 (figs 2-3), pl. 15
(fig. 3). Du Toit, 1954: 384. Klinger & Kennedy, 1979: 17, fig. 3F—H.

REVISION OF LATE VALANGINIAN CEPHALOPODA 159

Material

Numerous specimens, including BM-—C25227-9, AM-844, 846, SAM-—
PCU1586—88, 5706, and 12784-88.

Holotype

By the lectotype designation of Spath (1930), the original of Tate’s (1867)
plate 7, figure 5a, BM—C25228, from Prince Alfred’s Rest at the Sundays
River mouth.

Diagnosis

A coarsely ribbed species of Bochianites with 4-7 prorsiradiate ribs in a
distance equal to twice the whorl width and an elliptical to subtrigonal whorl
section.

Fig. 6. Bochianites africanus (Tate) The syntypes, of which the top right-hand
specimen was selected as lectotype (after Tate 1867; composite suture after
Spath 1930). x 1.

Description

A slowly expanding, coarsely ribbed Bochianites with an elliptical whorl
section in immaturity, becoming subtrigonal at large sizes when the maximum
width is near the dorsal shoulders. The dorsum is flattened in all but the youngest
individuals. Ornament comprises coarse annular ribs, strongly projected on the
flanks and sharply arched across the venter. Because of the oblique angle at
which the ribbing crosses the distinctly tabulate venter of mature individuals,
the ribs are broadest across the venter as well as being asymmetrical with a
gentle adapical slope and a steep adoral slope. Rib density varies between
4 and 7 in a distance equal to twice the costal whorl width.

Discussion

Bochianites africanus (Tate) is closest to B. neocomiensis (d’Orbigny)
(1842a, pl. 138 (figs 1-5)) which has, however, an almost perfectly circular
whorl section. Spath (1930: 154) considered the present species to be very close
to Baculites granatensis Karsten (1856: 105, pl. 2 (fig. 1)), to which illustration
it certainly bears a considerable resemblance but, according to J.-P. Thieuloy
(in litt. 1980), the Columbian species (Fig. 8) is a late Cretaceous Baculites.

160 ANNALS OF THE SOUTH AFRICAN MUSEUM

ARE

ce
*

Fig. 7. Bochianites africanus (Tate). A. Lateral view of AM-844, x 0,75. B—D. Dorsal, ventral
and lateral views of AM-846, x 0,75. E-F. Lateral and ventral views of AM-868, x 0,75.
G-I. Ventral, lateral and dorsal views of a specimen in the South African Museum, x 1.
J-K. Lateral and cross-sectional views of an unnumbered specimen in the South African
Museum, x 0,75. L. Lateral view of SAM-—PCU1586, x 1. M-—O. Lateral, ventral and dorsal
views of a specimen in the South African Museum, x 1. P. Dorsal view of SAM—PCU1587,
x 1. Q-R. Lateral and ventral views of a specimen in the South African Museum, x 1.
S-T. Ventral and lateral views of a specimen in the South African Museum, xX 1. U. Lateral
view of an unnumbered specimen in the South African Museum, x 1.

Fig. 8. Baculites granatensis Karsten (after Karsten
1856); Oe

REVISION OF LATE VALANGINIAN CEPHALOPODA 161

Imlay (1938: 585, pl. 6 (figs 1,11-13)) figured specimens of a Bochianites
sp. from the Upper Member of the Taraises Formation which he compared
with B. neocomiensis (d’Orbigny). The ornament is closely comparable with
that of B. africanus, but since the whorl section was not figured or mentioned
its specific assignment is unknown.

Occurrence

Bochianites africanus (Tate) is as yet known only from northern Mozam-
bique and South Africa. It seems surprising that it has not yet been recorded
from Madagascar in view of the relative abundance of this species in the
Sundays River Formation.

Superfamily PERISPHINCTACEAE Steinmann, 1890
Family Olcostephanidae Haug, 1910
Subfamily Olcostephaninae Haug, 1910
Genus Olcostephanus Neumayr, 1875

Type species Ammonites astieri d’Orbigny, 1840;
by original designation of Neumayr, 1875.
1875 Olcostephanus Neumayr
1889 Holcostephanus Sayn
1892 Astieria Pavlow
1923a Subastieria Spath
19236 Parastieria Spath
1924 Rogersites Spath
1938 Mexicanoceras Imlay
1938 Maderia Imlay
1964 Jeannoticeras Thieuloy
1966 Taraisites Cantu Chapa
1966 Satoites Cantu Chapa
1977a Lemurostephanus Thieuloy

Emended diagnosis

Compressed to strongly inflated cadicones, with strongly arched to well-
rounded venters. Primary ribs are usually present on the umbilical wall, com-
monly terminating in tubercles at the umbilical shoulder (except on the outer
whorls of Parastieria and Jeannoticeras), from which arise straight or slightly
curved secondary ribs, usually in fasciculate bundles (in pairs in Jeannoticeras).
There are commonly 3-4 secondaries per bundle, although there may be as
many as 6-9 or as few as 1-2. Secondary ribs may bifurcate on the flanks, while
intercalated ribs between bundles are the rule. Ribbing generally passes uninter-
rupted across the venter, although it may weaken considerably where a ventral
furrow is present (i.e. Mexicanoceras). Parabolae may or may not be present,
but never on the outer whorls of females. This genus is dimorphic; males small
and with lappets, females larger and with simple peristomes. Age: Upper
Valanginian—Middle Hauterivian.

162 ANNALS OF THE SOUTH AFRICAN MUSEUM

Discussion

In addition to the objective synonyms Holcostephanus and Astieria,
Wiedmann & Dieni (1968) also placed Rogersites within Olcostephanus s.s., as
well as including Parastieria and Subastieria within the genus as subgenera.
Capeloites was considered by these authors to be of dubious status, but, as
since shown by Thieuloy (1969), is distinct.

In 1924 Spath erected the new genus Rogersites, types species R. modderensis
(Kitchin), without giving a formal diagnosis. In his revision of the Uitenhage
Cephalopoda in 1930, Spath had still not formulated a generic diagnosis for
Rogersites, but included into it all previously described olcostephanids from the
Uitenhage Group. In his monograph of the Cephalopoda of the Neocomian
Belemnite Beds of the Salt Range, he (Spath 1939: 11) diagnosed Rogersites,
stating that ‘... although there are no typical Rogersites (e.g. R. modderensis
Kitchin sp., R. baini Sharpe sp. and R. kitchini Spath), with few and very coarse
primary and secondary ribs and prominent umbilical edge’, while further on
(p. 31) he refers to ‘...the typical Rogersites characters, namely a coronate
cadicone and vertical umbilical wall, at large diameters, while retaining coarse
ribbing’. At this time Spath was using Rogersites both as a genus and a sub-
genus. Thus (Spath 1939: 11), ‘...there is one common and widely quoted
transitional species between Rogersites and Olcostephanus. This is O. (Roger-
sites) schenki (Oppel). On page 16 he refers to Rogersites sphaeroidalis and
R. atherstoni, but on page 19 states *...I previously referred O. uitenhagensis
to Rogersites, but like R. atherstoni, R. sphaeroidalis and the many passage
forms between these species, O. uitenhagensis is one of the transitions from
Rogersites to Olcostephanus’. On page 31 Spath states ‘...O. (R.) schenki is
the most strongly and distantly ribbed form of Olcostephanus from the Salt
Range and the only species that may be compared with such typical Rogersites
as R. modderensis (Kitchin). .. . Since however, no large examples of O. schenki
have yet been found or recognized, it is uncertain whether it develops the typical
Rogersites characters.’

Imlay (1938) erected three new genera, all comprising entirely septate
pyritic nuclei, within the Olcostephanidae, viz. Maderia, Mexicanoceras, and
Ceratotuberculus. Mexicanoceras was considered ‘... similar in form, sculpture
and suture-line to Olcostephanus. It differs from Olcostephanus in having a
ventral furrow on the outer whorls, a more inflated form, broader whorl section,
and its much smaller size.’ These features are herein considered of only sub-
generic value and Mexicanoceras is accordingly placed within Olcostephanus
s. 1. Maderia was defined by Imlay (1938) as *. . . similar to Subastieria (defined
by Spath, 1923, p. 32) but exhibits a slight thinning of the ribs along the mid-
ventral line of the outer whorls. Compared with Mexicanoceras its whorl
section is more depressed, its umbilicus wider, and the ventral thinning of the
ribs less pronounced.’ The importance attached to the very slight weakening of
the ribs across the siphonal line in Maderia by Imlay (1938) appears unjustified
and, in the author’s opinion, Maderia comprises an heterogeneous assemblage

REVISION OF LATE VALANGINIAN CEPHALOPODA 163

of Olcostephanus s.s. and possibly O. (Subastieria), and is, therefore, superfluous.
The genus Ceratotuberculus is characterized by the appearance, after the third
whorl, of high, thick, ventrolateral bullae, as well as by possessing a ventral
furrow, whilst retaining the other olcostephanid characters. If, as suggested by
Imlay (1938), this genus is of early Hauterivian age, then it is probably descended
from Saynoceras. If, however, it is of late Valanginian age, then the differences
probably warrant at most subgeneric separation from Saynoceras.

Spath (1924) erected the genus Suwbastieria for Olcostephanus sulcosus
Pavlow, differentiated from Olcostephanus by its younger age and highly
coronate whorl section. According to Wright (in Arkell et al. 1957), Subastieria
closely resembles the inner whorls of some Rogersites which, as just shown,
is a synonym of Olcostephanus s.s. Moreover, Wiedmann & Dieni (1968) have
recently shown Subastieria to range into the Upper Valanginian, whilst
Olcostephanus is common in the Lower Hauterivian of the Swiss Jura (Debekmas
& Thieuloy 1963). Consequently, not only do Subastieria and Olcostephanus
have the same stratigraphic range, but they are also morphologically very
similar. However, in Subastieria the whorl section is coronate at all growth
stages, whereas microconch forms of Olcostephanus become coronate only with
the egression of the umbilical seam on the body chamber. This appears to be a
subtle but distinct difference, and consequently Wiedmann & Dieni (1968) are
provisionally followed in treating it as a valid subgenus.

Fig. 9. Olcostephanus (Olcostephanus) cf. atherstoni

(Sharpe). The holotype of Taraisites bosei Cantu

Chapa, from the Taraises Formation of northern
Mexico (after Bose 1923). x 1.

Cantu Chapa (1966) erected the genus Taraisites for the specimen of

Astieria aff. baini Sharpe figured by Bose (1923: 76, pl. 2 (figs 3-5)) (Fig. 9
herein), renaming it 7. bosei Cantu Chapa. This genus was characterized by
distant secondary ribbing arising in bundles of 2-3 from the umbilical bullae.
As remarked by Riccardi et al. (1971), Cantu Chapa’s disregard for ontogenetic
variation resulted in his separation of, amongst others, the small, coarsely
ribbed paratypes of ‘Rogersites’ prorsiradiatus Imlay, under the new species

164 ANNALS OF THE SOUTH AFRICAN MUSEUM

name of 7. neoleonense, from the larger and more densely ribbed holotype.
Indeed, he included the type of Olcostephanus baini (Sharpe) within his new
genus, thus separating a microconch from its far more densely ribbed macro-
conch dimorph. There can be no hesitation in following Riccardi et al. (1971)
in considering Jaraisites a synonym of Olcostephanus s.s.

Cantu Chapa (1966) also erected the new genus Satoites for the supposedly
Berriasian Olcostephanus sp. nov. figured by Sato (1958: 590, pl. 28 (figs 1-3),
fig. 2) (Fig. 10 herein), and considered ancestral to Mexicanoceras from which

Fig. 10. Olcostephanus oshimensis (Cantu Chapa). The
type species of the genus Satoites, allegedly from the
Berriasian of Japan (after Sato 1958). x 1.

it was said to differ by the absence of a ventral furrow, whilst possessing
prominent parabolae. Since the main criterion by which Imlay (1938: 562)
distinguished Mexicanoceras from Olcostephanus was in the presence of
a ventral furrow, Satoites must be considered a junior synonym of Olco-
stephanus §.S.

Thieuloy (1964: 212) erected the subgenus Olcostephanus (Jeannoticeras)
for Ammonites jeannoti d’Orbigny (1840, pl. 56 (figs 3—-5)) (Fig. 11 herein),
characterized by the absence of umbilical tuberculation in the adult, and by the
secondary ribbing which commonly arises in pairs (rarely three) from the
primary ribs at the umbilical shoulder. Through C. W. Wright, the writer has
seen a collection of Olcostephanus (Parastieria) from the Lower Hauterivian at
Speeton. This subgenus is based upon a microconch species with lappets which
upto the body chamber shows numerous primaries giving rise to fine secondaries
in twos and threes. At this stage, Parastieria is indistinguishable from Jeannoticeras
and it is only on the body chamber that the broad, flat ribs characteristic of
Parastieria appear. Further work may show that Jeannoticeras is a junior sub-
jective synonym of Parastieria, though for the time being they are treated as
distinct.

The genus Dobrodgeiceras Nikolov (1962: 69) was established for the type
species D. ventrotuberculatum Nikolov from the Upper Valanginian of Bulgaria,
and said to differ from Valanginites mainly in the presence of ventral tubercles
and more prominent primaries (Nikolov 1962: 69). Thieuloy & Gazay (1967)
assigned their entire French fauna to Dobrodgeiceras wilfridi (Karakasch)

REVISION OF LATE VALANGINIAN CEPHALOPODA 165

Fig. 11. Olcostephanus (Jeannoticeras) jeannoti (D’Orbigny), < 1. A-B. A specimen in the

collections of the University of Paris from the Lower Hauterivian of Montclus, Hautes-

Alpes. C. The crushed lectotype in the Natural History Museum, Paris, R3114 (D’Orbigny

Collection No. 4865a) from the Lower Hauterivian of ?Serrais, Hautes-Alpes. Note the
parabola. —

Fig. 12. Dobrodgeiceras wilfridi (Karakasch). The holotype of Holco-
stephanus wilfridi Karakasch from the Upper Valanginian of Crimea
(after Karakasch 1902). x 1.

(Fig. 12) of which they considered ventrotuberculatum merely a subspecies.
Riccardi & Westermann (1970) considered that the French sample was derived
from a single population, and since ‘...the use of the subspecies category in
palaeontology is usually confined to stratigraphically or geographically distinct
taxa.... The interpretation of this variation as genetic polymorphism is

166 ANNALS OF THE SOUTH AFRICAN MUSEUM

probably correct.’ According to Riccardi & Westermann (1970), the position
of the flank tubercles in Dobrodgeiceras is ventrolateral, and not lateral or
periumbilical as described by Thieuloy & Gazay (1967: 77). The similarity with
Valanginites led Riccardi & Westermann to place Dobrodgeiceras within the
Polyptychitinae. However, since this subfamily is generally distinguished by
the branching of the secondaries, a feature unknown in both Valanginites and
Dobrodgeiceras, there appears no reason why they should not preferably be
included within the Olcostephaninae. .

Valanginites is a much misunderstood genus although the situation has
recently been clarified somewhat by Thieuloy (1977a). Thus, a survey of the
literature shows that at some time or other most strongly inflated, coarsely
ribbed olcostephanids have been referred to this genus. Spath (1930) included
O. perinflatus (Matheron) and O. stephanophorus (Matheron) in this genus,
rectifying his mistake in 1939, but now including O. crassus (Zwierzyck1) and,
tentatively, Holcostephanus bachelardi Sayn into Valanginites. Imlay (1938)
erected the species Valanginites angusticoronatus for a coarsely ribbed, inflated
form which is herein considered to be an Olcostephanus, very close to O. rogersi
(Kitchin) (2). The only undoubted species of Valanginites appear to be the type
species V. nucleus (ROmer), of which V. utriculus (Matheron) was considered
a synonym by Roch (1930), V. dolioliformis (Roch) (Fig. 13), V. tijerensis
Imlay, V. psaephoides (Mayer—Eymar) (including V. bachelardi (Sayn)), and
V. simplus (d’Orbigny). Of these, umbilical tubercles are present in V. dolioli-
formis and V. tijerensis.

Leanza (1957) described four species of the Hauterivian genus Simbirskites
from the Upper Valanginian of Argentina. Rawson (1971: 42), however, con-
siders this material generically misidentified since ‘...all four of Leanza’s

Fig. 13. Valanginites dolioliformis (Roch).
The holotype from Morocco (after Roch
1930)) x i

REVISION OF LATE VALANGINIAN CEPHALOPODA 167

species differ from Simbirskites in that the ribs extend straight across the venter
instead of curving forwards. Leanza’s species appear closer to Rogersites; the
recorded stratigraphical horizon (late Valanginian) would agree with this.’

The Argentinian material can immediately be precluded from the sub-
genera Mexicanoceras, Jeannoticeras, and Parastieria. They differ from Olco-
stephanus s.s. (= Rogersites) in having prorsiradiate primary ribs throughout
ontogeny, in having a gently sloping umbilical wall and rounded umbilical
shoulder throughout ontogeny, and in being more evolute. While any one, or
even all, of these characters may be found at some growth stage in Olcostephanus
s.s., they are not known to persist together throughout ontogeny. The sloping
umbilical wall, coronate whorl section and prorsiradiate primaries of Leanza’s
S. araucanus are, however, all to be found in Olcostephanus (Subastieria)
nicklesi Wiedmann & Dieni, and it is to this subgenus that the Argentinian
material is best referred. The Argentinian material would seem to comprise
some of the few adult examples of this subgenus yet recorded. In maturity,
therefore, Subastieria should be regarded a close homoeomorph of Simbirskites;
possibly the latter is descended from the former.

Thieuloy (1977a: 432) has recently introduced the new subgenus Lemuro-
stephanus within Olcostephanus for forms with a very wide umbilicus (40-45 %
of the diameter), well-developed parabolae and primary ribs terminating in
pointed bullae from which arise bundles of 2-4 secondary ribs. Besides the type
species, Holcostephanus madagascariensis Lemoine (Fig. 14), Thieuloy (1977a)

Fig. 14. Olcostephanus (Olcostephanus) madagas-

cariensis (Lemoine). The holotype from the Lower

Valanginian of Madagascar (after Collignon 1962).
Sade

168 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 15. Olcostephanus (Olcostephanus)

mitreanus (d’Orbigny) (3). The holotype

of Astieria detonii Rodighiero from Venice
(after Rodighiero 1919). x 1.

Fig. 16. Olcostephanus (Olcostephanus) mitreanus

(d’Orbigny) (3). The holotype of Olcostephanus

wynnei Spath from the Spiti Shales of Pakistan
(after Spath 1939). x 1.

also included in this subgenus Spiticeras detonii Rodighiero (Fig. 15), Olco-
stephanus wynnei Spath (Fig. 16), Olcostephanus mitreanus (d’Orbigny) (Fig. 17),
O. sanctifirminensis Thieuloy, Holcostephanus chaignoni Sayn, Maderia?
latiumbilicata Imlay and the species of ‘Simbirskites’ described by Leanza (1957)
and discussed above. Of these, Leanza’s ‘Simbirskites’ species together with
H. chaignoni Sayn, Maderia? latiumbilicata and O. (L.) sanctifirminensis can
adequately be included in the subgenus Suwbastieria, whilst Spiticeras detonii,
Olcostephanus wynnei and O. mitreanus are conspecific microconchs whose

REVISION OF LATE VALANGINIAN CEPHALOPODA 169

Fig. 17. Olcostephanus (Olcostephanus) mitreanus (d’Orbigny). The syntypes in the Natural

History Museum, Paris, R3118 (D’Orbigny Collection No. 4871), of which the smaller (a

microconch) has been selected as lectotype (Thieuloy 1977a). The paralectotype would seem
to be an immature macroconch. x 1.

probable macroconch, O. collignoni (Besairie) (Fig. 18), is a typical Olcostephanus
s.s. In the writer’s opinion, the subgenus Lemurostephanus comprises an hetero-
geneous assemblage of O. (Subastieria) and O. (Olcostephanus) and is of little
taxonomic significance. The author prefers not to use the name.

During the early ontogenetic stages (Fig. 19), the suture line of Olco-
stephanus is relatively simple with long, thin saddles and a trifid first lateral lobe
(L). With ontogeny the suture becomes very deeply incised (Fig. 20K) with
long, thin folioles and lobules. Olcostephanus (Subastieria) hispanicus (Mallada)
(Fig. 20H-I) shows a similar suture line, as does Dobrodgeiceras broggianum
(Lisson), although in the latter the saddles are broader and shorter (Fig. 20J).

The following subdivisions within the Olcostephaninae are here recognized:

S. (Saynoceras). Small, inflated microconchs with trapezoidal whorl sections.

ANNALS OF THE SOUTH AFRICAN MUSEUM

170

‘| x ‘IeoseSepeypy ‘Ayxiquiy jo uvluIsurjeA Jsddq oy} woul odAjojoy oy, (4) (al4Tesag) jvous)]/0

2 (snuDvydajso7Q) snuvydajso.jO °8{ *3I4

REVISION OF LATE VALANGINIAN CEPHALOPODA 171

30,5mm

12mm

6,5mm

3mm

2mm

Fig. 19. Sutural ontogeny of Olcostephanus (Olcostephanus) atherstoni (Sharpe) (3) (after
Riccardi et al. 1971).

172 ANNALS OF THE SOUTH AFRICAN MUSEUM

My
® up

|
/

Fig. 20. Olcostephaninid suture lines. A. Olcostephanus (Olcostephanus) guebhardi (Kilian)
(after Baumberger 1908). B. Olcostephanus (Olcostephanus) imbricatus (Baumberger) (after
Baumberger 1908). C. Olcostephanus (Olcostephanus) leptoplanus (Baumberger) (after
Baumberger 1907). D. Olcostephanus (Olcostephanus) ventricosus (von. Koenen) (after
Tzankov 1943). E. Olcostephanus (Olcostephanus) inordinatus (Tzankov) (after Tzankov
1943). EF. Olcostephanus (Olcostephanus) inordinatus (Yzankov) (after Pictet & Campiche
1860). G. Valanginites bachelardi (Sayn) (after Tzankov 1943). H,J. Olcostephanus (Sub-
astieria) hispanicus (Mallada) (after Tzankov 1943). I. Dobrodgeiceras broggianum (Lisson)
(after Riccardi & Westerman 1970). K. Olcostephanus (Olcostephanus) baini var. sphaeroidalis
(Spath); drawn from the holotype, x 1.

REVISION OF LATE VALANGINIAN CEPHALOPODA 173

Umbilical tubercles give rise to 1-2 weak ribs leading to ventrolateral
tubercles; ribbing normally subordinate to tubercles on outer whorl, but
may be sharp at some stages (after Wright in Arkell et al. 1957). Age:
low Upper Valanginian.

S. (Ceratotuberculus). Small, inflated, like Saynoceras, but with more, 2-4,
secondaries per tubercle, and with a smooth ventral furrow. Age: Lower
Hauterivian.

O. (Olcostephanus). Compressed to inflated forms, usually with primary ribs
terminating in tubercles on the umbilical shoulder, from which secondary
ribs diverge in fasciculate bundles to pass uninterrupted across the venter.
Parabolae may or may not be present. Dimorphic; males small, with
lappets, females large and with simple peristomes. Age: Valanginian—
Middle Hauterivian.

O. (Subastieria). Similar to O. (Olcostephanus), but with sloping umbilical wall,
prorsiradiate primaries and a coronote whorl section throughout ontogeny.
Adults homoeomorph Simbirskites. Age: Upper Valanginian—Lower
Hauterivian.

O. (Parastieria). Based upon a lappeted microconch species. Inner whorls with
numerous, fine primaries giving rise to 2—3 flexuous secondary ribs with
intercalatories between bundles. Body chamber high-whorled, compressed,
with slightly sinuous, broad, flat-topped ribs and no umbilical tubercles,
Age: Lower Hauterivian.

O. (Jeannoticeras). Similar to O. (Parastieria) from which it is doubtfully
separable, but with fine bundled ribbing retained on to the adult body
chamber. Dimorphic. Age: Lower Hauterivian.

O. (Mexicanoceras). As for O. (Olcostephanus) but small, with ribs interrupted
on outer whorls by a ventral furrow. Age: ?Lower Hauterivian.

Dobrogeiceras. Like O. (Olcostephanus) but very involute; umbilical tubercles
have moved to the ventrolateral position, with a consequent lengthening
of the prominent primary ribs. Ventral tubercles (?dimorphic) may he
present on the body chamber. Age: Upper Valanginian.

Jeanthieuloyites. Similar to Olcostephanus, but with long, radial, nontuberculate
primaries extending to mid-flank where they regularly bifurcate. Prominent,
oblique parabolae. Age: Upper Valanginian.

Valanginites. Extremely inflated with very narrow umbilicus. Umbilical tubercles
weak to absent. Secondary ribbing coarse, simple. Age: Upper Valanginian.

Capeloites. Small, compressed, with lappets (microconch). Inner whorls with
prominent umbilical bullae from which pairs of ribs (the adoral one thick
and robust and the adapical one fine and delicate) are looped to siphonal
clavi. Intercalatories occur. On the body chamber siphonal clavi disappear,
umbilical bullae weaken considerably, and all secondaries become fine,
sinuous, passing uninterrupted across the venter. Age: Uppermost
Valanginian—basal Hauterivian.

174 ANNALS OF THE SOUTH AFRICAN MUSEUM

SEXUAL DIMORPHISM IN AMMONITES

It has long been noted that many ammonites occur in both large and small
forms within the same stratum. Whereas the large forms have simple peristomes,
the persitomes of the small forms frequently have various types of apertural
adornment, e.g. lateral lappets, ventral rostra, ventral horns, etc. Furthermore,
many such pairs were seen to have identical early whorls. This led to the concept
of sexual dimorphism and, by analogy with extant invertebrates, the large
forms were taken to represent the females and the smaller forms the males.
Within recent years much attention has been given to the phenomenon of
sexual dimorphism within fossil Cephalopoda (Callomon 1957, 1963; Makowski
1962a, 1962b; Westermann 1964, 1969; HouSa 1965; Lehmann 1966, 1969;
Palframann 1966, 1967; Cope 1967; Cobban 1969; Zacharov 1969; Reyment
1971; Riccardi et al. 1971; Kennedy & Cobban 1976), and it is now a widely
accepted phenomenon.

It has often been noted, and Makowski (1962qa) in his classical study placed
much emphasis on the fact, that the inner whorls of the female are invariably
identical to the male dimorph.

Douvillé (1880), in his study of Morphoceras pseudoanceps, noted that it
had inner whorls identical to those of M. polymorphus and suggested that they
possibly represented a dimorphic pair. Haug (1893) found that the inner whorls
of Sonninia sowerbyi and S. sulcata were identical, and hence indistinguishable,
up to a diameter of 30 mm, after which the later ontogenetic stages differed
markedly. Makowski (1962a) has provided numerous examples from such widely
diversified groups as the haploceratids, stephanoceratids, hildoceratids, cheilo-
ceratids, and the scaphitids, where the inner whorls of the large macroconchs
(2) were identical to the smaller microconchs (9). Palframann (1966) showed
Creniceras renggeri (Oppel) and Taramelliceras richei (de Lorial) to be identical
in every feature up to a diameter of about 8 mm. On the body chamber 7. richei
differed from C. renggeri in the development of ventrolateral spines. Further-
more, the peristome of C. renggeri was highly ornate, whereas that of 7. richei
was relatively simple. Palframann concluded that C. renggeri and T. richei were
merely male and female of the same species. Similar identity of the early growth
stages of Distichoceras bicostatum (Stahl) and Horioceras baugieri (d’Orbigny)
led Palframann (1967) to consider them to represent a sexually dimorphic pair.
Sutural approximation within a juvenile macroconch of D. bicostatum led
Palframann (1967: 73) to suggest *. . .it may be that the specimen in question has
changed sex during life as do some living molluscs. The latter explanation is con-
sidered unlikely though, as this feature has not been seen in other specimens
studied here, the explanation of the phenomenon is itself, no doubt, unusual.’

Cope (1967), in a re-examination of the fauna of the Upper Kimmeridge
Clay of Dorset, observed a unique type of dimorphism in the perisphinctid
Pectinatites, in which the apertural ornament of the microconch is in the form
of a ventral horn. As a general rule, he found that in this genus the microconch
was usually slightly coarser ribbed than its macroconch at a similar diameter.

REVISION OF LATE VALANGINIAN CEPHALOPODA 175

Riccardi et al. (1971) observed that identity of ornament between
Olcostephanus atherstoni (Sharpe) dimorphs was restricted to the nucleus under
20 mm in diameter.

It is significant, however, that in all cases where there is identity of early
ontogenetic stages in ammonites the change is always from a younger, typically
male-type morphology to an older female-type morphology. According to
Fretter & Graham (1964: 130) ‘...consecutive hermaphrodites change sex
once in their lives, usually from a younger male to an older female phase’,
whilst Henderson & Henderson (1967: 478) define protandry as the ‘.. . con-
dition of hermaphrodite plants and animals where male elements mature and
are shed before female elements mature’.

A necessary prerequisite for the hypothesis of protandrism is, therefore,
that the forms involved are simultaneous hermaphrodites. To prove this it
would be necessary to study the soft parts of an ammonite, an obvious
impossibility.

Whilst hermaphroditism is virtually unknown among extant Cephalopoda,
having been recorded from but a single specimen of Octopus vulgaris (Pickford
1947: 522), it is known to occur in many species of Bivalvia and Gastropoda.
Indeed, a tendency towards protandrism is frequent in many simultaneous herm-
aphrodites. Amongst the Bivalvia simultaneous hermaphroditism is known in
Pecten, Chlamys, Cardium, Teredo, Poromya, etc. Not all species of these genera
are, however, hermaphroditic. Thus Cardium edule is dioecious; whilst in some
simultaneous hermaphrodites, e.g. Teredo diegensis, a certain number of young
males never change sex and must therefore be regarded as true males.

In the Gastropoda simultaneous hermaphroditism is well known amongst
the opisthobranchs and the pulmonates, as well as occurring to a limited extent
in the prosobranchs, e.g. Diodora, Puncturella, Patella, etc. Whilst Diodora
is predominantly dioecious, Bacci (1947) has shown about 12 per cent to be
protandrous hermaphrodites. Orton (1920) claimed that 90 per cent of a popu-
lation of Patella vulgata changed sex from male to female. Orton et al. (1956)
found small specimens of P. vu/gata (16-25 mm shell length) to be 90 per cent
males; those with shells 40 mm long were male and female in equal proportions,
whilst those with a shell length of 60 mm were 60-70 per cent females. With
the exception of the well-known protandrous hermaphroditism of Crepidula
(Coe 1936), little is known of the conditions in the other hermaphroditic
prosobranchs, beyond the fact that most seem to be protandrous consecutive
hermaphrodites.

It can be seen, therefore, that protandrous hermaphroditism, and conse-
quently also simultaneous hermaphroditism, are well known in the other
molluscan classes and it seems possible that it was at some stage equally common
in the Cephalopoda. The fact that extant cephalopods are dioecious is not,
therefore, in itself significant since, according to Fretter & Graham (1964: 128),
‘... there is some evidence for regarding the hermaphroditic state as the
primitive one, especially in the phylum Mollusca’.

176 ANNALS OF THE SOUTH AFRICAN MUSEUM

When the literature on sexual dimorphism in ammonites is studied, it is
found that the occurrence of rare, aberrant mutants supports the possibility
that some ammonites were, in fact, hermaphroditic.

Makowski (1962a: 23) considers the genus Hecticoceras to exhibit type ‘A’
dimorphism, i.e. microconchs have 5-6 whorls and macroconchs at least 7,
with a morphological hiatus of one whorl between the two dimorphs. How-
ever, of the 21 specimens studied, Makowski noted that 3 examples had ‘.. . 64
whorls each, their aperture is not quite simple, but is nearing that of the growth
stages of large forms and provided with small broad lappets. However, the
spiral pattern followed is that of large forms, completely different from the
spiral in the small forms. Hence, they should be regarded as large forms whose
growth halted at the stage with about 64 whorls.’ Makowski (1962a) makes
no mention of whether he is dealing with a single species, and thus the true
value of the observation is lost. If, however, the Hecticoceras in question were
all referable to a single species, then this combination of male (lappets) and
female (spiral form) characteristics would seem to suggest hermaphroditism.

Cope (1967: 53) recorded 4 specimens of Pectinatites (Virgatosphinctoides)
reisiformis densicostatus Cope which he considered to be ‘intersexual’. Thus,
‘... One specimen is intermediate in size between macroconch and microconch
and has rib density of a typical microconch up to a diameter of 30 mm. There-
after it becomes more finely ribbed, and is intermediate between macroconch
and microconch in rib density’.

‘At a diameter of 94 mm a horn is developed, and beyond this there is about
three-eights of a whorl of coarsely ribbed shell with sculpture similar to the
outer whorls of a macroconch, but bearing four further horns... in addition
to the above specimen which is absolutely intermediate in character between
macroconch and microconch, three other specimens show a slight degree of
intersexuality. These three specimens are apparently normal macroconchs to
judge by their size, rib density and sculpture. They do, however, develop a type
of horn in the later stages of development; this appears at a diameter of 140-150
mm and is unlike the true microconch horn in that it is developed from a single
rib, has negligible ventral projection but projects laterally some distance down
the whorl side. In addition, the diameter at which these structures are developed
is much greater than that at which the true horn of the microconch occurs.’

Whilst the intermediate size of the mutants described by Cope (1967) need
not necessarily be significant in view of the possible size overlap between
dimorphs, as was noted by Cobban (1969: 9) in his study of Scaphites leei and
S. hippocrepis, the typical female-type ribbing, associated with male-type
ornament is, and must surely be, interpreted not as intersexuality but rather
as bisexuality.

It is suggested that these rare observations of apparent bisexuality, and thus
hermaphroditism, support the evidence provided by the shells of sexually
dimorphic ammonites that some ammonites changed sex from a younger male
phase to an older female state, and thus provide the first examples of protandrism

REVISION OF LATE VALANGINIAN CEPHALOPODA 177

within the class Cephalopoda. It should be noted, however, that by analogy
with extant Mollusca, not all species of even a single genus need be protandrous,
nor even hermaphroditic.

Little is known of the exact reasons for protandric changes within those
molluscs which are simultantous hermaphrodites. Pellegrini (1948), in his study
of Patella coerulea, concluded that the change of sex was restricted to the
resting period between successive breeding seasons and liable to affect animals
of any age. Fretter & Graham (1964) consider the change to take place in
such a way as to render the animals one sex early in the breeding season, and
of the other sex later, or there may have been a winter pause between the two
phases. According to Barnes (1968: 309) the sex of the older individuals is
influenced at least partly by the presence or absence of other sexes in teh
association. Thus, in Crepidula ‘... young specimens are always males. This
initial male phase is followed by a period of transition in which the male
reproductive tract degenerates; the animal now develops into a female or
another male. ... An older male will remain male as long as it is attached to a
female. If such a male is removed or isolated it will develop into a female. The
presence of a large number of males influences certain of the males to become
females. When the individual once becomes female, it remains in that state.’

The work of Gould (1919, 1947) and Coe (1938a, 1938b, 1944, 1948)
suggests that the transition from male to female occurs at different times in
different individuals, indicating sex changes to be influenced by other animals
in the chain, and by external stimuli.

When immature limpets are cultured in association with mature females,
the great majority assume the functional male phase. Gould (1919, 1952)
presented evidence, later supported by Coe (1953), to show that the formation
and maintenance of the male phase is influenced by a substance or substances
secreted in the water by mature females. Not all young males react in the same
way to the mating stimulus.

SEXUAL DIMORPHISM IN OLCOSTEPHANUS

Within the Uitenhage olcostephanid fauna, sexual dimorphism is very
apparent due to the unusually large size attained by the macroconch forms,
some of which exceed 300 mm in diameter, whereas the largest undoubted
microconch so far recorded from these beds is only slightly more than 100 mm
in diameter, with the average far less. As recognized within many of the Euro-
pean Jurassic faunas, there are two distinct size groups (Fig. 21)—small forms
with lappets and large forms with simple peristomes. However, these two distinct
size groups comprise three morphological components (see Figs 22-23). There
are small forms, both with and without parabolae, which bear lateral lappets
and represent microconchs; there are moderately large strongly inflated forms,
falling both into the microconch and macroconch size groups, both with and
without parabolae and invariably without the peristome preserved; and,

178 ANNALS OF THE SOUTH AFRICAN MUSEUM

No. of specimens ————

100 200 300 mm
Diameter ————»—

Fig. 21. Size-frequency histogram of the Uitenhage Olcostephanus fauna, showing the double
peak corresponding to microconch and macroconch dimorphs.

finally, there are gigantic, generally strongly inflated forms lacking parabolae
and with simple peristomes representing mature macroconchs. The inter-
mediate groups have one feature in common; they are all inflated beyond the
limits of microconchs and hence must be considered immature macroconchs.
However, many of them differ from mature macroconchs in possessing para-
bolae, features which never occur on the outer whorls, except in the form of the
simple peristome, of mature female forms. That some macroconchs had para-
bolae on their earliest whorls is revealed by a gigantic phragmocone of O. rogersi
(Kitchin) (Fig. 71A—D), corresponding to a diameter of approximately 200 mm,
which shows the impression on an inner whorl, corresponding to a diameter
of about 60 mm, of a prominent parabola. Parabolae would seem, therefore,
to be restricted to the inner whorls of certain macroconchs, and to certain
microconchs. Whether parabolae are the product of male genetic control alone,
thereby providing evidence of protandry in Olcostephanus, is uncertain in view
of the slight but significant morphological differences between the inner whorls
of macroconch forms and the microconch dimorph beyond a diameter of 20 mm.
As concerns O. baini (Sharpe) dimorphs, the macroconch is already distinguish-
able at 30 mm diameter from the microconch by its slightly more numerous
secondaries, with invariably 3 secondaries per bulla as against the 2-3 of the
microconch. This is in accord with the observations of Cope (1967) on dimor-

DIAMETER : WIDTH RATIO ———»

2.60

2.20

1.80

1.40

REVISION OF LATE VALANGINIAN CEPHALOPODA 179

SIZE LIMIT OF MATURE _...., i
MICROCONCHS a

“SIZE LIMIT OF
; CONSTRICTED FORMS

=| ee

—

—
—
—
SS Ss See

100 200 300mm
DIAMETER ———»

Fig. 22. Diameter/inflation plot for the unconstricted individuals of Olcostephanus from the
Sundays River Formation. Squares = unconstricted forms; open triangles = unconstricted
forms with lappets; circles = unconstricted forms with simple peristome.

phism in the Upper Jurassic genus Pectinatites, and also those of Riccardi et al.
(1971) on Olcostephanus, that the inner whorls of the macroconch forms tend
to be slightly more densely ribbed than the microconch dimorph.

The largest diameter at which parabolae have been observed is approxi-
mately 120 mm diameter, whilst an immature macroconch of O. baini (Sharpe),
recognizable by its denser ribbing, shows a noticeable increase in inflation
immediately after a parabola at 60 mm diameter (Fig. 151B-D).

Within the Uitenhage fauna there is no size overlap between corresponding
macroconchs and microconchs.

THE PERISTOME IN OLCOSTEPHANUS

The commonest modification to the microconch aperture is the develop-
ment of lateral lappets although, as already noted, in some ammonites this
takes the form of a rostrum, a ventral lappet, or a ventral horn.

SIZE LIMIT OF MATURE MACROCONCHS

DIAMETER : WIDTH RATIO ———e

180 ANNALS OF THE SOUTH AFRICAN MUSEUM

SIZE LIMIT OF MATURE

i MACROCONCHS

2.60

SIZE LIMIT OF UNCONSTRICTED FORMS

2.20

{| SIZE LIMIT OF
(2 == MATURE
“———e———_ MICROCONCHS

1,80

— ~~
——

1.40

=—
=
—
—
—
=—_—
=—
=—

—_——
=—_—

—
oe
—
—
a
—
SS
_—— ee

1.00

(0) 100 200 300

DIAMETER ———&

Fig. 23. Diameter/inflation plot for the constricted individuals of Olcostephanus from the
Sundays River Formation. Dots = constricted forms; closed triangles = constricted forms
with lappets.

The function of these apertural extensions has long been a subject of
contention. Thus, Cope (1967: 17) considered the ventral horn in Pectinatites
to possibly have assisted in copulation by housing the spadix. According to
Bidder (in Westermann 1971), however, such a horn was more likely a median
glandular modification equivalent to Van der Hoeven’s organ. Lappets have
even been compared with the claspers of insects, but according to Arkell (in
Arkell et al. 1957: L92) ‘... their only conceivable function seems to be pro-
tective. In planulate and sphaerocone shells Westermann (1954) has noted that
the microconchs frequently have exceptionally large, converging, lateral lappets,
which caused the aperture to become occluded. According to Westermann (1971)
such ‘... occluded apertures are obviously “‘specialized’’ features preventing
macrophagous predation’. None the less, he was led to conclude that ‘. . . the
function of the diverse apertural shapes, however, remains unknown’. It seems

REVISION OF LATE VALANGINIAN CEPHALOPODA 181

to the writer most reasonable to interpret them as display characters, designed
to attract females.

Lateral lappets are preserved in a number of examples of Olcostephanus
from the Uitenhage Group, usually as internal moulds, although in a single
specimen (Fig. 131A-—B) the peristome is preserved as recrystallized test. In
this microconch the peristome comprises a deep, slightly flexuous constriction,
bordered adapically and adorally by prominent parabolic ribs. The adapical
rib is prominently flared, whilst the adoral rib is associated with well-developed
lateral lappets. On this same specimen, SAM-—PCU1527, the outer whorl is
seen to bear a parabola which takes the form of a prominent, deep, oblique
constriction bordered by strongly developed parabolic ribs, the adapical rib
being more strongly flared than that to the anterior. The parabola truncates
ribbing adapically, but is parallel to the adoral ribbing. That such parabolae
are associated with halts in growth is evidenced by the change in ribbing direc-
tion adorally, and often by a distinct change in inflation immediately after
such a feature. Such parabolae are thus virtually identical to the peristome
and must surely have an identical mode of formation. However, according to
Arkell (in Arkell et al. 1957: L93) ‘...they [parabolae] are not, however, the
same as the peristome of the adult shell, for often no such constriction or other
features may be found at the end of the adult body chamber’. In Olcostephanus
this is never the case, and it seems inconceivable that they owe their origins to
two unrelated processes. Consequently, parabolae in Olcostephanus, at least,
are interpreted as relict peristomes.

Accepting parabolae in Olcostephanus to represent relict peristomes, it is
of interest to note that in many extant gastropods, e.g. Charonia tritonis tritonis
(Linnaeus), the whorls are ornamented with varices which represent the position
of relict apertures developed during halts in growth. They would appear, there-
fore, to be absolutely analogous to the parabolae occurring in Olcostephanus.
What is especially interesting is the fact that in extant Gastropoda they are
known to be of specific importance.

HOMOEOMORPHY IN OLCOSTEPHANUS

A significant feature associated with sexual dimorphism in Olcostephanus
is a striking degree of convergence in the outer whorls of macroconch forms.
This was already noted by Makowski (1962a: 21) who wrote ‘.. . we may note
the side by side existence of large forms (macroconchs) whose last whorls and
particularly the last body chamber are identical, while their young forms differ
in section or in character of ornamentation. These differences, being not very
striking, are not taken into account in the specific delimitation of large forms
(macroconchs), they are, however, very readily discernible in small forms
(microconchs) which repeat the character of the young whorls of large forms
(macroconchs).’

It was this pitfall that led Riccardi et al. (1971) to regard O. schenki (Oppel)

182 ANNALS OF THE SOUTH AFRICAN MUSEUM

(= O. baini 2), a species with prominent parabolae, as merely the inner whorls
of the large O. atherstoni macroconch, whilst their microconch forms of
O. atherstoni, viz. O. psilostomus Neumayr & Uhlig, O. wilmanae (Kitchin),
and O. midas (Leanza), do not possess such features.

With regard to convergence within macroconch forms of Olcostephanus,
Spath (1930: 143) noted that °... examples like those figured by Burckhardt or
by Bése from Mexico as Astieria cfr. atherstoni and A. ex. aff. atherstoni repre-
sent the outer whorls of Olcostephanus of the astierianus—filosus group such as
are common in the south of France’ and that ‘...it is probable that in each
area that had its Olcostephanus fauna there were developed ‘“‘atherstoni” forms
which thus do not constitute a true species but are merely homoeomorphous
local variants of the common root-stock’ (Spath 1930: 34).

This convergence is very evident in the macroconch forms of the Uitenhage
olcostephanid fauna. Thus, the macroconch forms of O. atherstoni (Sharpe)
and O. baini (Sharpe) differ only in degree of inflation, whilst with a broad
specific interpretation it would be possible to group most Olcostephanus macro-
conchs within a single species. By far the most important factor in the matching
of sexual dimorphs is a close similarity between the inner whorls of the macro-
conch and the microconch dimorph. In those forms studied by Makowski
(1962a) there was complete identity. In Olcostephanus, however, as noted by
Riccardi et al. (1971: 96), identity of ornament is restricted to the nucleus under
20 mm diameter, with the inner whorls of the O. baini macroconch being slightly
more densely ribbed at 30 mm diameter than the microconch at 50 mm diameter.

The fact that three morphological types, i.e. the microconch, the macro-
conch, and the inner whorls of the macroconch, may be distinguished within
a single species has led to a proliferation of names, the majority endemic, which
together with an almost complete disregard for intraspecific variation has led
to taxonomic confusion.

DESCRIPTION OF THE SUNDAYS RIVER SPECIES OF OLCOSTEPHANUS

Olcostephanus (Olcostephanus) atherstoni (Sharpe, 1856)
Figs 9, 19, 24-26, 27A—D, 28-43, 55, 118, 143C—D, 151A

Microconch (6)

Ammonites astieri Pictet & Campiche (non d’Orbigny), 1860: 298, pl. 43 (figs 1, 3 only).

Olcostephanus psilostomus Neumayr & Uhlig, 1881: 149, pl. 32 (fig. 2).

Astieria aff. psilostoma (Neumayr & Uhlig) von Koenen, 1902: 151, pl. 54 (fig. 21).

Astieria psilostoma (Neumayr & Uhlig) von Koenen, 1902: 151. Baumberger, 1907: 35,
pl. 24 (fig. 6), pl. 21 (fig. 4), figs 111-113.

Astieria atherstoni (Sharpe) Baumberger, 1907: 39, pl. 21 (fig. 3), pl. 24 (figs 2, 5 only).

Astieria leptoplana Baumberger, 1908: 9, pl. 28 (fig. 2 only).

Holcostephanus wilmanae Kitchin, 1908: 195, pl. 9 (fig. 1).

Holcostephanus (Astieria) psilostomus (Neumayr & Uhlig) Wegner, 1909: 85.

Holcostephanus (Astieria) psilostomus var. picteti Wegner, 1909: 85.

Holcostephanus (Astieria) psilostomus var. wilmanae Kitchin, Wegner, 1909: 86.

Holcostephanus (Astieria) psilostomus var. koeneni Wegner, 1909: 86.

REVISION OF LATE VALANGINIAN CEPHALOPODA 183

Fig. 24. Olcostephanus (Olcostephanus) atherstoni (Sharpe) (2). Lateral view of the holotype,
BM-C32202, x 1. Photo W. J. Kennedy.

184 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 25. Olcostephanus (Olcostephanus) atherstoni (Sharpe) (2). Ventral view of the holotype,
BM-C32202. x 1. Photo W. J. Kennedy.

185

REVISION OF LATE VALANGINIAN CEPHALOPODA

‘py'O X ‘ouWO}sIIod payors
-U09 PUL UONLPUL JO 9y¥1 JULISUOD 9JON “SZILPO-INA JO SMOIA [esUDA pur [eJoyeT “(b) (odseYg) toss4ays0 (snuvydajsoajQ) snuvydajsoz1O “97 “314

ee ee
: : Feo Spal ~ See

186 ANNALS OF THE SOUTH AFRICAN MUSEUM

ee
ok ee Peas
NSS

® .
ERS See era

oe
8 Bea SN
as

we
Ss ae

Fig. 27. A-C. Olcostephanus (Olcostephanus) atherstoni (Sharpe) (2), < 0,66. Lateral, ventral

and front views of SAM—PCU1590. D. Lateral view of a slightly crushed specimen. SAM-—

PCU1585. E-F. Olcostephanus (Olcostephanus) ?densicostatus (Wegner) sp. juv. Front and
lateral views of SAM-—PCU1612, x 1.

187

REVISION OF LATE VALANGINIAN CEPHALOPODA

‘py * ‘SIE-INVS JO SMOIA [eAQUDA puR [es9Ve'T “(4) (Odueyg) Mojssayio “JO (snunydajsor1Q) snuoydajsor]O ‘87 sf te |

188 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 29. Olcostephanus (Olcostephanus) atherstoni (Sharpe). A—-B. Lateral and front views of
SAM-PCU1532, a microconch, x 0,75. C—E. Ventral, lateral and front views of SAM-—
PCU1608, a juvenile, = 0,66 F-H. Ventral, lateral and front views of SAM—PCU1526, a
microconch, x 0,66. I-K. Front, ventral and lateral views of AM-—839, a juvenile, x 0,68.

REVISION OF LATE VALANGINIAN CEPHALOPODA 189

Fig. 30. Olcostephanus (Olcostephanus) atherstoni (Sharpe) (3). The holotype of Rogersites
otoitoides Spath, SAM-—9242. x 0,75.

Astieria psilostoma var. veneto Rodighiero, 1919: 88, pl. 9 (fig. 11).

? Astieria aff. baini (Sharpe) Bose, 1923: 76, pl. 2 (figs 3-5).

Astieria psilostoma var. crassa Roch, 1930: 315.

Astieria psilostoma var. lateumbilicata Roch, 1930: 314, pl. 16 (fig. 3).

Rogersites otoitoides Spath, 1930: 149, pl. 14 (fig. 1).

Rogersites wilmanae (Kitchin) Spath, 1930: 145, pl. 13 (fig. 3), pl. 14 (fig. 4), pl. 15 (fig. 2).

Rogersites tenuicostatus Imlay, 1937: 562, pl. 73 (figs 3-9).

Holcostephanus midas Leanza, 1944: 16, pl. 1 (fig. 1).

? Taraisites bosei Cantu Chapa, 1966: 16.

Taraisites tenuicostatus (Imlay) Cantu Chapa, 1966: 16.

Olcostephanus atherstoni (Sharpe) (¢), Riccardi et al., 1971: 91, pl. 12 (fig. 4), pl. 13 (figs 2-3).

Macroconch (Q)

Ammonites atherstoni Sharpe, 1856: 196, pl. 23 (fig. 1).

Olcostephanus atherstoni (Sharpe) Holub & Neumayr, 1882: 272. Riccardi et al., 1971: 91,
pl. 12 (fig. 3), pl. 13 (figs 1, 4 only).

Non Olcostephanus (Astieria) atherstoni (Sharpe) Pavlow & Lamplugh, 1892: 495 (= O.
(Subastieria) decipiens Spath).

Holcostephanus (Astieria) atherstoni (Sharpe) Kilian & Leerhardt, 1895: 973. Wegner, 1909:
81. Kilian, 1910: 213.

Non Holcostephanus (Astieria) atherstoni (Sharpe) Kilian, 1902: 865, pl. 57 (fig. 1)

(= ? O. ventricosus (von Koenen)).

Holcostephanus atherstoni (Sharpe) Uhlig, 1903: 132. Kitchin, 1908: 185. Collignon, 1962:
38, pl. 188 (fig. 860).

Non Holcostephanus (Astieria) cf. atherstoni (Sharpe) Karakasch, 1902: 103, pl. 1 (fig. 3)
(= O. sharpei Karakasch).

Non Holcostephanus atherstoni (Sharpe) Hatch & Corstorphine, 1909: 303, fig. 76a (= O. baini
(Sharpe)).

? Astieria cf. atherstoni (Sharpe) Baumberger, 1907: 39, pl. 23 (fig. 1), fig. 114 only.

Non Holcostephanus cf. atherstoni (Sharpe) Kitchin, 1908: 193 (= O. baini (Sharpe)).

190 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 31. Olcostephanus (Olcostephanus) atherstoni (Sharpe). A—B, E. Lateral, ventral and front

views of an immature macroconch, AM-4292, x 0,75. C—D, F. Lateral, ventral and front

views of an immature macroconch, SAM-—PCU1589, x 0,44. G—H. Ventral and lateral
views of SAM—PCU1529, a microconch, x 0,66.

REVISION OF LATE VALANGINIAN CEPHALOPODA 19]

Astieria cf. atherstoni (Sharpe) Bose, 1923: 77, pl. 3 (figs 1-2).

Rogersites curvicostatus Besairie, 1936: 141, pl. 12 (figs 7, 10), pl. 13 (fig. 8).

? Astieria aff. atherstoni (Sharpe) Riedel, 1938: 13, pl. 3 (figs 5-6), pl. 12 (fig. 3).

Rogersites prorsiradiatus Imlay, 1937: 561, pl. 17 (figs 1-7).

Taraisites neoleonense Cantu Chapa, 1966: 16.

Olcostephanus (Rogersites) atherstoni (Sharpe) Spath, 1939: 32, pl. 20 (fig. 4).

? Non Olcostephanus (Rogersites) cf. atherstoni (Sharpe) Spath, 1939: 32, pl. 20 (fig. 3)
(=? O. baini (Sharpe)).

Rogersites atherstoni (Sharpe) Tzankov, 1943: 196, pl. 8 (figs 1-2, 4 only).

Non Olcostephanus cf. O. atherstoni Baumberger (non Sharpe), Imlay & Jones, 1970: B38,
pl. 9 (figs 1-3, 6-10).

Material

18 specimens; 8 microconchs (SAM-PCU1526, 1529, SAM -9242,
BM-C32199, 32204), 7 macroconchs (SAM-PCU1585, 1589-90, 1604,
AM-4292, BM-—C32202, 47128), and 3 juveniles (SAM-PCU1608, AM-839,
4293).

Holotype

By monotypy, the original of the specimen of Ammonites atherstoni
figured by Sharpe (1856: 196, pl. 21 (fig. 1)) from the Sundays River, now in
the British Museum, BM—C32202.

Diagnosis
Dimorphic. Microconch fairly small (60-100 mm in diameter), with

peristome bearing lateral lappets. Primary ribs rursiradiate, terminating in
about 18 bullae on the final whorl from which arise bundles of 3 prorsiradiate

Fig. 32. Olcostephanus (Olcostephanus) atherstoni (Sharpe) (3). The holotype of Holcostephanus
wilmanae Kitchin, BM—C32199, a laterally crushed microconch. x 0,75.

192 ANNALS OF THE SOUTH AFRICAN MUSEUM

secondaries. There are 25-37 ribs per half-whorl. Parabolae lacking. Whorl
section apparently variable. Macroconch large (+ 200 mm in diameter),
strongly inflated, with well-rounded venter and depressed, semicircular whorl
section. Primary ribs rursiradiate, terminating in 17—25 bullae at the umbilical
shoulder from which arise bundles of 3—4 secondaries, with 1—2 intercalatories
between bundles. There are 40-50 secondary ribs per half whorl. Parabolae
absent at all growth stages. Peristome simple.

Description

Microconch (3): the shell is rather small, with lappets present at diameters
from 60-100 mm, and comprises somewhat inflated to rather compressed
cadicones, involute up to the umbilical bullae so that about 75 per cent of the
previous whorl is covered. The shell becomes slightly more evolute as the
umbilical seam egresses on the adoral portion of the body chamber. About
18 rursiradiate primary ribs terminate in prominent bullae on the umbilical
shoulder of the final whorl, from which fasciculate bundles of commonly three,
rarely only two, secondary ribs arise, generally with an intercalated rib between
bundles. The coarse secondaries are prorsiradiate, recurving slightly so as to
cross the venter transversely. There may be a slight inflexion of the secondaries
as they cross the siphonal line. The umbilical wall is steep, with a subrounded
umbilical shoulder. The whorl section is rather variable. Whilst parabolae
are lacking on the phragmocone, the peristome is provided with a typical
parabola, viz. a deep, oblique constriction bordered both adorally and adapically
by prominent ribs, the adapical one of which is prominently flared. The adoral
rib is provided with well-developed, slightly converging lateral lappets. There
are between 8-10 secondaries per 3 bullae on the final whorl with between
7 and 9 secondaries within a 30 mm distance along the venter.

Macroconch (@): shell large, with peristome preserved at 200 mm diameter ;
strongly inflated, cadicone, involute up to the umbilical bullae on the inner
whorls, with about 80 per cent of the preceding whorl being covered, but
becoming slightly evolute on the final whorl as the umbilical seam egresses, so as
to make a short distance of secondary ribbing visible below the umbilical seam.
Prominent primary ribs begin at, or close to, the umbilical seam and curve back-
wards (virguliform) to terminate in 17-25 prominent bullae on the umbilical
shoulder. Each bulla gives rise to bundles of 3-5 prorsiradiate secondaries,
generally with |—2 intercalated ribs between bundles, so that there are between
10 and 17 secondaries per 3 bullae. The secondaries recurve slightly so as to
cross the venter transversely. At diameters greater than 90 mm there are almost
invariably 4 secondaries per bulla. At 75 mm diameter there are 10 ribs per
3 bullae, with a rib spacing across the venter of 3-4 mm. The whorl section is
semicircular and rather depressed, with a broad, evenly-rounded venter. Para-
bolae are lacking at all growth stages. The peristome is seen to be preserved in
a single specimen, BM—C47128, and is simple. The whorls of the macroconch
show a steady rate of inflation. SAM-—PCU1604 is preserved as an internal

REVISION OF LATE VALANGINIAN CEPHALOPODA 193

Fig. 33. Olcostephanus (Olcostephanus) atherstoni (Sharpe) (3). The specimen figured by Spath
(1930) as Rogersites aff. wilmanae (Kitchin), BM-—C32204, a probable microconch. x 1.

mould. It is moderately large and somewhat inflated, with a rather narrow
umbilicus, and steep umbilical walls. Well-developed primaries terminate in
small but distinct bullae on the umbilical shoulder, from which arise bundles
of fine, prorsiradiate secondaries which very occasionally bifurcate at about
mid-flank, with 1-2 intercalated ribs between bundles. There are 20 umbilical
bullae on the outer whorl. Parabolae are lacking.

The following description of the holotype of O. atherstoni is based on a
plastotype supplied by M. K. Howarth: the specimen is moderately large, and
somewhat inflated, and appears to have much of the shell material preserved.
The umbilicus is narrow and deep, with convex umbilical walls and a subrounded
umbilical shoulder. Primary ribs begin at, or close to, the umbilical seam and
pass strongly backwards (rursiradiate) to the umbilical shoulder where they
terminate in small but prominent umbilical bullae, of which there are about
20 on the outer whorl. There are fewer umbilical bullae on the earlier whorls.
From the umbilical bullae arise bundles of usually 4 fine secondary ribs, rarely
3 or 5, with commonly 2 intercalated ribs between bundles, although there
may occasionally be one only. Thus, on the adoral portion of the outer whorl
there are 37 secondaries per 7 bullae, with 26 ribs within a 100 mm distance
along the venter. The outer whorl is not complete, but there were probably
in all about 100 secondaries. Portion of an earlier whorl, evident in the broken
umbilicus, shows ribbing to have been coarser at earlier growth stages, with
secondaries generally arising in bundles of 3, with an intercalated rib between
bundles. The whorl section is semicircular and depressed, with an evenly
rounded venter. The whorls show a constant rate of inflation. Although the
specimen is entirely septate, matrix on the outer whorl shows that the umbilical

194 ANNALS OF THE SOUTH AFRICAN MUSEUM

seam of the following whorl egressed markedly, and that it represented the
body whorl. There is no evidence for parabolae at any growth stage.

Specimen AM-—4293 represents a typical juvenile. In this example, there
are 17 bullae on the outer whorl, from which arise bundles of 2-3 prorsiradiate
secondaries with an intercalated rib between bundles. There are 10 ribs per
3 bullae, with 13 secondaries within a 20 mm distance along the venter.

Measurements
No. D H Wi - W/E Ui
BM-C47128 197 81 C295 L117 95 61 (31)

Bs c.-130 56 li nu a 34 (26) (9)
SAM-PCU1590 102 49 75 53 45 25 (25)

a 80 34 52 3 29 16 (20) (9)
SAM-—PCU 1604 98 48 67 1,40 35 Ee, 2A (Q1) (2)
PEM-1468/74 72 33 36 1,09 30 21 (29) (3)
SAM-—PCU1529 58 G22 hk ik 28 eee (cS)
SAM-PCU1598 66 28 ? ? 32 19 (29) (3)
SAM-—PCU1526 72 Sil 41 32 30 17 (24) (3)
SAM-9242t 94 36 47 1.30 42 26 (28) (3)
BM-C32202* 134 61 80 iil 48 27 (20) (8)

Rs 106 Sil 66 1,29 36 D322)

c. 80 40 53 132 28 WFQ)
BM-C32204¢ 44 19 24 1,26 17 10 (23) (3)

+ The holotype of Rogersites otoitoides Spath.
* Plastotype of O. atherstoni (Sharpe).
t Specimen figured by Spath (1930) as Rogersites aff. wilmanae Kitchin.

Discussion

It is hardly necessary to point out the confusion that has surrounded
Sharpe’s species since its inception in 1856. This has been due to a number of
factors. First and foremost is the marked homoeomorphy between O. atherstoni
(2) and macroconchs of other species. This, together with the comparison of
different growth stages, the failure to recognize sexual dimorphism, and the
placing of constricted forms within this species have all added to the confusion.
Thus, this name has been used for species from the Crimea (Karakasch 1902),
England (Pavlow in Pavlow & Lamplugh 1892), Mexico (Burckhardt 1906;
Bose 1923), the Swiss Jura (Baumberger 1907), Pakistan (Spath 1939), South
Africa (Kitchin 1908; Spath 1930), France (Collignon 1962), and Argentina
(Riccardi et al. 1971).

Riccardi et al. (1971), in describing forms of Olcostephanus from the Lower.
Cretaceous of west-central Argentina, assigned their entire olcostephanid fauna
*...to the almost cosmopolitan Olcostephanus atherstoni (Sharpe)’. They place
into synonymy with Sharpe’s holotype, which they take to represent a macro-
conch, the following species: O. schenki (Oppel), O. sudandina (Windhausen),

REVISION OF LATE VALANGINIAN CEPHALOPODA 195

Fig. 34. Olcostephanus (Olcostephanus) atherstoni (Sharpe) (3).
The holotype of Rogersites tenuicostatus Imlay from the Taraises
Formation of northern Mexico (after Imlay 1937). x 1.

and with reservation O. curacoensis (Weaver) and O. sublaevis Spath. The
microconch is taken to be represented by the synonymous species O. psilostomus
Neumayr & Uhlig, O. wilmanae (Kitchin), and O. midas (Leanza). Moreover,
these authors hint that O. otoitoides (Spath), O. wynnei Spath, O. baini (Sharpe),
O. baini var. ambikyi (Besairie), O. auritus (Leanza), O. salinarius Spath,
O. sphaeroidalis (Spath), O. glaucus Spath, O. rigidus (Baumberger), O. lepto-
planus (Baumberger), O. imbricatus (Baumberger), and O. modderensis (Kitchin)
were a group of doubtfully distinct species over which O. atherstoni had priority.
These authors have, however, been misled both by the convergence between
macroconch forms, and in neglecting the specific importance of parabolic
constrictions.
The microconch is represented, amongst South African material, by
O. otoitoides (Spath) (Fig. 30), the crushed O. wilmanae (Kitchin) (Fig. 32) and
the specimen figured by Spath (1930) as Rogersites aff. wilmanae (Fig. 33).
Other synonyms certainly include O. tenuicostatus (Imlay) (Fig. 34), O. prorsi-
radiatus (Imlay) (Fig. 35), and O. neoleonensis (Cantu Chapa) (Fig. 36), the
latter species based upon the inner whorls of O. prorsiradiatus, as well as
O. psilostomus (Pictet) (Fig. 37), O. midas (Leanza) (Fig. 38), O. leptoplanus
(Baumberger) (Fig. 39), of which Astieria psilostoma var. picteti Wegner (1909)
is a junior objective synonym, and possibly O. curacoensis (Weaver) (Fig. 40).
_ Astieria sudandina Windhausen (1931) (Fig. 41) was based upon a specimen
illustrated only in lateral view, without scale, description or locality, and is thus
a nomen nudum. According to Riccardi et al. (1971), it is a synonym of
O. atherstoni.
Maderia altiumbilicata Imlay (1938) was based upon a strongly inflated,
globose, pyritic nucleus with a strongly depressed, semilunate whorl section.

196 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 35. Olcostephanus (Olcostephanus) atherstoni (Sharpe) (2). The holotype of Rogersites
prorsiradiatus Imlay from the Upper Valanginian of the Taraises Formation of northern
Mexico (after Imlay 1937). x 1.

Fig. 36. Olcostephanus (Olcostephanus) atherstoni (Sharpe). The
holotype of Taraisites neoleonense Cantu Chapa from the Taraises
Formation of northern Mexico (after Imlay 1937). x 1.

REVISION OF LATE VALANGINIAN CEPHALOPODA 197

Fig. 37. Olcostephanus (Olcostephanus) atherstoni (Sharpe) (3). The holotype of Olcostephanus
psilostomus Neumayr & Uhlig from northern Germany (after Neumayr & Uhlig 1881). x 1.

Fig. 38. Olcostephanus (Olcostephanus) atherstoni (Sharpe) (3). The holotype of Holcoste-
phanus midas Leanza from the Upper Valanginian of Neuquén, Argentina (after Leanza
1944). x 1.

198 ANNALS OF THE SOUTH AFRICAN MUSEUM

A . a
. re a ( ee, nie ae
iol EE ers
a A 4 Lie Hh
Brae, Hee i AO: é
2 ry a OK ae
22 2 OF ¢ ard
232 toys = ©. Soe
SGA pe wget
Z SE 2 > co ’ Nes
oe ‘° CUR ‘ E
dee & E PES roy,
eee ~ . wee oy hip!
PD Fen Ae LG SS 27 5p
Vow = 8 Nive <2
wee Oy aS. a. ar
oes eeZs 4 x 4345 as
Pw Eee E83 a Se ;
PER ERE <$ *8 -$ 54 - Bi
= Ef Eek 14 : -
ROW Stes 9 3 nek Been ke
Ae pete wey =
* 8 Peete Tey HS

Fig. 39. Olcostephanus (Olcostephanus) atherstoni (Sharpe) (¢). The original of this specimen

is the type of Holcostephanus leptoplanus Baumberger, by lectotype designation herein, and

also of Holcostephanus (Astieria) psilostomus var. picteti Wegner (after Pictet & Campiche
L858) aoe

Ornament comprises 16-17 primary ribs which terminate in bullae on the
umbilical shoulder from which arise bundles of three prorsiradiate secondaries.
Parabolae are lacking, and the writer believes this species to be based upon a
nucleus of O. atherstoni.

As can be seen from Figures 42-43, Rogersites curvicostatus Besairie
(1936) is merely based upon a macroconch of O. atherstoni whilst, according
to Thieuloy (19776: 126), Astieria carpathica Jekelius (1913) is also a synonym
of Sharpe’s species.

Taraisites bosei Cantu Chapa (Fig. 9) was erected for the poorly preserved
specimen figured by Bése (1923: 76, pl. 2, figs 3-5) as Astieria aff. baini, and
made the type of the genus Jaraisites. This fragment appears to be specifically
indeterminate; it may be either O. baini or a juvenile O. atherstoni. It is best
regarded as a nomen dubium, not to be used.

Olcostephanus actinotus (Baumberger) (Fig. 44) is a rather large species,
probably an immature macroconch, with a deep umbilicus and steep walls
ornamented with 17-18 slender, rursiradiate primaries terminating in bullae.
From these commonly arise 4, occasionally 3 or 5, secondaries with 1-2 inter-
calated ribs between bundles. Parabolae are lacking. This species seems to
differ from O. atherstoni in its more compressed form with a subtrigonal whorl
section, and in its straight not sinuous secondary ribs.

Olcostephanus andartae Thieuloy (1972) (Fig. 45) has a depressed whorl
section with steep umbilical walls. On the outer whorl, 14-15 primary ribs

199

REVISION OF LATE VALANGINIAN CEPHALOPODA

"TX “(LE6T JOAROMA
Jaye) vunussiy ‘ugnbNeN WoI JOABOMA, SISUBOIDAND DIdaISp JO IdAyoJoOY 9YJ, (4) (ed4eYyg) IWosssaysy “Jo (SnuDydajsoz]Q) snuvYydajso71O ‘Op “SIA

200 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 41. Olcostephanus (Olcostephanus) cf. atherstoni (Sharpe) ().

The holotype, by monotypy, of Astieria sudandina Windhausen,

a nomen nudem since it was not described, whilst the scale and
locality are also unknown (after Windhausen 1931). x 1.

terminate in prominent rounded tubercles on the umbilical shoulder and give
rise to bundles of 3-4 coarse, prorsiradiate secondaries characterized by fre-
quent bifurcation at midflank. Parabolae are lacking. This species differs from
O. atherstoni in its coarser, more distant ribbing with frequent bifurcation and
in possessing swollen, rounded umbilical tubercles. Olcostephanus bossingaulti
(d’Orbigny) (Fig. 46), of which O. laticosta (Gerth) (Fig. 47) is merely based
upon a juvenile, is an Hauterivian species which is very close to O. andartae,
but seems to lack bifurcating secondaries.

‘Simbirskites’ araucanus Leanza (1957) is a moderately large, fairly evolute
species of Olcostephanus, with inclined umbilical walls ornamented with about
20-23 prorsiradiate primaries per whorl. These terminate in prominent umbilical
bullae which generally give rise to three radial, to slightly prorsiradiate
secondaries, frequently with an intercalated rib between bundles. There are about
seventy secondary ribs per whorl, a short distance of which are exposed in the
umbilicus, beneath the umbilical seam. Parabolae lacking. This species differs
from the microconch of O. atherstoni in its more evolute form, with sloping
umbilical walls and prorsiradiate primaries, and should be assigned to the
subgenus Subastieria. It seems likely that the Sardinian O. (Subastieria) nicklesi
Wiedmann & Dieni (Fig. 48) is merely based upon juveniles of this species.

REVISION OF LATE VALANGINIAN CEPHALOPODA 201

Fig. 42. Olcostephanus (Olcostephanus) atherstoni (Sharpe) (@). Lateral view of the holotype
of Rogersites curvicostatus Besairie, in the collections of the University of Paris. x 1.

202 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 43. Olcostephanus (Olcostephanus) atherstoni (Sharpe) (2). Front view of the holotype of
Rogersites curvicostatus Besairie, in the collections of the University of Paris. x 1.

203

REVISION OF LATE VALANGINIAN CEPHALOPODA

ORs
Ps

acre:
POURED

Fig. 44. Olcostephanus (Olcostephanus) actinotus (Baumberger) (2), from the Swiss Jura
(after Baumberger, 1908). x 1.

loy. The paratype, possibly a macro-

ieu
conch, from the Lower Hauterivian of Rottier (Dréme) (after Thieuloy 1972). x 1.

Fig. 45. Olcostephanus (Olcostephanus) andartae Th

204 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 46. Olcostephanus (Olcostephanus) bossingaulti (d’Orbigny) (2). The holotype, by mono-
typy, from Colombia (after d’Orbigny 18425). x 1.

Fig. 47. Olcostephanus (Olcostephanus) bossingaulti (d’Orbigny). The holotype of Astieria
laticosta Gerth from Neuquén, Argentina (after Gerth 1925). x 1.

REVISION OF LATE VALANGINIAN CEPHALOPODA 205

Fig. 48. Olcostephanus (Subastieria) nicklesi Wiedmann & Dieni. The holotype and front view
of a paratype from the Upper Valanginian of Sardinia (after Wiedmann & Dieni 1968). x 1.

Fig. 49. Olcostephanus (Olcostephanus) bernardensis (Lory) (2). A cast of the holotype
MNHP-R3111, from the Middle Hauterivian of the Hautes-Alpes. x 1.

206 ANNALS OF THE SOUTH AFRICAN MUSEUM

A cast of the holotype of Olcostephanus bernardensis (Lory) (Fig. 49)* in
the Natural History Museum, Paris, with the number MNHP-R3111, shows
this species to be moderately involute (umbilicus 28% of the diameter), with a
strongly compressed whorl section (W/H = 0,75). On the outer whorl, nineteen
rursiradiate primaries terminate in bullae which give rise to bundles of prorsi-
radiate secondary ribs. The secondaries sometimes bifurcate on the flanks so
that there are about 115 ribs across the venter of the final whorl. Parabolae
are lacking and the peristome appears to be simple. This species seems to differ
from the O. atherstoni macroconch in being somewhat more densely ribbed,
with more frequent bifurcation of secondaries.

‘Maderia’ cupidinensis Imlay (1938) was based upon a pyritic nucleus with a
strongly depressed, semilunate whorl section. Between 20 and 22 primary ribs
terminate in umbilical bullae from which arise 2-3 slightly prorsiradiate
secondaries. Parabolae are lacking. This species seems to differ from
O. atherstoni in the shape of the whorl section, and in being less inflated. It
closely approaches O. andartae Thieuloy.

Fig. 50. Olcostephanus (Olcostephanus) crassus (Zwierzycki)
(2). The holotype, by monotypy, from the Tendaguru
Formation of Tanzania (after Zwierzycki 1914). x 1.

Olcostephanus crassus (Zwierzyck1) (Fig. 50) is based upon a poorly pre-
served macroconch which approaches O. atherstoni, but should be regarded as
specifically indeterminate, a nomen dubium.

Olcostephanus discoideus Imlay (Fig. 51) is a probable macroconch with a
compressed ovate whorl section, the compression seemingly enhanced by
crushing. About 20 primaries terminate in small bullae on the umbilical shoulder
from which arise 3, rarely 2 or 4, radial secondaries with 1—2 intercalated ribs

* This figure and various others are not up to our usual standard; some are from published
illustrations and others cannot be replaced—Ione Rudner, Editor.

REVISION OF LATE VALANGINIAN CEPHALOPODA 207

Fig. 51 Olcostephanus (Olcostephanus) discoideus
Imlay (2). The holotype from the Taraises Formation
of northern Mexico. (after Imlay 1938). x 1.

between bundles. On the adoral three-quarters of the outer whorl, the adapical
rib of most bundles bifurcates. Its strongly compressed form separates Imlay’s
(1938) species from O. atherstoni.

Olcostephanus irregularis (Wegner) (Fig. 52) is a strongly compressed
(?crushed), moderately evolute form with steep umbilical walls. About fifteen
umbilical bullae per half whorl give rise to bundles of commonly three, slightly
prorsiradiate secondaries which frequently bifurcate near midflank. There are
occasional intercalatories between bundles. Parabolae lacking. This species
differs from O. atherstoni in its more compressed form, more numerous
umbilical bullae and frequent bifurcation of secondaries.

A cast of Olcostephanus lamberti (Kilian) (Fig. 53), MNHP-R3110, shows
the original to have been somewhat eroded and slightly distorted. The whorl
section is distinctly compressed, elliptical, and the umbilicus fairly wide with
convex, rather sloping, umbilical walls. There are thirteen slightly rursiradiate,
almost radial, primary ribs per whorl which terminate in rather prominent
bullae at the umbilical shoulder. From these arise bundles of 3—4 rather coarse,
prorsiradiate secondaries with intercalated ribs between bundles. The secondary
ribs are about as wide as the interspaces and there are 65—70 on the outer whorl.

208 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 52. Olcostephanus (Olcostephanus) irregularis (Wegner)
(2). The holotype, by monotypy, from the Lower Hauterivian
of Marignac (Dréme) (after Wegner 1909). x 1.

Parabolae are lacking but there is a prominent apertural constriction. This
species resembles O. atherstoni but may be distinguished by its more compressed
form, fewer umbilical bullae and its younger (mid-Hauterivian) age.

Olcostephanus fascigerus Spath (Figs 92-97) differs from O. atherstoni
in its finer, thread-like secondary ribs with more intercalatories between bundles,
its more prominent bullae, and its constant rate of inflation which gives Spath’s
(1939) species a cylindrical aspect in ventral view.

The microconch of O. baini baini (Sharpe) (Fig. 114) differs from that of
O. atherstoni in being smaller with somewhat fewer secondary ribs per whorl,
and in possessing parabolae. The O. baini var. sphaeroidalis (Spath) microconch
(Figs 145-146, 149) differs from O. atherstoni (3) in possessing parabolae.

Within the Uitenhage fauna, O. atherstoni macroconchs are rather rare
and only five further specimens, one of which is fragmentary, are without
hesitation assigned to Sharpe’s species. However, very common are large
macroconchs which differ from the holotype of O. atherstoni in the greater
inflation of their middle whorls and their more depressed whorl section
(Fig. 118). Whilst the differences may appear slight, the holotype of O. baini

REVISION OF LATE VALANGINIAN CEPHALOPODA 209

Fig. 53. Olcostephanus (Olcostephanus) lamberti (Kilian). A cast of the holotype, MNHP-—
R3110, from the Middle Hauterivian of Montbrand, Hautes-Alpes. x 1.

var. sphaeroidalis (Spath) shows that in addition the inner whorls of these more
inflated forms bear parabolae. As such, the similarities merely provide further
evidence of the perturbing homoeomorphy between adult macroconchs of
Olcostephanus.

Occurrence

Olcostephanus atherstoni is a widely distributed species which is at present
recorded from South Africa, Madagascar, Argentina, northern Mexico,
Morocco, France, Austria, Switzerland and Germany but, rather curiously,
not yet from Pakistan.

Olcostephanus (Olcostephanus) densicostatus (Wegner, 1909)
Figs 27E-F, 54-55

Microconch (3)

? Olcostephanus salinarius Spath, 1939: 13, pl. 1 (figs 1-3, 6-7 only), pl. 2 (fig. 5), pl. 19 (fig. 4),
pl. 20 (fig. 2). Fatmi, 1977: 266, pl. 1 (figs 5-6), pl. 2 (fig. 4), pl. 3 (fig. 1).

? Holcostephanus auritus Leanza, 1944: 18, pl. 2 (fig. 1).

? Holcostephanus busnardoi Collignon, 1962: 43, pl. 191 (fig. 868).

210 ANNALS OF THE SOUTH AFRICAN MUSEUM

Macroconch (Q)

Non Holcostephanus atherstoni Sharpe var. nov., Kilian, 1902: 865, pl. 57 (fig. 1) (= ? O. ventri-
cosus (von Koenen)).

Holcostephanus (Astieria) atherstoni var. densicostata Wegner, 1909: 82, pl. 16 (fig. 3).

? Astieria multistriata Zwierzycki, 1914: 53, pl. 6 (figs 6-9, 16).

? Rogersites sakalavensis Besairie, 1936: 139, pl. 13 (figs 10-12).

? Rogersites spathi Besairie, 1936: 140, pl. 12 (figs 1-2).

? Rogersites filifer Imlay, 1937: 559, pl. 73 (figs 1-2), pl. 74 (figs 4-5).

Olcostephanus densicostatus (Wegner) Spath, 1939: 27.

Rogersites atherstoni var. densicostatus (Wegner) Tzankov, 1943: 197, pl. 9 (figs 1-3).

Material

A single juvenile, SAM-—PCU1612, without locality data but presumably
from the Sundays River Formation.

Holotype

By lectotype designation herein, the original of the specimen of Hol-
costephanus (Astieria) atherstoni var. densicostata figured by Wegner (1909: 82,
pl. 16 (fig. 3)) (Fig. 54) from Escragnolles (Alpes Maritimes).

Diagnosis ,

A species resembling O. atherstoni (Sharpe) but with finer, more numerous
secondaries (Fig. 55). In the presumed microconch there are between 54 and 75
secondaries per half whorl, with well in excess of 60 per half whorl in the
macroconch.

Fig. 54. Olcostephanus (Olcostephanus) densicostatus

(Wegner). The holotype, by lectotype designation herein,

from the Lower Hauterivian of Escragnolles (after Wegner
1909). x 1.

REVISION OF LATE VALANGINIAN CEPHALOPODA 211

Zé
O.densicostatus :

No. of ribs per half whorl

50 100 150mm

Diameter

Fig. 55. Plot of rib density versus diameter for microconchs of O. atherstoni and the
O. densicostatus plexus. 1 = O. salinarius var. crassa Spath, 2 = O. midas (Leanza),
3 = O. densicostatus (a probable macroconch), 4 = O. salinarius Spath (the holotype),
5 = O. salinarius var. involuta Spath, 6 = O. salinarius var. obesa Spath, 7 = O. salinarius
sp. juv. (in Spath 1939), 8 = O. salinarius sp. juv. (in Spath 1939), 9 = O. auritus (Leanza),
10 = O. atherstoni (3) (in Riccardi et al. 1971), 11 = O. aff. wilmanae (in Spath 1930),
12 = SAM-PCU1526, 13 =O. wilmanae (Kitchin), 14 =O. otoitoides (Spath),
15 = O. tenuicostatus (Imlay), 16 = O. prorsiradiatus sp. juv. (in Imlay 1937), 17 = O. lepto-
planus (in Pictet & Campiche 1860).

Description

A single juvenile (Fig. 27E-F), preserved as a ferruginous internal mould,
is referred, with reservations, to this species. The umbilicus is rather narrow
and ornamented with about 18 bullae on the outer whorl. There are about 70
fine secondary ribs on the outer whorl (at a comparable diameter a juvenile
O. atherstoni has only 60) but their connections with the umbilical bullae are
indistinct. The whorl section is semicircular.

Discussion

_ The widespread occurrence of forms closely resembling O. atherstoni
(Sharpe) but differing in the possession of denser secondary ribbing would
seem to justify Spath’s (1939: 27) elevation of Wegner’s (1909) variety to specific
rank. In his original description, Wegner also included the juvenile figured
by Kilian (1902, pl. 57 (fig. 1)) into this species but Kilian’s specimen shows a
prominent parabola and is excluded from O. densicostatus as herein interpreted.

ANNALS OF THE SOUTH AFRICAN MUSEUM

212

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REVISION OF LATE VALANGINIAN CEPHALOPODA 213

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te,

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i,
a

Fig. 57. Olcostephanus (Olcostephanus) sakalavensis (Besairie). A hypotype
in the collections of the University of Paris, from Ambiky, Madagascar.
Kel:

Olcostephanus sakalavensis (Besairie) (Figs 56-57) is a strongly inflated
macroconch species with a very depressed whorl section. The steep umbilical
walls are ornamented with about 12 rursiradiate primaries per half whorl
which terminate in bullae on the umbilical shoulder. From these arise bundles
of 4-6, fine, prorsiradiate secondaries with intercalated ribs between bundles,
so that there are 36 secondaries across the venter per 7 umbilical bullae on the
outer whorl of the holotype. Parabolae are lacking at all growth stages. Modern
revision of the European material will probably show that O. sakalavensis is
a synonym of O. densicostatus.

Olcostephanus filifer (Imlay) (Fig. 58) was created for a large, strongly
inflated macroconch with a broadly convex venter. The umbilicus of this
species is narrow and deep, with vertical walls ornamented by 20 weak, rursi-
radiate primaries which terminate in rather weak tubercles at the umbilical
shoulder. Each tubercle gives rise to 4-5 fine, prorsiradiate secondary ribs
with 1—2 intercalated ribs between bundles. Parabolae are lacking. This species
was said to differ from O. densicostatus in its finer ribbing and more numerous
umbilical tubercles, but the differences are slight and probably do not
warrant specific separation. Olcostephanus filifer is certainly a synonym of
O. sakalavensis.

Olcostephanus spathi (Besairie) (Fig. 59) is based upon the inner whorls
of a macroconch. It is a moderately inflated species with a semilunate, depressed
whorl section. The umbilical walls are strongly convex and ornamented with
25 rursiradiate primaries which terminate in rather small umbilical bullae.
From these arise bundles of slightly prorsiradiate, almost rectiradiate, second-
aries whose connections with the umbilical bullae are indistinct. Between
bundles there are several intercalated ribs, so that there are 33 secondaries across

ANNALS OF THE SOUTH AFRICAN MUSEUM

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: Wray ri

REVISION OF LATE VALANGINIAN CEPHALOPODA

‘| X ‘Ivosesepep|
‘Kyiqury jo ueluisueyed Joddy oy} Woy o1sesog 14j0ds sajisdasoy Jo adAyo[oy OYJ, (4) (Ollesog) sisuaanjoyps (snuvYydajsoz]Q) snupyda{soz]C *6¢ “S14

216 ANNALS OF THE SOUTH AFRICAN MUSEUM

the venter per 8 umbilical bullae on the outer whorl of the holotype. Parabolae
are lacking. This species is based upon the inner whorls of O. sakalavensis and
is probably, therefore, a junior subjective synonym of O. densicostatus.

The holotype of O. salinarius Spath (Fig. 60) is a microconch, somewhat
inflated and with a rather wide umbilicus. Rursiradiate primaries terminate
in 20-24 umbilical bullae from which arise bundles of 4-5 fine, prorsiradiate
secondaries. Parabolae are lacking. In view of the side-by-side occurrence of
this species with O. sakalavensis in the Spiti Shales (Fatmi 1977) they are herein

Fig. 60. Olcostephanus (Olcostephanus) densicostatus (Wegner) (3). The
holotype of Olcostephanus salinarius Spath from the Spiti Shales of
Pakistan (after Spath 1939). x 1.

regarded as sexual dimorphs and O. salinarius is likely to become a junior
subjective synonym of O. densicostatus. According to Fatmi (1977), O. geei
Spath (Fig. 61) is a synonym of O. sakalavensis. As can be seen from Figure 62,
Holcostephanus auritus Leanza is undoubtedly a synonym of O. salinarius and
hence also probably based upon the microconch of O. densicostatus.

Olcostephanus bakeri (Imlay) (Fig. 63) is based upon an inflated macro-
conch with a semicircular whorl section and rursiradiate primary ribs which
terminate in 11-12 small, pointed umbilical bullae per whorl. From these arise
bundles of 4-5 secondaries which frequently bifurcate on the lower part of the
flanks and with 1-2 intercalated ribs between bundles. Thus, there are about
120 ribs across the venter of the outer whorl. Parabolae are lacking. This
species differs from O. densicostatus in possessing far fewer umbilical bullae
and by showing more frequent bifurcation of the secondary ribs.

REVISION OF LATE VALANGINIAN CEPHALOPODA 27

Fig. 61. Olcostephanus (Olcostephanus) sakalavensis (Besairie). The holo-
type of Olcostephanus geei Spath from the Spiti Shales of Pakistan (after
Spath 1939). x 1.-

Fig. 62. Olcostephanus (Olcostephanus) densicostatus (Wegner) (3). The holotype of Holcoste-
Phanus auritus Leanza, from the Lower Hauterivian of Argentina (after Leanza 1944). x 1.

218 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 63. Olcostephanus (Olcostephanus) bakeri (Imlay) (@).
The holotype from the Taraises Formation of northern
Mexico (after Imlay 1937). x 1.

Fig. 64. Spiticeras? balestrai (Rodighiero). The holotype of
Astieria balestrai Rodighiero, from Venice (after Rodighiero
1919) ele

REVISION OF LATE VALANGINIAN CEPHALOPODA 219

‘Astieria balestrai Rodighiero (Fig. 64) resembles the presumed micro-
conchs of O. densicostatus but frequent trifurcation of the secondary ribs
suggests it is better referred to the genus Spiticeras.

Olcostephanus busnardoi (Collignon 1962) is based upon what appears to be
a microconch showing a wide, deep umbilicus with vertical walls. Small rounded
tubercles on the umbilical shoulder give rise to bundles of 3—5 slightly prorsi-
radiate ribs, with intercalatories between bundles. Parabolae are lacking. It is
doubtful whether this species can be satisfactorily distinguished from the
suspected microconchs of O. densicostatus.

Fig. 65. Olcostephanus delicatecostatus Haas. The holotype
from Colombia (after Haas 1960). x 1.

Olcostephanus delicatecostatus Haas (Fig. 65) from the Upper Valanginian
of Colombia is characterized by ‘.. . the fineness and density of costation, there
being nearly 25 ribs on a quarter whorl. ... Some ribs bifurcate at about mid-
flank. In one individual only 3 circumumbilical tubercles of medium strength
are present. In the holotype... one or two narrow, extremely shallow con-
strictions are recognizable’ (Haas 1960: 9). The density of ribbing in Haas’s
species is comparable to that of O. densicostatus but it can be distinguished by
the irregular (?pathological) development of umbilical tubercles. What Haas
(1960) refers to as constrictions in O. delicatecostatus do not seem to be para-
bolae but merely irregularities in growth.

Olcostephanus latiflexus (Baumberger 1908) (Fig. 66) is a moderately large,
compressed form, with a wide rather shallow umbilicus and a very depressed
whorl section. About 21 rursiradiate primaries on the outer whorl terminate
in bullae from which arise 4-5 prorsiradiate secondaries with intercalated ribs
between bundles. Parabolae appear to be lacking. This species is disturbingly
close to O. salinarius and the other suspected microconchs of O. densicostatus
and, as it has priority over Wegner’s (1909) species, its affinities warrant closer
scrutiny.

220 ANNALS OF THE SOUTH AFRICAN MUSEUM

STITT Ff p>
ae. j ; A < D>

z2
v4 ¢
a 2

Fig. 66. Olcostephanus (Olcostephanus) latiflexus (Baumberger), (3). The holotype, by lecto-
type designation herein, from the Swiss Jura (after Baumberger 1908). x 1.

Fig. 67. Olcostephanus (Olcostephanus) multistriatus (Zwierzycki).
The holotype, by lectotype designation herein, from the Lower
Hauterivian of Tanzania (after Zwierzycki 1914). x 1.

REVISION OF LATE VALANGINIAN CEPHALOPODA 221

Fig. 68. Olcostephanus (Olcostephanus) rebillyi (Collignon). The holotype
from the Lower Valanginian of Madagascar. (after Collignon 1962). x 1.

Olcostephanus multistriatus (Zwierzycki) (Fig. 67) is based upon a juvenile
which does not seem to warrant specific separation from O. densicostatus.

Olcostephanus rebillyi (Collignon) (Fig. 68) is a species, allegedly from the
Lower Valanginian, which is characterized by its rounded whorl section,
relatively flat inner flanks, and by the fineness of its ornament which consists of
extremely small tubercles from which arise bundles of very fine, prorsiradiate
secondaries. Parabolae are apparently lacking. It was said to differ from
O. salinarius in its narrower umbilicus, less inflated form and more flexuous
ribs, but the differences are slight. In view of its age, however, Collignon’s
(1962) species is for the present maintained as distinct.

Olcostephanus rabei (Besairie) (Fig. 69) shows few features to distinguish
it from O. sakalavensis and may also prove to a synonym of O. densicostatus.

Occurrence

Olcostephanus densicostatus is present in the Swiss Jura and perhaps
Pakistan, Madagascar, Tanzania, South Africa, and Mexico.

Olcostephanus (Olcostephanus) rogersi (Kitchin, 1908)
Figs 70-73, 74A—B, 75A-—B, 76, 80, IS0E-G

Microconch (2)

Holcostephanus rogersi Kitchin, 1908: 201, pl. 9 (fig. 1), pl. 10 (fig. 3). Hatch & Corstorphine,
1909: 303, fig. 76c. Giovine, 1950: 39.

Holcostephanus (Astieria) rogersi Kitchin, Wegner, 1909: 88.

Rogersites rogersi (Kitchen) Spath, 1930: 147. Besairie, 1936: 141.

Rogersites crassicostatus Spath, 1930: 199.

Olcostephanus (Olcostephanus) madagascariensis (Lemoine) Fatmi, 1977: 271, pl. 5 (fig. 4).

PDs ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 69. Olcostephanus (Olcostephanus) rabei (Besairie), x 1. A—B. The holotype. C—F. Para-
types. From the Upper Valanginian of Madagascar.

REVISION OF LATE VALANGINIAN CEPHALOPODA 223

Macroconch (@)

Holcostephanus modderensis Kitchin, 1908: 202, pl. 10 (fig. 3). Giovine 1950: 39.

? Astieria imbricata Baumberger, 1908: 14, figs 123-125, pl. 26 (figs 2-3).
Holcostephanus (Astieria) modderensis Kitchin, Wegner, 1909: 89. Kilian 1910: 214.
Rogersites modderensis (Kitchin) Spath, 1924: 86. Spath, 1930: 148.

? Holcostephanus (Rogersites) leanzai Giovine, 1950: 38, pl. 2 (figs 1-3).

Rogersites kitchini Spath, 1930: 148, pl. 15 (fig. 4).

Olcostephanus modderensis (Kitchin) Riccardi et al., 1971: 90.

Olcostephanus kitchini (Spath) Riccardi et al., 1971: 90.

Material

14 specimens; 6 microconchs (SAM-—4698, 5071, 11004, SAM-—PCU1527,
LJE-989b, PEM-—1468/42), 7 macroconchs (SAM-—PCU1542, 1566, LJE—989e,
AAS-370, PEM-1465/81, BM—C47127, SAM-—5070), and 1 juvenile (AM-—4028).

Holotype

By monotypy, the original of Holcostephanus rogersi, SAM-—5071 (Fig.
70D-G), figured by Kitchin (1908: 201, pl. 9 (fig. 2)) from the Sundays River
Formation.

Diagnosis

Dimorphic. Microconch small (about 70 mm diameter), with 16 rursiradiate
primaries per whorl, terminating in prominent umbilical bullae, from which
arise invariably 3 coarse, radial secondaries, usually with an intercalated rib
between bundles. Parabolae prominent. Macroconch large (300+ mm diameter),
extremely inflated, globose. Inner whorls only with parabolae. About 18 very
prominent umbilical tubercles on the outer whorls give rise to very coarse,
radial secondaries, 3—4 per bulla, with 1-2 intercalated ribs between bundles.

Description

Microconch (3): Kitchin’s holotype, which is taken to represent a micro-
conch, is a poorly preserved internal mould, with odd patches of recrystallized
shell. Moreover, it has been slightly crushed, thereby leading Kitchin (1908)
to believe the secondary ribbing to be rursiradiate. A near perfect example of
this dimorph, LJE—989e, with the lappets preserved, shows, however, the true
characters. In this latter specimen, the whorl section is rather depressed, with
a fairly broad, evenly rounded venter, as in the holotype. The whorls are involute
up to the umbilical bullae, except on the anterior portion of the body chamber
when the umbilical seam egresses slightly to reveal a short distance of secondary
ribbing of the penultimate whorl. The umbilicus is moderately wide and deep,
with steep umbilical walls. The slope of the latter decreases slightly as the
umbilical seam egresses.

The primary ribs begin at the umbilical seam and are prominently
developed, curving backwards (rursiradiate) to sharp, prominent bullae on the
umbilical shoulder. With the egression of the umbilical seam the primary

224 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 70. Olcostephanus (Olcostephanus) rogersi (Kitchin) (3), x 0,75. A-C. Ventral, lateral

and front views of LJE-989b, with recrystallized test preserved, showing prominent umbilical

bullae, radial secondaries, parabolae and lappets. D-G. Right lateral, front, ventral and left

lateral views of the holotype, by monotypy, SAM-—5071, based upon a poorly preserved
internal mould.

REVISION OF LATE VALANGINIAN CEPHALOPODA DS

Fig. 71. Olcostephanus (Olcostephanus) rogersi (Kitchin) (2), x 0,44. A-D. Lateral, dorsal,
and ventral views, and ventral view of a plasticine mould of the internal whorls of SAM-
PCU1542. E-F. Lateral and ventral views of PEM-—1468/81, an immature macroconch.

226 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 72. Olcostephanus (Olcostephanus) rogersi (Kitchin) (2). The holotype of Rogersites
modderensis (Kitchin), BM—C32201. x 0,75.

ribs become almost radial. From the umbilical bullae, bundles of three, very
rarely only two, radial secondaries pass straight across the venter, although
on the adoral portion of the body chamber, i.e. with the egression of the
umbilical seam, they become distinctly prorsiradiate. There is a deep oblique
parabola about 180° from the peristome, the adapical rib of which is prominently
flared. The peristome is virtually identical to this parabola except that the
posterior margin is even more prominently flared, while the adoral rib bears
well-preserved lateral lappets which have a subhorizontal twist to them.

Macroconch (Q): in the slightly crushed holotype of O. modderensis (Kitchin)
(Fig. 72), which represents merely the inner whorls of the macroconch of this
species, well-developed rursiradiate primary ribs terminate in about 16 sharp
umbilical bullae on the outer whorl. These give rise to bundles of commonly 3,
occasionally 4, very coarse, radial secondaries, generally with an intercalated
rib between bundles. The secondary ribbing may become slightly prorsiradiate
as a parabola is approached. There is a very prominent, deep, oblique parabola
on the outer whorl.

The following is a description of the holotype of O. kitchini (Spath) based
on a plastotype supplied by M. K. Howarth: the shell is gigantic, extremely
inflated, with a very depressed whorl section. The umbilical seam of the outer
whorl has already begun to egress, showing it to represent part of the body
chamber. The umbilicus is narrow and very deep. The umbilical walls of the
penultimate whorl are very steep, convex, and ornamented with 17 prominent
rursiradiate primaries which begin at, or close to, the umbilical seam and
terminate in prominent bullae on the umbilical shoulder. On the final whorl

REVISION OF LATE VALANGINIAN CEPHALOPODA 227

Fig. 73. Olcostephanus (Olcostephanus) rogersi (Kitchin) (2). A—B. Ventral and lateral views

of AAS—989e, x 0,75. C—D. Lateral and front views of AAS-—370, a specimen which is only

tentatively included here, x 0,75. E-G. Lateral, front and ventral views of a juvenile in the
Albany Museum, x 1.

228 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 74. A, C. Olcostephanus (Olcostephanus) rogersi (Kitchin) (2). Lateral and ventral views
of SAM-S5070, x 0,44. B, D. Olcostephanus (Olcostephanus) baini baini (Sharpe) (Q).
Lateral and ventral views of PEM-1463/41. Note the rapid rate of inflation, x 0,44.

REVISION OF LATE VALANGINIAN CEPHALOPODA 229

Fig. 75. A-B. Olcostephanus (Olcostephanus) rogersi (Kitchin) (2). Lateral and ventral views

of a specimen in the Geological Survey, Pretoria, x 0,75. C—D. Olcostephanus sp. Lateral

and ventral views of a macroconch fragment which approaches O. saintoursi (Collignon),
but appears to lack constrictions, SAM-—PCU1562. x 0,44.

230 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 76. Olcostephanus (Olcostephanus)
rogersi (Kitchin) (3). Whorl section of
the holotype. x 1.

the bullae become rounded and swollen, and very prominent, while the primaries
weaken considerably on the umbilical wall, being almost entirely effaced on the
adoral portion of the outer whorl. Extremely coarse secondaries arise from the
umbilical tubercles and vary from almost radial to slightly prorsiradiate in
direction. There are commonly 3 secondary ribs per tubercle, with an inter-
calated rib between bundles, although on the adoral portion of the outer whorl
there are commonly 4 ribs per bundle. There is not the slightest doubt that this
species merely represents a mature growth stage of O. modderensis.

Measurements

No. D H Wi W/H_ Ue Ui
PEM-1468/81 85 32 2 2 4] 28 (33) (9)

= TS 28 5) 1,96 29 ?

“5 55 23 aT 1,61 21 8 (15)

7 38 59 [E55 25 ? (2)

Plastotype of O. modderensis 65 28 50 1,79 1 10 (15)
SAM-PCU 1542 e220 ) 155 2,06 ? ” (9)

ms NYS ? 84 ? ? y

99 c. 60 2 45 ? ? i
SAM-—5070 e170 77 119 los 85 45 (26) (2)

' c. 140 60 ? ? 65 is
LJE-989e 15 28 48 Lew Di 13 (17) )

*- 54 33 35 [52 20 122)
SAM-5071 58 24 6)5) 1,46 26 16 (28) (3)

ae 46 26 30 1,50 21 11 (24)

. 35 LS 24 1,60 18 ?
LJE-989b 59 17 30 1,76 27 16 (27) (3)

99 47 15 29 1,93 19 1123)

55 35 et 22 2,00 15 (KCAS)
Plastotype of O. kitchini 235 90 c.195 par 95 69 (29) (9)

REVISION OF LATE VALANGINIAN CEPHALOPODA Za"

Discussion

The only significant difference between the holotypes of O. modderensis
and O. rogersi is the extreme inflation of the former and consequently they are
regarded as sexual dimorphs.

In 1930, Spath, when describing Rogersites kitchini, referred to a large but
fragmentary specimen which he considered ‘... closer to the present species
(R. kitchini) than to any other described form of Rogersites’, while considering
it to have *...the general appearance of what a gigantic R. schenki may be
supposed to be like’ (Spath 1930: 149). This specimen, SAM -—S070, is here
figured (Fig. 74A, C) and can be seen to show all the characteristics of O. rogersi
(2). It is still septate at a diameter of 175 mm and thus represents merely the
inner whorls of a much larger specimen. It has rursiradiate primary ribs termi-
nating in umbilical bullae on the umbilical shoulder, from which arise 3-4
radial secondaries. At this diameter there are 14 secondary ribs per 4 bullae.
A prominent deep parabola is evident at a diameter corresponding to about
130 mm. There seems little doubt that this example represents merely a larger
growth stage than that represented by the holotype of O. modderensis.

Another gigantic phragmocone (Fig. 71A—C), approximately 220 mm in
diameter, shows an even later growth stage in the ontogeny of this species.
The secondary ribbing is still remarkably coarse, with 3-4 radial secondaries
arising from each umbilical bulla and 17 secondaries per 4 bullae on the outer
whorl. The umbilicus is narrow and very deep, with broad, almost vertical,
slightly convex umbilical walls ornamented with prominent rursiradiate
primaries. Whilst the outer whorl, admittedly not complete, shows no sign of
parabolae, an impression of an inner whorl corresponding to a diameter of
about 60 mm shows a deep, oblique parabola, and at this growth stage would
appear to be almost identical to the holotype of O. modderensis. On the adoral
portion of the outer whorl of this specimen the umbilical bullae are seen to
become more prominent, a trend which is continued in the even larger,
approximately 300 mm diameter, holotype of O. kitchini (Spath), a species which
is undoubtedly conspecific with the present form.

Kitchin (1908) noted a slight forward inclination of the secondary ribbing
on the adoral portion of the outer whorl of O. modderensis, but thought it might
have been due to crushing. However, the habit of the ribbing becoming prorsi-
radiate as it approaches the peristome, and thus also parabolae (which are
interpreted as relict peristomes), is common to many species of Olcostephanus.
Indeed, O. tenuicostatus Imlay, a synonym of O. atherstoni, was distinguished
by *... the forward inclination of the ribs near the aperture’ (Imlay 1937: 562).

_Olcostephanus rogersi is undoubtedly closely related to O. baini (Sharpe),
as 1s evidenced by the close similarity of the microconchs. Olcostephanus baini
(3) differs from O. rogersi ($) in having slightly fewer secondaries per bulla,
while the direction of ribbing is prorsiradiate and not radial as in O. rogersi.
A subtle, but distinctive difference is also seen in the nature of the whorl
sections, that of O. rogersi (3) being more depressed, with a broader, flatter

232 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 77. Olcostephanus (Olcostephanus) angusticoronatus (Imlay) (2). The holotype from the
Taraises Formation of northern Mexico (after Imlay 1938). x 1.

venter, whereas the whorl section of O. baini baini (3) is almost semicircular.
Kitchin (1908) considered the weak umbilical tubercles of O. rogersi important
in the separation of this species from O. baini but this feature is due to the fact
that the holotype is an internal mould.

Olcostephanus baini var. sphaeroidalis (Spath) (4) has a less inflated form
while the secondary ribbing is also prorsiradiate. Whilst the differences between
O. rogersi and O. baini may seem slight, the very marked differences between
the macroconch forms of these two species suggests that the separation is a
valid one.

In describing the ammonite fauna of the Taraises Formation of northern
Mexico, Imlay (1938) created the new species Valanginites angusticoronatus
(Fig. 77) for very involute, globose forms with rursiradiate primary ribs termi-
nating in 16 umbilical bullae from which radiate bundles of 3-4 ribs. There are
two prominent parabolae per whorl. This species is very close to the inner whorls
of O. rogersi macroconch from which it appears to differ in being somewhat
more involute and more finely ribbed.

Although originally assigned to the.genus Holcostephanus, “H. bachelardi
Sayn (Fig. 78) has recently been included in the synonymy of Valanginites
psaephoides (Mayer—Eymar) (Thieuloy 1977a).

Olcostephanus imbricatus (Baumberger) (Fig. 79) is a strongly inflated,
globose species with coarse almost radial secondary ribs which arise from 19-20
umbilical bullae. According to Baumberger (1908: 17), the secondaries arise
in pairs from the bullae but quickly bifurcate so that 4 ribs from each bulla
cross the venter. There are prominent parabolae on the inner whorls. This
species is very close to the macroconch of O. rogersi and this relationship
bears closer scrutiny.

REVISION OF LATE VALANGINIAN CEPHALOPODA 233

Fig. 78. Valanginites psaephoides (Mayer-
Eymar). The holotype, by monotypy, of Holco-
stephanus bachelardi Sayn allegedly from the
Barremian of France (after Sayn 1889). x 1.

Fig. 79. Olcostephanus (Olcostephanus) imbricatus (Baumberger) (2). A hypotype from the
Swiss Jura (after Baumberger 1908). x 1.

The resemblance between O. crassicostatus (Spath) (Fig. 80) and the
O. baini microconch was first noted by Kitchin (1908) when he described the
former as Holcostephanus cf. baini (Sharpe). The large number of O. baini
microconchs now available allows a better judgement of the intraspecific
variation within this dimorph, and hence a closer comparison with O. crassi-
costatus. The latter species differs from O. baini (3) in its larger adult size,
generally radial secondary ribbing although, as in most species of Olcostephanus,
this tends to become prorsiradiate near the aperture, and in commonly having
three secondary ribs arising from each bulla. In addition, the whorl section of
O. crassicostatus is strongly depressed and with a broad, flattish, gently rounded
venter. The features which separate O. crassicostatus from O. baini are all
characters typical of the O. rogersi microconch and hence O. crassicostatus

ANNALS OF THE SOUTH AFRICAN MUSEUM

234

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REVISION OF LATE VALANGINIAN CEPHALOPODA 235

is considered to be based upon a gerontic individual of O. rogersi (3). Two
other individuals, SAM-—PCU1527 and PEM-1463/42, show these same
features, the former at 72 mm diameter and with the peristome and lappets
preserved and the latter preserved as an internal mould.

‘Subastieria chanchelula (Anderson 1938) shows a depressed reniform
whorl section, with a deep, narrow umbilicus and steep umbilical walls.
Prorsiradiate primaries terminate in bullae on the umbilical shoulder from which
arise 2-3 rectiradiate secondaries with an occasional intercalated rib between
bundles. Parabolae appear to be lacking. The absence of parabolae and the
prorsiradiate primary ribs serve to distinguish this species from O. rogersi.
According to Imlay (1960) it may be a juvenile Simbirskites.

Fig. 81. Olcostephanus (Olcostephanus) radiatus Spath. The holotype from the Spiti Shales of
Pakistan (after Spath 1939). x 1.

Olcostephanus radiatus Spath (Fig. 81) is very close to O. rogersi (3) but
differs in having more umbilical bullae (twenty-four) and, in the holotype, by
the absence of parabolae.

Olcostephanus madagascariensis (Lemoine) (Fig. 14), the type of the sub-
genus Lemurostephanus, resembles the O. rogersi microconch but can be
distinguished by having distinctly sloping umbilical walls and more numerous
bullae. The specimen figured by Fatmi (1977: 271, pl. 5 (fig. 4)) is indistinguish-
able from the present material of O. rogersi (compare Fatmi’s specimen with
Figure 70A-C), and is included in the synonymy of the latter species. Olco-
stephanus mitreanus (d’Orbigny) (Fig. 17), which includes amongst its synonyms
O. detonii (Rodighiero) (Fig. 15), O. wynnei Spath (Fig. 16), and possibly the
specimen figured by Thieuloy (1977a, pl. 9, fig. 27) as O. (Lemurostephanus)
aff. sanctifirminensis Thieuloy, differs from O. rogersi ($) in being more evolute.

236 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 82. Olcostephanus (Olcostephanus) leanzai (Giovine) (2). The holotype from the Lower
Hauterivian of Neuquén, Argentina (after Giovine 1950). x 0,66.

‘Maderia’ multituberculata Imlay (1938) is based upon an inflated, globose,
pyritic nucleus with a strongly depressed, semilunate whorl section. About 26
rursiradiate primary ribs terminate in bullae on the umbilical wall and give
rise to 2-3 prorsiradiate secondary ribs. Parabolae are prominent and thinning
of the ribs across the siphonal line is insignificant. This species differs from
O. rogersi in its more numerous primary ribs and prorsiradiate secondaries. It
is probably based upon the innermost whorls of O. globosus Spath.

Olcostephanus leanzai (Giovine) (Fig. 82) is a coarsely ribbed, globose
species which was compared with O. rogersi and O. modderensis. It is unknown
whether the inner whorls of this species bear parabolae. So far as can be judged,
it differs from the O. rogersi macroconch only in the possession of more
numerous umbilical bullae but the differences are slight.

Olcostephanus saintoursi (Collignon) (Fig. 83) is too fragmentary to allow
a proper comparison, but has finer, more numerous secondaries and is
apparently from Lower Valanginian strata. The large size and strong inflation

REVISION OF LATE VALANGINIAN CEPHALOPODA O37

Fig. 83. Olcostephanus (Olcostephanus) saintoursi (Collignon) (2). The holotype from the Upper
Valanginian of Madagascar (after Collignon 1962). x 1.

of this species clearly indicate it to be an immature macroconch but topotype
material is necessary to prove its validity.

Dobrodgeiceras wilfridi (Karakasch) (Fig. 12) is another strongly inflated
form but differs from the O. rogersi macroconch in its small adult size, extremely
narrow umbilicus, and the shifting of the umbilical tubercles to a ventrolateral
position with a consequent lengthening of the primaries.

Olcostephanus laticostus (Gerth) (Fig. 47) has swollen umbilical tubercles
and lacks parabolae and hence cannot be confused with O. rogersi. It is, how-

Fig. 84. Olcostephanus (Olcostephanus) singularis (Baumberger).

The holotype, by monotypy, of Holcostephanus klaatschi Wegner,

from the Lower Hauterivian of Basses-Alpes, France (after
Wegner 1909). x 1.

238 ANNALS OF THE SOUTH AFRICAN MUSEUM

ever, close to O. klaatschi (Wegner) (Fig. 84) from which it differs in the posses-
sion of much coarser ribbing. Wegner’s (1909) species differs from O. rogersi
in being more finely and densely ribbed, lacking parabolae, and with swollen
umbilical tubercles at early growth stages. It is almost certainly merely based
upon the inner whorls of O. singularis (Baumberger).

Astieria dolioliformis Roch (Fig. 13) was erected for extremely inflated,
globose forms with a very narrow umbilicus ornamented with nineteen primaries,
each terminating in a small umbilical tubercle. Each bulla gives rise to two
secondary ribs separated by an intercalatory. As suggested by Spath (1939: 25),
this species is best assigned to the genus Valanginites.

Fig. 85. Olcostephanus (Olcostephanus) globosus Spath (2). The holotype of Olcostephanus
pachycyclus Spath from the Spiti Shales of Pakistan (after Spath 1939). x 1.

Olcostephanus pachycyclus (Folgner MS) Spath (Fig. 85) is very close to
the holotype of O. modderensis, i.e. the O. rogersi macroconch, from which it
differs only in the possession of far more umbilical bullae (twenty-three at
70 mm diameter). Fatmi (1977) has included this species in the synonymy of
O. globosus Spath (Fig. 86).

Olcostephanus drumensis (Sayn MS) (Kilian) (Fig. 87) differs from O. rogersi
in the possession of more numerous umbilical tubercles and in its denser, finer
secondary ribbing.

Olcostephanus psilostomus quadricostatus (Tzankov) (Fig. 88) was based
upon two individuals, a macroconch and a microconch. The latter is herein
selected as lectotype and seems very close to O. mitreanus (3), although it is
unclear from Tzankov’s (1943) figure if the Bulgarian form has parabolae. It
differs from the O. rogersi microconch in its smaller adult size, wider umbilicus

239

REVISION OF LATE VALANGINIAN CEPHALOPODA

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240 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 87. Olcostephanus (Olcostephanus) drumensis (Sayn MS) Kilian.
The holotype, by monotypy, from the Middle Valanginian of
Fontanil, France (after Kilian 1910). x 1.

and apparent lack of parabolae. It still remains to be demonstrated that the
macroconch which forms part of the syntype series is, in fact, conspecific with
the microconch. Since the Bulgarian microconch is certainly specifically distinct
from ‘O. psilostomus’, the writer would suggest elevating Tzankov’s (1943) |
trivial name to full specific status, i.e. O. quadricostatus (Tzankov).

Occurrence

This species is known with certainty only from South Africa and Pakistan,
but may also be present in the Swiss Jura and Argentina.

Olcostephanus (Olcostephanus) victoris Spath, 1939 (9)
Figs 89-90

Olcostephanus victoris Spath, 1939: 20, pl. 19 (fig. 7a—b). Fatmi, 1977: 268.
Material

A single specimen, LJE-991, from Addo Drift East B farm in the Uitenhage
district.
Holotype

The original of Olcostephanus victoris figured by Spath (1939: 20, pl. 19
(fig. 7a only)) (Fig. 89) from the Chichali Pass of Pakistan.
Diagnosis

A rather evolute species of Olcostephanus, probably representing the inner
whorls of a larger macroconch, with a somewhat compressed form. Primary

REVISION OF LATE VALANGINIAN CEPHALOPODA 241

Fig. 88. Olcostephanus (Olcostephanus) quadricostatus (Tzankov). The syntypesi from, Bulgaria,
of which the smaller specimen, a microconch, is selected as lectotype (after Tzankov 1943). x 1.

242 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 89. Olcostephanus (Olcostephanus) victoris Spath (2). Lateral view of the holotype and
ventral view of a paratype, from the Spiti Shales of Pakistan (after Spath 1939). x 1.

ribs rursiradiate, terminating in about 28 umbilical bullae on the outer whorl,
from which arise bundles of 3-4 fine, prorsiradiate secondaries. Whorl section
suboval, depressed. Parabolae present. Umbilical shoulder rounded, with
maximum width some way above the umbilical shoulder.

Description

A single entirely septate specimen (Fig. 90) from the Sundays River is
assignable to Spath’s species.

This example, LJE-991, has recrystallized test preserved. The shell is
somewhat inflated and rather evolute, with a strongly depressed whorl section.
At 120 mm diameter the specimen is entirely septate and thus almost certainly
represents the inner whorls of a larger probably compressed macroconch. The
umbilicus is rather wide and moderately deep, with steep umbilical walls and
a well-rounded umbilical shoulder. The venter is rather broad and flattish, but
evenly rounded.

REVISION OF LATE VALANGINIAN CEPHALOPODA 243

On the adoral one-quarter of the outer whorl there is already a distinct
egression of the umbilical seam, so that a short distance of secondary ribbing
is visible on at least the penultimate whorl. Primary ribs begin at the umbilical
seam and pass backwards (rursiradiate) to 26 rather inconspicuous bullae on
the umbilical shoulder. These in turn give rise to bundles of 3-4 fine, prorsi-
radiate secondaries, commonly with an intercalated rib between bundles. The
secondaries recurve slightly so as to cross the venter transversely. There are
25 secondaries per 6 bullae on the outer whorl, with 15 ribs within a 50 mm
distance along the venter. Maximum width is some way above the umbilical
bullae. There are two prominent parabolae on the outer whorl, with a third
partially hidden by the adoral portion of the outer whorl.

Fig. 90. Olcostephanus victoris Spath (2). Lateral, front and ventral views of LJE-991. Note
the convex flanks and the egression of the umbilical seam at a stage when the shell is still
entirely septate. x 0,44.

Measurements
No. D H Wi W/H Uo Ui
LJE-991 ELS 41 69 1,68 51 30 (26)
be 95 38 56 1,47 42 23 (24)
Me 64 30 37 123 26 16 (25)
Discussion

The Uitenhage example closely resembles Spath’s holotype. The latter
does not, however, show the characteristic egression of the umbilical seam on
the outer whorls. None the less, it represents an earlier growth stage than the
present specimen and thus may well have become more evolute with ontogeny.
Fatmi (1977) considered O. victoris a synonym of O. sakalavensis Besairie but
it differs from that species in being somewhat more evolute, with maximum
width somewhat above the umbilical tubercles and in possessing constrictions.

244 ANNALS OF THE SOUTH AFRICAN MUSEUM

Olcostephanus uitenhagensis (Kitchin) also becomes markedly evolute on
the final whorl, hence at a much later stage than O. victoris, whilst its inner
whorls are more compressed, with broader, flatter flanks. It is unknown whether
the inner whorls of O. witenhagensis bear parabolae.

Fig. 91. Olcostephanus (Olcostephanus) schafarziki (Somogyi) (2). The
holotype from Martonkat (after Somogyi 1916). x 1.

Olcostephanus schafarziki (Somogyi) (Fig. 91) would also appear to be
rather evolute at moderate diameters but differs from O. victoris in lacking
parabolae and in the frequent bifurcation of secondaries. In this respect it
closely resembles O. irregularis (Wegner), with which it may well be conspecific.

Occurrence
This species is currently known only from the Sundays River Formation
and the Spiti Shales of Pakistan.

Olcostephanus (Olcostephanus) fascigerus Spath, 1939 (Q)
Figs 92-97
Holcostephanus (Astieria) cf. convoluta von Koenen, Uhlig, 1903: 394, pl. 78 (fig. 1).

Olcostephanus fascigerus Spath, 1939: 18, pl. 4 (figs 1-3). Fatmi, 1977: 268, pl. 3 (fig. 3).
Holcostephanus uhligi Collignon, 1962: 23, fig. 827.

REVISION OF LATE VALANGINIAN CEPHALOPODA 245

Material
Four adult macroconchs, SAM-PCU1568, 1611, SAM-—5074, BM-—C7126.

Holotype

The original of the specimen figured by Spath (1939: 18, pl. 14 (figs 1-3))
and here refigured (Fig. 92) from the top bed of the middle member of the
Chichali Formation of northern Pakistan.

Diagnosis

Large inflated cadicone macroconchs (+ 150 mm diameter), with a
depressed whorl section and evenly rounded venter. Between 18 and 22 pro-
minent umbilical bullae on the final whorl give rise to 4-5 radial to prorsiradiate,
fine, flexuous, secondary ribs, occasionally bifurcating or with intercalatories,
and with 2-5 intercalated ribs between bundles. Parabolae lacking at all growth
stages and with a simple peristome in maturity.

Description

The material assigned to this species comprises large, inflated cadicones
with depressed whorls and subquadratic whorl sections. Characteristic are the
distinct flanks which merge into the broad, evenly rounded venter and provide
the whorl section with its subquadrate appearance. The rursiradiate primaries
terminate on the umbilical shoulder in about 20-22 somewhat bullate, very
prominent tubercles which generally give rise to 5, less commonly 4 or 6, almost
radial (SAM-PCUI1611) to prorsiradiate (BM-—C47126) fine secondaries,
usually with 3 intercalatories between bundles. Another feature which dis-
tinguishes this species from all other macroconchs from the Uitenhage Group
is the almost constant width of the final whorl, which tends to give the shell a
cylindrical aspect when viewed ventrally.

In a second specimen, SAM-—5074 which shows the same prominent
umbilical bullae and cylindrical form as SAM—PCU1611, the secondary ribbing
on the adoral portion of the body chamber is very fine and thread-like, with
fine secondaries occurring intercalated high up on the flanks.

On SAM-PCUI611 there are 23 secondaries per 3 bullae on the outer
whorl, with 18 secondaries within a 100 mm distance along the venter. On the
adoral portion of the body chamber the secondary ribbing becomes distinctly
prorsiradiate.

Measurements

~ No. D H Wi W/H Uo Ui
SAM-PCUI1611 226 70 Ft} 1,59 1h) di
SAM-5074 D335 78 110 1,41 85 ?

5; 165 15 100 35 65 v
BM-C47126 ZS 100 ID 12) 88 62 (29)

ANNALS OF THE SOUTH AFRICAN MUSEUM

246

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247

REVISION OF LATE VALANGINIAN CEPHALOPODA

f the holotype of

1eWw O

Lateral v
1903)

igerus Spath (Q)
ignon (after Uhlig

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Holcostephanus uh

Fig. 93. Olcostephanus (Olcostephanus) fasc

248 ANNALS OF THE SOUTH AFRICAN MUSEUM

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Fig. 94. Olcostephanus (Olcostephanus) fascigerus Spath (9).
Ventral view of the holotype of Holcostephanus uhligi Collignon
(after Uhlig 1903). x 1.

249

REVISION OF LATE VALANGINIAN CEPHALOPODA

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ANNALS OF THE SOUTH AFRICAN MUSEUM

250

201

REVISION OF LATE VALANGINIAN CEPHALOPODA

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P50) ANNALS OF THE SOUTH AFRICAN MUSEUM

Discussion A

The writer would agree with Spath (1939: 19) that this species should
include the adult figured by Uhlig (1903: 394, pl. 78 (fig. 1)) as Holcostephanus
(Astieria) cf. convoluta von Koenen (Figs 93-94). Since, however, Uhlig’s
specimen was subsequently made the type of the new species H. uhligi Collignon,
the latter name becomes a junior subjective synonym of O. fascigerus.

Although Spath (1939) and Fatmi (1977) mention constrictions in their
discussions of O. fascigerus, none of the figured material shows parabolae and
it is herein assumed that parabolae are lacking in the present interpretation of
this species.

Olcostephanus fascigerus can be distinguished from all other macroconch
dimorphs occurring in the Sundays River Formation by the weak inflation of
the whorls which provides it with a cylindrical aspect in ventral view, its promi-
nent umbilical bullae and its fine, irregular secondary ribbing.

Occurrence
This species is at present recorded only from northern Pakistan, Mada-
gascar, and South Africa.

Olcostephanus (Olcostephanus) aff. durangensis (Cantu Chapa) (3)
Figs 99-100

Hoplites symonensis Bose, 1923: 96, pl. 5 (figs S—16).
Taraisites durangense Cantu Chapa, 1966: 16.

Material

Two microconchs, SAM—PCU1547, 1549, collected in the Algoa Brick &
Tile quarries at Coega (see Fig. 1).

Fig. 98. Olcostephanus (Olcostephanus)
durangensis (Cantu Chapa). The
syntypes from the Taraises Formation
of northern Mexico, of which the
smaller specimen is herein selected as
lectotype (after Bose 1923). x 1.

REVISION OF LATE VALANGINIAN CEPHALOPODA 253

Holotype

By lectotype designation herein, the smaller and more complete of the
two individuals figured by Bése (1923) and here refigured (Fig. 98) from
northern Mexico.

Description

In the only complete example, SAM—PCU1549, which has recrystallized
test preserved, the shell is moderately inflated and involute up to the umbilical
bullae, with about 80 per cent of the preceding whorl covered. However, the
umbilical seam egresses noticeably on the final whorl. The umbilicus is rather
wide and moderately deep, with a steep umbilical wall which becomes somewhat
inclined with the egression of the umbilical seam. The umbilical shoulder is
subrounded. The whorl section is semicircular, with a broadly rounded venter.

The primary ribs begin at the umbilical seam and pass strongly backwards
(rursiradiate) to nineteen prominent bullae on the umbilical shoulder of the
final whorl. These bullae give rise to thick, robust secondaries, almost invariably
in pairs, although very occasionally there is only a single secondary rib arising
from a bulla. The rib direction changes significantly on the final whorl. On the
adapical portion of the outer whorl the secondaries are very slightly rursiradiate.
Half-way round the final whorl the secondary ribbing has become radial, while
at the peristome it is distinctly prorsiradiate. This latter feature is not, however,
considered characteristic since most olcostephanids show the same tendency
for the secondary ribbing to become more inclined near the peristome. There
is an occasional intercalated rib between bullae. A rib which occurs intercalated
on the one flank is frequently seen to arise from a bulla on the other side.

There are 15 secondaries per 6 bullae on the adoral portion of the body
chamber, with 10 ribs within a 40 mm distance along the venter. Prior to the
deep parabola on the outer whorl the ribbing is somewhat closer, with 6
secondaries within a 20 mm distance along the venter.

Measurements
No. D H Wi W/H Uo Ui
SAM-K 1549 65 26 36 1,38 30 20 (31)
bie 49 7a | 32 i SZ 23 14 (29)
SAM-K1547 45 16 D5) 1,56 19 12 (27)
e 35 14 v 2 17 10 (29)
Discussion

The two Uitenhage examples are characterized by robust, distant, pre-
dominantly radial secondaries, numerous umbilical bullae, paired secondaries
and semicircular whorl sections. The robust nature of the ribbing together with
the prominent umbilical bullae suggests better reference to Olcostephanus s:s.
rather than to the subgenus Jeannoticeras which lacks bullae and has finer,
more delicate secondaries.

254 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 99. Olcostephanus (Olcostephanus) aff. durangensis (Cantu Chapa). A—D. Ventral, front,

left lateral, and right lateral views of SAM-PCU1459, a microconch, x 0,66. E-F. Ventral

and lateral views of SAM-PCU1547, x 0,75. G. Lateral view of a badly crushed individual,

AAS-381, x 0,66. H-J. Front, ventral and lateral views of a juvenile in the Albany Museum
which may belong here, x 0,75.

_ &

REVISION OF LATE VALANGINIAN CEPHALOPODA QS)

Fig. 100. Olcostephanus (Olcostephanus) aff. durangensis (Cantu Chapa) (3).
A reconstruction based upon SAM-PCU1549. x 1.

Olcostephanus crassicostatus (Spath) is similar, but has fewer umbilical
bullae and commonly three secondaries per bulla, while O. rogersi (3) has
fewer, more prominent bullae, a more depressed whorl section and characteris-
tically three secondaries per bulla.

Olcostephanus durangensis (Cantu Chapa) (Fig. 98), erected for the Hoplites
symonensis described by Bose (1923) is very similar, but as the largest specimen
is only 16,2 mm in diameter comparison is rather difficult. The Mexican species
shows the characteristic radial bifurcation of coarse secondaries from the
umbilical bullae, of which there are about 24 per whorl (at 16 mm diameter),
but the umbilical walls of the Mexican form slope more. Olcostephanus baini
baini (3) has fewer umbilical bullae, from which arise 2-3 prorsiradiate
secondaries.

Fig. 101. Olcostephanus (Jeannoticeras) frequens (Zwierzycki) (2).
The holotype from the Rutitrigonia schwarzi Beds of Tanzania
(after Zwierzycki 1914). x 1.

256 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 102. Olcostephanus (Jeannoticeras) frequens (Zwierzycki) (3). Two of the syntypes of
Astieria auriculatus Zwierzycki from Tanzania, of which the right-hand specimen is herein
selected as the lectotype (after Zwierzycki 1914). x 1.

Olcostephanus frequens (Zwierzyck1) (Fig. 101), O. auriculatus (Zwierzyck1)
(Fig. 102), O. pecki Imlay (Fig. 103), and O. popenoei Imlay (Fig. 104) are all
characterized by abundant primary ribs from which the secondaries commonly
bifurcate and should, therefore, be included in the subgenus Jeannoticeras, as
should O. colorinensis Imlay (Fig. 105). It is doubtful whether more than one
species is involved in the above list. |

Bose (1923) created Hoplites aquilerae for a juvenile form with a rather
narrow umbilicus and a depressed, almost semicircular whorl section. On the
outer whorl, between 27 and 28 primary ribs terminate in weak bullae on the
umbilical shoulder and generally give rise to 2 prorsiradiate secondaries, with
occasional intercalatories between bundles. Parabolae are apparently lacking.
This species clearly belongs to the subgenus Jeannoticeras and differs from the
present material in its much finer ribbing, more numerous umbilical bullae and
the absence of parabolae.

Whiteaves (1893) introduced the species Olcostephanus (Astieria) deansii
(Fig. 106) for a compressed form lacking parabolae and with flexuous secondary
ribs which bifurcate just above the umbilical shoulder. This form is perhaps
better assigned to the genus Homolsomites.

Olcostephanus huizachensis (Cantu Chapa) (Fig. 107) is a small, somewhat
compressed species with an oval whorl section. Fifteen primaries terminate in
bullae from the umbilical shoulder from which secondary ribs arise in pairs, so
that there are thirty distant secondaries across the venter of the outer whorl.
This species is close to the present material from which it differs in lacking
parabolae and in having more distant secondaries. It is probably the microconch
of O. raricostatus (Bose).

‘Astieria’ neohispanica (Bose 1923) is a very involute, compressed form with
about one-quarter of the penultimate whorl visible in the umbilicus. There
are about 30-35 primary ribs which seem to lack umbilical tubercles. The lower
part of the flanks, immediately above the umbilical shoulder, are smooth.

REVISION OF LATE VALANGINIAN CEPHALOPODA

251,

°°

Bs
?

Fig. 103. Olcostephanus (Jeannoticeras) frequens (Zwierzycki) (2). The holotype of Olcoste-
Phanus pecki Imlay from the Upper Valanginian of Oregon (after Imlay 1960). x 1.

258 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 104. Olcostephanus (Jeannoticeras)
frequens (Zwierzycki) (3). The holotype of
Olcostephanus popenoei Imlay from the Upper
Valanginian of Oregon (after Imlay 1960). x 1.

Fig. 105. Olcostephanus (Jeannoticeras) colorinensis Imlay (@).
The holotype from the Taraises Formation of northern Mexico
(after Imlay 1938). x 1.

REVISION OF LATE VALANGINIAN CEPHALOPODA 259

Fig. 106. Homolsomites? deansii (Whiteaves).

The holotype, by monotypy, of Olcostephanus

(Astieria) deansii Whiteaves from Queen

Charlotte Island, British Columbia (after
Whiteaves 1893). x 1.

Fig. 107. Olcostephanus (Olcostephanus)

huizachensis (Cantu Chapa) (¢). The

holotype from the Taraises Formation

of northern Mexico (after Cantu Chapa
1966). x 1.

Above this smooth ‘periumbilical band’ fine secondaries arise and occasionally
bifurcate. Parabolae are lacking. In the writer’s opinion, this is a species of
O. (Jeannoticeras) which differs from the Uitenhage material in its much finer
ribbing.

-Olcostephanus elegans (Karakasch) (Fig. 108) has an inflated shell with a
strongly depressed whorl section and well-rounded venter. The umbilicus is
fairly wide, with sloping walls and well-rounded shoulders. There are about
thirty long primaries on the outer whorl which give rise to pairs of radial second-
aries with occasional intercalated ribs. This species is more finely ribbed than

260 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 108. Olcostephanus (?Jeannoticeras) elegans (Karakasch). The holotype,
by monotypy, from the Upper Valanginian of the Crimea (after Karakasch
1907). x 1.

Fig. 109. Olcostephanus? cadoceroides (Kara-

kasch). The holotype, by monotypy, from the

Upper Valanginian of Crimea (after Karakasch
1907). x 1.

the present material and lacks parabolae and is undoubtedly closely related to
‘Astieria’ cadoceroides Karakasch (Fig. 109) which seems to differ only in the
presence of a thick parabolic rib on the outer whorl which is said to form two
low tubercles on the venter. Whilst both these species show features of O. (Jean-
noticeras), the sloping umbilical walls with prorsiradiate primaries are features
of O. (Subastieria). However, the alleged ventral tubercles are not a feature of
Olcostephanus and hence the generic classification of these two forms is
uncertain.

The holotype of ‘Rogersites’ quinquestriatus Besairie (Fig. 110) shows a
subcoronate, depressed whorl section and a rather narrow umbilicus. The
umbilical shoulders appear evenly rounded and seem to lack umbilical bullae.
Simple rectiradiate ribs, about as wide as the interspaces, arise from the
umbilical shoulder (the umbilicus is plugged with matrix) and almost invariably
bifurcate above midflank, although the occasional rib remains single. There
are five prominent oblique parabolae on the outer whorl. This species differs
so markedly from the rather consistent characters shown by the genus Olco-
stephanus that it warrants a new generic name. In consequence, the new generic
name Jeanthieuloyites, for Dr J. P. Thieuloy of the University of Grenoble, is
proposed, with ‘R.’ quinquestriatus Besairie (Fig. 110) as type species.

Olcostephanus bangei (Bose) (Fig. 111) is a very involute species with a
narrow umbilicus and steep umbilical walls. Prominent radial primaries termi-
nate in about 10 tubercles on the outer whorl from which arise bundles of 3-4

REVISION OF LATE VALANGINIAN CEPHALOPODA 261

Fig. 110. Jeanthieuloyites quinquestriatus (Besairie). The holotype in the collections of the
University of Paris, from the Upper Valanginian of Ambiky, Malagasy Republic. x 1.

prorsiradiate secondaries, commonly with an intercalated rib between bundles.
There are 45 ribs across the venter of the outer whorl. This species differs from
the present material in lacking parabolae, and in having fewer primary and
secondary ribs. ‘Taraisites’ carillense Cantu Chapa (Fig. 112) and ‘Rogersites’
paucicostatus Imlay (Fig. 113) merely seem to be based upon different growth
stages of Bése’s (1923) species.

Occurrence

Olcostephanus durangensis is known with certainty only from Mexico,
though it may also be present in South Africa.

Fig. 111. Olcostephanus (Olcostephanus) bangei (Boése). The
holotype, by monotypy, from the Taraises Formation of northern
Mexico (after Bose 1923). x 1.

262 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 112. Olcostephanus (Olcostephanus)

bangei (Bose). The holotype of Taraisites

carillensis Cantu Chapa from the Taraises

Formation of northern Mexico (after Imlay
1937), <e

Fig. 113. Olcostephanus (Olcostephanus) bangei (Bose) (2). The holotype of Rogersites paucico-
status Imlay from the Taraises Formation of northern Mexico (after Imlay 1937). x 1.

REVISION OF LATE VALANGINIAN CEPHALOPODA 263

Olcostephanus (Olcostephanus) baini baini (Sharpe, 1856)

Figs 114-129, 130A-B, 131C-J, 132-136, 143A-B, 144A—D, 148, 150C—D,
151B-D

Microconch (3)

Ammonites baini Sharpe, 1856: 197, pl. 23 (fig. 2a-b).

Astieria baini (Sharpe) Kilian, 1902: 866.

Holcostephanus baini (Sharpe) Kitchin, 1908: 187.

? non Astieria aff. baini (Sharpe) Bése, 1923: 76, pl. 2 (figs 3-5) (= ? O. atherstoni).
Rogersites baini (Sharpe) Spath, 1930: 146.

Taraisites baini (Sharpe) Cantu Chapa, 1966: 16.

Olcostephanus baini (Sharpe) Neumayr & Uhlig, 1881: 156; Riccardi ef a/., 1971: 100.
Olcostephanus (Rogersites) schenki (Oppel) Fatmi, 1977: 270, pl. 5 (fig. 1).

Macroconch (Q)

Ammonites schenki Oppel, 1863: 286, pl. 81 (fig. 4a—c).

Astieria schenki (Oppel) Pavlow (in Pavlow & Lamplugh), 1892: 493.

Holcostephanus (Astieria) schenki (Oppel) Uhlig, 1903: 130, pl. 18 (fig. 2a—c only).

Astieria atherstoni (Sharpe) Baumberger, 1907: 39, figs 115-116 only.

Holcostephanus schenki (Oppel) Kitchin, 1908: 193, 198, 202-204. Kilian, 1910: 177. Spath,
1930: 150. Besairie, 1930: 629, pl. 64 (fig. 1—-la).

Holcostephanus cf. atherstoni (Sharpe) Kitchin, 1908: 193.

Rogersites douvillei Besairie, 1932: 44, pl. 5 (fig. 9-9a), fig. 2; 1936: 138, fig. 9 no. 2.

Rogersites baini var. ambikyi Besairie, 1936: 138, pl. 13 (fig. 5), fig. 9 no. 3.

Olcostephanus (Rogersites) schenki (Oppel) Spath, 1939: 30, pl. 2 (? fig. 6), pl. 18 (figs 9-10).

? Olcostephanus sublaevis Spath, 1939: 21, pl. 3 (figs 1-3), pl. 19 (fig. 2).

Holcostephanus douvillei (Besairie) Collignon, 1962: 43, fig. 869.

Olcostephanus atherstoni (Sharpe) Riccardi er al., 1971, pl. 13 (fig. Sa-c only).

Olcostephanus schenki (Oppel) Riccardi et al., 1971: 91, 97.

Material

42 specimens; 16 microconchs (PEM—1463/40a, b, 1462/76, SAM-—PCU1528,
1530, 1535, 1540, 1548, AM-429c, BM-—C52052, AAS-369a, SAM-525, 581,
583), 23 macroconchs (SAM-316, 1579, 5072, 6157, LJE-989d, PEM-1463/41,
1468/79, 1468/89, SAM—PCU1533, 1538, 1546, 1565, 1570, 1591, 1600, 1602,
1609, AAS—369b, 370, BM—C47122, AM-—2345, 2346), and 3 juveniles (SAM—
PCU1536, 1579, LJE—989f).

Holotype

By monotypy, the original of Ammonites baini figured by Sharpe (1856: 197,
pl. 23 (fig. 2a—b)) (Fig. 114) from the Sundays River.

Diagnosis

Dimorphic. Microconch small (about 50 mm diameter), rather inflated.
Primary ribs rursiradiate, terminating in 14-18 sharp umbilical bullae, from
which arise 2-3 coarse, prorsiradiate secondaries, often with an intercalated
rib between bundles. Usually two prominent parabolae per whorl. Peristome
with lateral lappets. Macroconch very large (about 250 mm diameter), very
strongly inflated, with a greatly depressed whorl section. Primary ribs rursi-

264 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 114. Olcostephanus (Olcostephanus) baini baini (Sharpe) (3). The holotype, in the British
Museum (Natural History). x 1. Photo: W. J. Kennedy.

radiate, terminating in 17-22 bullae on the body chamber, from which arise
bundles of 3-5 prorsiradiate secondaries, with 1-2 intercalatories between
bundles. Parabolae present on inner whorls only. Inflation decreases on adoral
portion of the body chamber.

Description

Microconch (3): the shell is small, rarely larger than 60 mm diameter, and .

involute up to the umbilical bullae on the inner whorls, so that about 80 per
cent of the preceding whorl is covered, but becoming slightly more evolute as
the umbilical seam egresses on the adoral portion of the body chamber. Conse-
quently, a short distance of the secondary ribbing of the penultimate whorl is
visible below the umbilical seam at this stage. The whorls are rather inflated,
with a depressed whorl section, the latter tending to become coronate as the
umbilical seam egresses. Prominent primary ribs begin at the umbilical seam
and curve backwards (rursiradiate) to distinct, sharp umbilical bullae on the
umbilical shoulder. Each bulla gives rise to two or three coarse, prorsiradiate
secondaries, frequently with an intercalated rib between bundles. Where there
are three secondaries per bundle the intercalated rib is usually absent. Often
a rib which is seen to arise from a bulla on the one flank is found to be inter-
calated on the other side. The secondaries recurve slightly so as to cross the
venter transversely. In some specimens there may be a slight concave inflexion
of the secondaries along the siphonal line. There are 14-18 bullae on the final
whorl, with 17-18 secondaries per 6 bullae. The umbilicus is rather narrow and
moderately deep, with steep umbilical walls and a subrounded shoulder. The
slope of the umbilical wall lessens as the umbilical seam egresses. There are
usually two deep parabolae per whorl, commonly about 180° apart. As is
characteristic of parabolae, they truncate the ribbing adapically, and are
parallel to the adoral ribs. The peristome is virtually identical to these para-

REVISION OF LATE VALANGINIAN CEPHALOPODA 265

Fig. 115. Olcostephanus (Olcostephanus) baini baini (Sharpe). A-C. Lateral, front and lateral

views of the inner whorls of AM—2346, an immature macroconch, x 075. D-E. Lateral and

front views of AM-4292c, a microconch, x 0,75. F-H. Ventral, front and lateral views of a

juvenile, SAM—PCU1536, doubtfully included here. Note the flattened venter and quadrate

whorl section, <x 1. I-J. Ventral and lateral views of a microconch in the Albany Museum,
x 1. K-L. Lateral and front views of LJE—989f, x 0,75.

266 ANNALS OF THE SOUTH AFRICAN MUSEUM

Sete FA.
we

Bh a bar a al

Fig. 116. Olcostephanus (Olcostephanus) baini baini (Sharpe) (3S). Reconstruction of the
peristome of the microconch. x 1.

(YOY

Fig. 117. Olcostephanus (Olcostephanus) baini baini (Sharpe) (3).
Whorl sections of SAM-—PCU1528 (left), PEM—1462/76 (right).
els

210
210

90 300
[a

Fig. 118. Plot showing the differing rates of inflation for macroconchs of O. baini (left) and
O. atherstoni (right). Note increased rate of inflation in middle whorls of O. baini.

bolae but, when preserved, the adoral margin is seen to bear slightly converging
lateral lappets. The whorl section varies somewhat, from almost semicircular
to coronate, with a well-rounded venter. The suture line is unknown. As in
other olcostephanids the body chamber is almost a full whorl in length.
Macroconch (Q): The shell is very large, commonly around 250 mm
diameter, and extremely inflated, with a strongly depressed whorl section. The
earliest whorls are only moderately inflated and up to a diameter of about
60 mm closely resemble the microconch, from which they differ only in being
slightly more inflated and in having invariably three secondaries per bulla,

a. a

REVISION OF LATE VALANGINIAN CEPHALOPODA 267

o
- |e
a
8
5
a
oO
A iz
Ss) A B | | &
= wad z
= aver A =
Aft +
ane 7 @ juveniles
@er

+ microconchs

A immature macroconchs

@ mature macroconchs

0 150 300

Diameter (mm) ————____»»

Fig. 119. Plot showing increase in rib density with ontogeny, and between dimorphs, in
O. b. baini (Sharpe).

commonly with an intercalated rib between bundles. At this stage, the macro-
conch is, therefore, slightly more densely ribbed than the microconch. After
about 60 mm diameter the whorl section rapidly becomes extremely inflated,
reaching a maximum about one-third of a whorl behind the peristome. The
umbilicus is very narrow and deep, with a vertical umbilical wall and acute
shoulder. The slope of the umbilical wall becomes less steep with the egression
of the umbilical seam while the umbilical shoulder also becomes distinctly
rounded at this growth stage. Prominent primary ribs begin at the umbilical
seam and curve backwards (rursiradiate) to terminate in 19-22 bullae on the
final whorl. One specimen, SAM-—PCU1591, has only 14 bullae on the outer
whorl, which become extremely swollen and large near the peristome, and is
with reservations referred to this species. Each bulla gives rise to bundles of
usually 3-4 coarse prorsiradiate secondaries, rarely 5, with 1-2 intercalated
ribs between bundles. There are between 23 and 28 secondaries per 5 bullae
on the adoral portion of the body chamber, with generally fewer at earlier
growth stages. The secondary ribbing is slightly flexuous, recurving on the
flanks so as to cross the venter transversely. At 155 mm diameter (PEM-—1468/79)
there are 24 secondaries per 6 bullae, and 9 ribs within a 50 mm distance along
the venter. In the same specimen at 210 mm diameter there are 21 secondaries
per 4 bullae, with a rib spacing of 7 mm. The inner whorls of macroconch
dimorphs of O. baini have prominent, deep parabolae, the largest diameter at
which they are seen to occur being 130 mm. However, in some examples taken
to belong to this dimorph by virtue of their very strong inflation, there is no
sign of parabolae at smaller diameters. This would seem to suggest that the
growth stage at which parabolae ceased to be produced varies from individual
to individual, although it would generally seem to be at about 90-120 mm

ANNALS OF THE SOUTH AFRICAN MUSEUM

268

‘py'0 X “MojsdsayjD °C 04 BdUR[QUIASeI SIYdioW
“O90WOY SION “ZLOS-WVS JO SMAIA [e1Oye] uw JOLT (5) (edaeYys) sg JuIDq (snuvydajsoz{Q) snuvydajsoz1Q “ZT ‘BLY

REVISION OF LATE VALANGINIAN CEPHALOPODA 269

oI
N

Width ———————a=—-

Diameter/

No. of bullae per VY whor| ——»

Fig. 121. Plot showing the relationship between inflation (Diameter/

Width) and number of umbilical bullae per half whorl in O. b. baini

dimorphs. Dots = microconchs, triangles = immature macroconchs,
squares = mature macroconchs.

diameter. Parabolae never occur on the outer whorls of macroconch forms,
although the peristome takes the form of a parabola.

Measurements

No. D H Wi W/H Uo Ui
AM-2344 50 Di 3h 1,48 ile 9 (18)
AM-2345 68 26 46 197, Ss) 13 (19) (Q)

5 56 22 32 1,45 18 10 (18)

270 ANNALS OF THE SOUTH AFRICAN MUSEUM |

Fig. 122. Olcostephanus (Olcostephanus) cf. baini (Sharpe) (2). Ventral and lateral views of a
specimen in the British Museum (Natural History). x 0,66.

No. D H Wi WH 9 Ue Ui
PEM-1468/73 180 wey 195 1,65 70 4424) (9)
135 62 90 1,45 50 - 27 @0)
SAM-PCUI1591 255 115 166 1,44 90 52 (20) (9)
z 174 85 150 1,76 69 ?
PEM-1468/89 250 92 155 1,68 110 a W823)
r 167 83 126 1,52 65 4326)
PEM-1468/79 225 87 OD 1,40 83. 46 (20) (2)
5 184 79 120 52 56 Wiha?)
SAM-316 208 93 138 1,48 18° a (2)(@)
: 145 60 100 1,67 55 ?
PEM-1468/80 170 100 ? ee Os RD
i 140 75 ? ? ?
AM-2346 55 23 37 1,61 Di 1731) (Q
. 38 13 21 1,61 13 7(18)
2 24 11 17 1,55 10 4(17)
SAM-PCU1546 —_ 226 108 170 1,57 55 4319). )
Ae 190 85 132 1,55 ? ?

271

REVISION OF LATE VALANGINIAN CEPHALOPODA

“bY

0 X ‘Snor[Iquin oy} S|] YOIYM Jo ouO ‘s1o}ShO

BUIJSNIOUS 9}ION ‘9PSTNOd-WYVS JO SMIIA JUOIS pue [B19}e'T (4) (odaeys) mig (snupydajso7Q) snuvydajso2/O

ETL “SIA

Mi AE me oe

ANNALS OF THE SOUTH AFRICAN MUSEUM

272

‘pro X “‘suIO}SIIed 94) puIYysq [IOYM ¥& JO PIIY}-oUO ynoge UOTeyUI
WNUNXeU SUIMOYS ‘6//89PI-INdd JO SMOIA [eIO}e] pue JUOI *(S) (edreys) tung (snuvydajso7jQ) snuvydajso71Q “pT “3

REVISION OF LATE VALANGINIAN CEPHALOPODA DAB

Fig. 125. Olcostephanus (Olcostephanus) baini baini (Sharpe) (2). A-—B. Front and lateral views

of a specimen in the South African Museum, x 0,44. C—D. Lateral and front views of SAM-—

PCU1533, an immature macroconch, x 0,75. E-G. Lateral, front and cross-sectional views
of inner whorls of SAM—PCU1533, x 0,75.

ANNALS OF THE SOUTH AFRICAN MUSEUM

274

‘pro X “oRT[Ng Jeorquin
Ud|[OMS BY} ION “T6STMOd-WVS JO SMOIA [es9;e] pue UOT “(4) (edseYs) Hq JUDG “Jo (snuvydo}so21Q) snunydajso21Q ‘9Z{ ‘B14

REVISION OF LATE VALANGINIAN CEPHALOPODA 215

Fig. 127. Olcostephanus (Olcostephanus) baini baini (Sharpe) (2), < 0,44. A-—B. Ventral and
lateral views of SAM-—PCU1609. C—D. Lateral and ventral views of SAM-—PCU1570.

ANNALS OF THE SOUTH AFRICAN MUSEUM

276

‘yr'0 X (08/897 I-INAd
‘UsWUIDEdS poYysn4io JeYMOWOS B JO SMOIA [BIJUDA puke [eIOJeT “(d) (Odreys) mg (snuvYydajsozjQ) snuvydajso7jC ‘3Z{ ‘3Iq

REVISION OF LATE VALANGINIAN CEPHALOPODA 277

Fig. 129. Olcostephanus (Olcostephanus) baini baini (Sharpe). A. A nucleus which possibly

belongs here, in the South African Museum. Note the prominent parabolae at a stage when

ribs are still absent, x 2,5. B. Lateral view of a microconch, BM-52052, x 1. C. Ventral
view of an immature macroconch, SAM—PCU1533, x 1.

278 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 130. A-B. Olcostephanus (Olcostephanus) cf. baini (Sharpe) (2). Lateral and ventral views
of AAS-369b, somewhat resembling O. actinotus (Baumberger), but with parabolae. x 0,75.
C. Olcostephanus (Olcostephanus) sp. (2). Lateral view of a crushed macroconch, SAM-—
PCU1551. Note peristome and coarse ribbing on adoral portion of the body chamber. x 0,44

REVISION OF LATE VALANGINIAN CEPHALOPODA 279

Fig. 131. A-B. Olcostephanus (Olcostephanus) cf. rogersi (Kitchin) (3). Ventral and lateral

views of SAM—PCU1527. Note inflexion in the apertural constriction which is lacking in the

parabola, x 0,75. C—J. Olcostephanus (Olcostephanus) baini baini (Sharpe) (3). C—E. Front,

ventral and lateral views of SAM-—PCU1528. F—H. Lateral and two ventral views of PEM-—

1463/40a; note peculiar chevron-shaped crushing of peristomal region suggestive of a bite-
mark. x 0,75. I-J. Ventral and lateral views of SAM-—525, x 1.

280

No.

SAM-PCU1544
SAM-PCU1538

SAM-PCU1594
AM-_2978
PEM_1463/40a
LJE_989f
AAS-369a
PEM_1462/76
SAM_PCU1528

PEM-1468/76

De)

ANNALS OF THE SOUTH AFRICAN MUSEUM

105
120

H Wi
50 81
50 85
40 oi
70 115
52 80
15 27
16 22
17 dpe

9 16
7 14

5 10
lS 2)
9 16
15 24
11 20
17 24
12 18
22 33)
17 22

28 (19) (2)

27 (21)

12 (24) (3)
7 (18)

11 (28) (dg)
:

6 (22) (3)
3 (18)
10 (23) (3)
9

12 (25) (3)
7 (23)

12 (24) (3)
7 (18)

13 (20) (3)
9 (18)

Fig. 132. Olcostephanus (Olcostephanus) baini baini (Sharpe) (Q). The
holotype of Olcostephanus schenki (Oppel) from the Salt Range of Pakistan

(after Uhlig 1903). x 1.

Sa a

> a ae

REVISION OF LATE VALANGINIAN CEPHALOPODA 281

hb Re
a ads wee
bs

PY
%
2

~
4

Fig. 133. Olcostephanus (Olcostephanus) baini baini (Sharpe) (2). The holotype of Rogersites
douvillei Besairie (fide Collignon 1962) and also Rogersites baini var. ambikyi Besairie, from
the Upper Valanginian of Ambiky, Madagascar (after Collignon 1962). x 1.

Discussion

As has been pointed out by many previous workers, O. baini (3) forma
typica bears a close resemblance to O. schenki (Oppel) (Fig. 132). It becomes
apparent from the present study that the differences are merely those due to a
comparison of the inner whorls of a macroconch with a microconch form.
Consequently, O. schenki (Oppel) is regarded asa junior subjective synonym of
O. baini baini. Riccardi et al. (1971) considered O. schenki to represent the inner
whorls of the O. atherstoni macroconch. However, as warned by Makowski
(1962a: 21), they failed to recognize the homoeomorphy between macroconch
forms. Hence the O. atherstoni fauna recorded by Riccardi et al. (1971) from
west-central Argentina includes both O. atherstoni and O. b. baini, as evidenced
by the presence of young macroconchs both with and without parabolae. This
same phenomenon recurs in the Uitenhage fauna, the homoeomorphy being
recognized by the differing rates of inflation of the two forms (Fig. 118).

Olcostephanus douvillei (Besairie) (Fig. 133) was erected for strongly inflated
forms with commonly three prorsiradiate secondaries per bulla, and parabolae
on the inner whorls. This species is undoubtedly conspecific with the O. b. baini
macroconch, representing merely an immature growth stage. Olcostephanus
baini var. ambikyi Besairie has the same holotype as Q. douvillei and is thus a
junior objective synonym.

In 1930, Besairie figured an example of O. schenki (Oppel) which he con-
sidered ‘...tout a fait semblable aux figures d’Uhlig’. The large size and
numerous secondaries, together with the presence of parabolae, suggest this

282 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 134. Olcostephanus (Olcostephanus) baini baini (Sharpe) (@).

The specimen figured by Spath (1939) as R. schenki (Oppel), and

considered to represent the holotype by Collignon (1962) (after
Spath 1939). This is an immature macroconch. x 1.

specimen represents the inner whorls of an O. baini macroconch. This example
does not, however, show the rapid inflation which characterizes the Uitenhage
forms at a similar growth stage. Collignon (1962: 36), in refiguring Besairie’s
specimen, made the interesting assertion that Uhlig had, in error, figured the
wrong specimen as Oppel’s holotype, and that the specimen of O. schenki
figured by Spath (1939, pl. 18 (fig. 9a—b)) (Fig. 134) was, in fact, the holotype.
There appears no justification, however, in this assertion since, according to
Oppel (1863: 287), the holotype came from ‘Schangra, west of Puling in Gnari-
Khorsum (Tibet)’, whereas Spath’s specimen came from ‘... near loc. 682’,
i.e. the northern side of Maranwal Nala, Makerwal Colliery. To suggest that
Spath’s specimen is the actual holotype would suggest that both Uhlig (1903)
and Spath (1939) had made the same mistake (A. C. Riccardi pointed out this
fact to the author).

Olcostephanus sphaeroidalis (Spath) is a macroconch form erected for those
species differing from O. atherstoni (2) in their greater inflation and more
depressed whorl section. These are the characters which distinguish the O. baini
macroconch from that of O. atherstoni. It is of interest to note, therefore, that
a re-examination of Spath’s holotype shows the inner whorls to bear parabolae.
Since the outer whorls of the macroconchs of O. baini (Sharpe) and O. sphaeroi-
dalis (Spath) cannot be distinguished, the latter is treated as a variety of the
former.

Olcostephanus (Subastieria) nicklesi Wiedmann & Dieni (Fig. 48) closely
resembles the microconch of O. baini baini, from which it differs largely in
having a sloping umbilical wall at all growth stages. This causes the umbilical
bullae to be situated at about mid-flank, resulting in a pentagonal whorl section.
This subtle difference is readily apparent, but whether it is of subgeneric rank

283

REVISION OF LATE VALANGINIAN CEPHALOPODA

‘T X “pomMosie vjoqeieg “Ieosesepeyy ‘AyIquiey jo uvlursurleA Joddy 94} wos;
OIMNVsog Sisuajayuis] Sajisdasoy JO sdAjo[oy OY, (6) (Od1eys) Ming 1u1ng (Snuvydajso2]Q) snuvydajso71Q

Cel “SIA

284 ANNALS OF THE SOUTH AFRICAN MUSEUM

would seem to depend on the recognition of sexual dimorphism in this sub-
genus, if it exists, and the morphology of the corresponding macroconch. It is
of interest to note that Wiedmann & Dieni (1968) included forms both with and
without parabolae in their species.

In 1923 Bose recorded a specimen of ‘Astieria aff. baini’ from the Taraises
Formation of northern Mexico. Cantu Chapa (1966) subsequently made this
specimen the type of his new genus Taraisites, renaming it 7. bosei (Fig. 9).
This genus was erected within the new subfamily Taraisitinae for those forms
of Olcostephanus in which 2-3 secondaries arise from each umbilical bulla.
Riccardi et al. (1971) have already shown this genus to be a synonym of
Olcostephanus s.s., also supported by the fact that the Ammonites baini of
Sharpe, assigned to Taraisites by Cantu Chapa (1966), represents nothing more
than the microconch of a much larger, more densely ribbed macroconch. The
preservation of ‘Taraisites bosei’ leaves much to be desired, but as it appears to
lack parabolae, and in view of the abundance of O. atherstoni in these beds, it
is probably best referred to that species.

The holotype of ‘Rogersites’ tsimihetensis Besairie (Fig. 135) is poorly
preserved but shows no features whereby it can be distinguished from the
O. baini macroconch. It is, therefore, a junior subjective synonym of Sharpe’s
(1856) species. |

Fig. 136. Olcostephanus (Olcostephanus) baini (Sharpe) (2). The holotype of Rogersites
sanlazarensis Imlay from the Taraises Formation of northern Mexico (after Imlay 1937). x 1.

REVISION OF LATE VALANGINIAN CEPHALOPODA 285

‘Rogersites’ sanlazarensis Imlay (Fig. 136) was created for large, fairly inflated
forms with a deep and narrow umbilicus, and steep umbilical walls. About 18
rursiradiate primaries terminate in small umbilical bullae from which arise
bundles of 3-4 prorsiradiate secondaries, with 1-2 intercalated ribs between
bundles. The inner whorls bear prominent parabolae. It is clear from the above
description and figure that ‘R.’ sanlazarensis should be considered a junior
subjective synonym of the O. baini macroconch, perhaps of the variety
sphaeroidalis (Spath).

“Astieria taurica Karakasch (Fig. 137) resembles the O. baini microconch,
but has more numerous constrictions and seems to lack umbilical bullae to most
secondary ribs. This species is very close to Jeanthieuloyites quinquestriatus
(Besairie), from which it seems to differ in that the secondary ribs do not
bifurcate before crossing the venter. It is, however, perhaps best referred to this
genus.

Fig. 137. Olcostephanus (?.Subastieria) tauricus (Karakasch). The syntypes,
of which the larger is herein selected as lectotype, from Crimea (after
Karakasch 1907). x 1.

porte)
i hp,
dee

Fig. 138. Olcostephanus (?Subastieria)

pavlowi (Karakasch). The holotype, by

monotypy, from the Upper Valanginian

of the Crimea (after Karakasch 1907).
alka

‘Astieria’ pavlowi Karakasch (Fig. 138) is based upon a tiny individual
with a strongly depressed whorl section, narrow umbilicus, rursiradiate
secondaries and prominent parabolae. It was compared with O. (Subastieria)
sulcosus Pavlow (in Pavlow & Lamplugh 1892) and differs from O. baini in the
rursiradiate direction of its secondary ribbing.

~ Olcostephanus hispanicus (Mallada 1882) was considered to lie at the
‘triple-junction’ of the subgenera Parastieria, Subastieria and Olcostephanus by
Wiedmann & Dieni (1968), although they placed the holotype within the sub-
genus Subastieria. The writer has not seen Mallada’s (1882) original figure or
description but, as figured by Nicklés (1890), this species is remarkably similar

286 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 139. Olcostephanus (Olcostephanus) dacquei
(Krenkel). The holotype, by monotopy, from the
Mikadi region of Tanzania (after Krenkel 1910). x 1.

to the inner whorls of the O. baini macroconch. Topotype material is necessary
to determine the specific and subgeneric status of the Spanish species.
Olcostephanus dacquei (Krenkel) (Fig. 139) is based upon a juvenile with a
rather narrow umbilicus and a depressed, coronate whorl section. Prominent
primary ribs terminate in 7 well-developed, somewhat rounded umbilical
tubercles from which arise 4-5 prorsiradiate secondaries, frequently with an
intercalated rib between bundles. There are about 37 ribs per half whorl, as
well as two prominent parabolae. This species differs from O. baini in possessing
fewer umbilical tubercles from which arise finer, more numerous secondary ribs.
The holotype of O. sublaevis Spath (Fig. 140) is entirely septate and seems
to be based upon the inner whorls of a macroconch. Based upon Fatmi’s (1977)
interpretation of this species, there are about 20 rursiradiate primaries (cer-
tainly more in Spath’s holotype) which terminate in bullae on the umbilical
shoulder and give rise to 4-6 slightly flexuous, prorsiradiate ribs. There is little
to distinguish this species from the O. baini macroconch, of which it seems

Fig. 140. Olcostephanus (Olcostephanus) sublaevis Spath (2). The holotype from the Spiti
Shales of Pakistan (after Spath 1939). x 1.

REVISION OF LATE VALANGINIAN CEPHALOPODA 287

Fig. 141. Olcostephanus (Olcostephanus) inordinatus (Tzankov). The syntypes from the Upper
Valanginian—Lower Hauterivian of Tchakantsi, Bulgaria, of which the upper specimen is
herein selected as lectotype (after Tzankov 1943). x 5.

288 ANNALS OF THE SOUTH AFRICAN MUSEUM

eS, 456
OO Bd cn |
a IRS

cote) UTD

RB. FZ. i 5:
ty j

Ss dl

> *

tae 5
Cea aie

oes

me » g
sEniauares eas F

H Bt eR a Feta i Se

Fig. 142. Olcostephanus (Olcostephanus) inordinatus (Tzankov) (2) (after Pictet 1860). x 1.

merely to represent an early growth stage. Olcostephanus sublaeyis is tentatively
included in the synonymy of O. baini.

Olcostephanus inordinatus (Tzankov) (Fig. 141) was based upon nuclei,
some of which are still partially smooth and hence comparison is difficult. At
this growth stage the whorl section is coronate, strongly depressed, and the
umbilicus is fairly narrow, with sloping umbilical walls. There seem to be
between 15 and 17 umbilical bullae on the outer whorl of the largest individual,
which is herein selected as lectotype, from which arise 3-4 prorsiradiate
secondary ribs. There are prominent parabolae. Since Tzankov (1943) included
the specimen figured by Pictet (in Pictet & Campiche 1860, pl. 17 (fig. 4),
pl. 18 (fig. 3)) (Fig. 142) into this species, this suggests that O. inordinatus loses its
parabolae in maturity. There is, thus, little to distinguish Tzankov’s (1943)
species from the O. baini macroconch, of which it may prove to be a synonym.

As already pointed out, O. crassicostatus (Spath) (Fig. 80) was originally
compared with O. baini but is herein interpreted as a gerontic O. rogersi
(Kitchin) microconch.

Occurrence

As interpreted here, this species appears to be known from Tibet, Pakistan,
?Spain, Madagascar, Argentina, northern Mexico, ?Oregon, and South Africa,
while the occurrence of strongly inflated forms with constricted inner whorls
in the Swiss Jura and Bulgaria suggests its presence in these areas as well.

REVISION OF LATE VALANGINIAN CEPHALOPODA 289

Fig. 143. A-B. Olcostephanus (Olcostephanus) baini (Sharpe) (2). Lateral and ventral views
of SAM-PCU1538, x 0,50. C-—D. Olcostephanus (Olcostephanus) atherstoni (Sharpe) (2)
Lateral and front views of SAM—PCU1604, x 0,66.

290 ANNALS OF THE SOUTH AFRICAN MUSEUM

Olcostephanus (Olcostephanus) baini var. sphaeroidalis (Spath, 1930)

Figs 144E-G, 145-150A-B, F, 153

Microconch (8)

? Astieria leptoplana Baumberger, 1908: 9, pl. 26 (fig. 4 only).

? Olcostephanus glaucus Spath, 1939: 17, pl. 6 (figs 7-8 only).

Holcostephanus ankaranensis Collignon, 1962: 40, pl. 191 (fig. 870).

Olcostephanus (Rogersites) madagascariensis var. isakhelensis Fatmi, 1977: 272, pl. 5 (fig. 3).

Macroconch ()

Rogersites sphaeroidalis Spath, 1930: 144, pl. 13 (fig. 5), pl. 15 (fig. 1).

? Olcostephanus sublaevis Spath, 1939: 21, pl. 3 (figs 1-3), pl. 19 (fig. 2); Fatmi, 1977: 269,
pl. 4 (fig. 2).

? Rogersites sanlazarensis Imlay, 1937: 560, pl. 72 (figs 1-3), pl. 74 (fig. 1).

Olcostephanus (Olcostephanus) sakalavensis Fatmi (non Besairie), 1977: 267, pl. 2 (fig. 3 only),
pl. 3 (fig. 2), pl. 4 (fig. 3).

Material

8 microconchs (SAM—PCUI1534, SAM-—PCU1592, SAM-PCU1593,
SAM-PCU1523, SAM-1525, PEM-—1468/76, BM—C41733, BM-C41731), and 1
macroconch (SAM-9241).

Holotype

By monotypy, the original of Rogersites sphaeroidalis figured by Spath
(1930: 144, pl. 13 (fig. 5), pl. 15 (fig. 1)) from the Sundays River and now in the
South African Museum, SAM-9241.

Diagnosis

Microconch moderately small, about 70-80 mm diameter, rather com-
pressed but with a depressed whorl section. Rursiradiate primaries terminate
in 18-22 umbilical bullae from which arise bundles of 3-4, rarely only 2,
prorsiradiate secondaries. Parabolae prominent. Macroconch large, outer
whorls indistinguishable from those of O. b. baini Inner whorls more finely
ribbed, with commonly 4 secondaries per bulla, and 1-2 intercalated ribs
between bundles. Parabolae present on inner whorls.

Description

Microconch (3): moderately small cadicones, about 70-80 mm diameter,
with somewhat compressed shells. The whorl section is depressed, with an
evenly arched venter. The whorls are involute up to the umbilical bullae,
covering about 80 per cent of the preceding whorl, except on the adoral portion
of the body chamber when the umbilical seam egresses slightly.

Primary ribs begin at, or close to, the umbilical seam and curve backwards
(rursiradiate) to 18-22 prominent bullae, on the umbilical shoulder of the final
whorl. Secondary ribbing commonly arises in prorsiradiate bundles of 3-4,
rarely only 2, often with an intercalated rib between bundles. The secondary
ribbing recurves on the flanks so as to cross the venter transversely. There
may bea slight adorally concave inflexion where the ribbing crosses the siphonal
line. Along the venter of the adoral portion of the body chamber there are

REVISION OF LATE VALANGINIAN CEPHALOPODA 291

Fig. 144. A-B, F-G. Olcostephanus (Olcostephanus) baini baini (Sharpe). A-B. Ventral and

lateral views of SAM-—581, a microconch, x 0,75. F—G. Front and lateral views of an immature

specimen, SAM-—PCU1579, x 0,66. Note fine ribbing and prominent parabola on smooth

nucleus. C—E. Olcostephanus (Olcostephanus) baini var. sphaeroidalis (Spath) (Q). Front,

ventral and lateral views of the holotype of Rogersites sphaeroidalis Spath. SAM-9241.
x 0,62.

292 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 145. Olcostephanus (Olcostephanus) baini var. sphaeroidalis (Spath) ($). A—B. Lateral and
ventral views of SAM-—PCU1592, x 0,75. C—E. Right lateral, left lateral and ventral views
of SAM-PCU1534, x 0,75. F-G. Lateral and ventral views of SAM-—PCU1575, x 0,66.

f
i
:
;

REVISION OF LATE VALANGINIAN CEPHALOPODA 293

Fig. 146. Olcostephanus (Olcostephanus) baini var. sphaeroidalis (Spath) (3). A-C. Front,

ventral and lateral views of SAM-—PCU1523, x 0,66. D-—F. Front, lateral and ventral views

of PEM-1468/76, a specimen transitional to O. b. baini, x 0,86. G—H. Ventral and lateral
views of SAM-—PCU1525, x 0,75.

294 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 147. Schematic representation of the rib pattern of Olco-
stephanus (Olcostephanus) baini var. sphaeroidalis (Spath) ($) on
PEM-—1462/76a (ac = apertural constriction).

60
E i i g7
= Ig 0.b. sphaeroidalis
dill
O0.b.baini
°50 390 mm

Diameter(mm) (9 ————-»>

Fig. 148. Diameter/width plot showing the relationship between microconchs of O. b. bain,
and O. baini var. sphaeroidalis.

REVISION OF LATE VALANGINIAN CEPHALOPODA 295

Fig. 149. Olcostephanus (Olcostephanus) baini var. sphaeroidalis (Spath)
(3). A-B. Lateral and ventral views of BM-—C47133. C—D. Ventral
and lateral views of BM—C47131. x 0,75.

12-14 secondaries within a 40 mm distance, with 19-22 secondaries per 5 bullae.
There are prominent parabolae on the outer whorl, while the peristome is
ornamented with lateral lappets.

Macroconch (@): as already pointed out, the outer whorls of the macro-
conch of this variety appear to be indistinguishable from those of O. baini baini.
Consequently, it is only the inner whorls of this macroconch form that can be
recognized. In SAM-9241 (the holotype of O. sphaeroidalis (Spath)), which
represents an immature macroconch, the whorls are moderately inflated to a
diameter of 55 mm, whereafter they increase very rapidly in width to become

296 ANNALS OF THE SOUTH AFRICAN MUSEUM

extremely inflated and strongly depressed. On the outer whorl, at 68 mm
diameter, about 16 rursiradiate primaries terminate in bullae from which
bundles of commonly 4, rarely 3 or 5, rather fine, prorsiradiate secondaries
arise and recurve slightly so as to cross the venter transversely. There are 26
secondaries per 5 bullae at this growth stage, with 16 ribs in a 40 mm distance
along the venter. Beyond this growth stage it seems unlikely that this variety
can be distinguished from O. baini baini (9).

Measurements
No. D H Wi W/H Uo Ui

SAM-—PCU1593 79 33 43 1,30 40 29 (37) (3)
SAM-—PCU1592 85 36 48 398) 41 L (3)
SAM-PCU1596 79 37 >)! 1,38 35 26 (33) (3)
SAM-PCU1595 13 53 41 1,24 q 2 (3)
SAM-—PCU1523 83 35) 42 1,20 40 26 (31) (3)

af 64 30 35 Peleg 29 18 (28)
SAM-—PCU1534 13 3 42 Heil 37 27 (37) (3)
SAM-PCU1525 70 30 355) Lol ? 2 (3)
PEM-1468/76 68 30 38 1,27 29 18 (26) (3)

2 43 20 28 1,40 22 b(26)
BM-C47133 68 PeA| 34 1,26 34 PIG2NCS)

Bi 58 20 30 1,50 23 15 (28) (3)
BM-C47131 68 Daf 35 1,30 32 19 (28) (3)
SAM-9241 c.130 65 110 1,69 ? ? (9)

ss 68 30 47 Lon 28 u

Do) 24 32 133 & u
Discussion

The microconch of this variety differs from the O. baini baini microconch
in its larger size (about 70-80 mm diameter as against 50-60 mm), more com-
pressed form, greater number of umbilical bullae (18-22 as against 14-18),
and more secondaries per bulla (3-4 as against 2-3). While these differences in
the microconchs were at first thought to be of specific value, the fact that the
macroconchs can be distinguished only during their early ontogenetic stages
suggests that the differences are of no more than varietal importance.

A single example, from the Sundays River Formation, PEM-1468/76
(Fig. 146D-F), appears to be transitional between O. baini baini (3) and
O. baini var. sphaeroidalis (3). In this specimen the whorl section is coronate
at the peristome and thus resembles O. baini baini in this respect, while there are
eighteen bullae on the final whorl, corresponding to the upper limit of O. baini
baini but below the average of about twenty for the O. baini var. sphaeroidalis
microconch. Moreover, the ribbing is finer and closer than in typical examples
of O. baini baini, while its dimensions are within the range of O. baini var.
sphaeroidalis.

REVISION OF LATE VALANGINIAN CEPHALOPODA 297

Fig. 150. A-B. Olcostephanus (Olcostephanus) baini var. sphaeroidalis (Spath) (3). Ventral

and lateral views of SAM-—PCU1592, x 0,75. C—D. Olcostephanus (Olcostephanus) baini

baini (Sharpe). Lateral and ventral views of AAS-370, an immature macroconch, x 0,75.

E, G. Olcostephanus (Olcostephanus) cf. rogersi (Kitchin) ($). Ventral and lateral views of

PEM-1468/42, x 0,75. F. Olcostephanus (Olcostephanus) cf. baini (Sharpe). Lateral view of

SAM-PCU1591, an immature macroconch, possibly referable to the variety sphaeroidalis
(Spath), x 0,66.

298 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 151. A. Olcostephanus (Olcostephanus) atherstoni (Sharpe) (2). Lateral view of the crushed

inner whorls of SAM-—320, x 0,66. B-D. Olcostephanus (Olcostephanus) baini baini (Sharpe)

(2). Ventral, lateral and front views of a specimen in the Albany Museum. Note the increase

in inflation immediately after a parabola, x 0,75. E-F. Olcostephanus (Olcostephanus) cf.

perinflatus (Matheron) (2). Lateral and front views of AAS—425. Note the extreme inflation
and fine ribbing, x 0,55.

REVISION OF LATE VALANGINIAN CEPHALOPODA 299

Fig. 152. Olcostephanus (Olcostephanus) glaucus Spath (2). The
holotype from the Spiti Shales of Pakistan. (after Spath 1939).
all

As can be seen from Fatmi’s (1977) figure and description, Olcostephanus
(Rogersites) madagascariensis var. isakhelensis Fatmi is a junior subjective
synonym of the microconch of this taxon.

Olcostephanus glaucus Spath (1939) (Fig. 152) differs from the present
species in being less inflated, more finely and densely ribbed and, in the holo-
type, lacking constrictions. The constricted paratype (Spath 1939, pl. 6 (fig. 7))
is very close, however, to the microconch of the present variant, and may prove
to be identical.

Collignon (1962: 44) considered O. ankaranensis (Fig. 153) to be
characterized by ‘...ses tours subcylindriques dont l’épaisseur reste a peu
prés constante, la réduction des flancs qui sont limités a la partie garnie de
tubercules ombilicaux, et par un ombilic profond et trés large’. So far as the
writer is able to judge, there are no features whereby this species can satis-
factorily be separated from the microconch of the present form, of which
O. ankaranensis (Collignon) thus becomes a probable junior subjective synonym.

300 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 153. Olcostephanus (Olcostephanus) baini var. sphaeroidalis Spath (3). The holotype of
Holcostephanus ankaranensis Collignon from the Upper Valanginian of Ambiky, Madagascar
(after Collignon 1962). x 1.

The constricted example of Astieria leptoplana figured by Baumberger
(1908, pl. 26 (fig. 4)) shows no features satisfactorily to distinguish it from the
present material with which it is here included.

Occurrence

This variety is known from the Swiss Jura, Pakistan, Madagascar, South
Africa, and possibly Mexico.

Olcostephanus (Olcostephanus) ventricosus (von Koenen, 1902)

Figs 154-156

Olcostephanus multiplicatus Neumayr & Uhlig (non Roemer), 1881: 150, pl. 33 (fig. 2).
Astieria ventricosa von Koenen, 1902: 144. Kitchin, 1908: 189.

? Astieria convoluta von Koenen, 1902: 146, pl. 39 (fig. 4a—b).

Astieria rigida Baumberger, 1908: 7, pl. 28 (fig. 1), fig. 121.

Holcostephanus (Astieria) ventricosus (von Koenen) Wegner, 1909: 87.

Material
Two specimens, LJE-989a, SAM-—PCU1607.

Holotype

The original of Olcostephanus multiplicatus Romer (Fig. 155) figured by
Neumayr & Uhlig (1881: 150, pl. 33 (fig. 2)) from H6heneggelsen, northern
Germany.

REVISION OF LATE VALANGINIAN CEPHALOPODA 301

Fig. 154. Olcostephanus (Olcostephanus) ventricosus (von Koenen), X 0,75. Lateral and
ventral views of SAM—PCU1607. C—D. Front and lateral views of LJE-989a.

302 ANNALS OF THE SOUTH AFRICAN MUSEUM

Diagnosis |

Medium-sized, somewhat inflated, with a semicircular whorl section.
About 18-22 rursiradiate primaries terminating in sharp bullae, from which arise
2-4, commonly 3, prorsiradiate, slightly flexuous, secondaries. Prominent deep
parabolae present. Possibly the inner whorls of a macroconch.

Description

Two specimens from the Sundays River beds, one rather crushed, are
referable to this species. LJE—989a has the recrystallized test preserved, and
closely resembles the type figured by Neumayr & Uhlig.

This example is somewhat inflated and moderately large. It is involute up
to the umbilical bullae and has a steep umbilical wall. The whorl section is
depressed, with an almost perfectly semicircular whorl section. Distinctly rursi-
radiate primaries terminate in 20 sharp bullae on the umbilical shoulder of the
outer whorl. These in turn give rise to 2-4, commonly 3, prorsiradiate
secondaries which recurve slightly on the upper part of the flanks so as to cross
the venter transversely. There are 25 secondaries per 7 bullae on the adapical
portion of the outer whorl (about 65 mm diameter) and 24 per 6 bullae on the
adoral portion. There are invariably 1-2 intercalated ribs between bundles,
with 11 secondaries within a 30 mm distance along the venter of the outer
whorl. The outer whorl is ornamented with 2 prominent parabolae.

SAM-PCU1607 is very similar, but rather crushed at the adapical portion
of the outer whorl. This specimen would appear to have part of a peristome
preserved, but there are no signs of lateral lappets.

ITNT OT A ne
oo peer nn aiine™ ese s

Fig. 155. Olcostephanus (Olcostephanus) ventricosus (von Koenen). The
holotype of Astieria ventricosa von Koenen, from north-west Germany
(after Neumayr & Uhlig 1881). x 1.

REVISION OF LATE VALANGINIAN CEPHALOPODA 303

Measurements
No. DD H Wi W/H Uo Ui
LJE-989a 80 2T 44 1,63 26 14 (18)
60 23 8. 1,43 25 12 (20)
se 45 17 23 135 7. 8 (18)
SAM-PCU1607 109 53 64 20 34 DI(19)
Discussion

This is a rather problematical form since it is uncertain whether it represents
the inner whorls of a macroconch, or if it is a large microconch species. Its
large size would seem to favour the former, whilst its lack of inflation suggests
the latter.

This species is clearly very closely related to O. baini from which it seems
to differ only in remaining relatively compressed to large diameters.

According to Von Koenen (1902), Astieria convoluta is a moderately
inflated form with a deep, narrow umbilicus. Rursiradiate primaries terminate
on the umbilical shoulder in bullae, of which there are about 8 per half whorl.
There are generally 3 prorsiradiate secondaries per bulla, with an intercalated
rib between bundles, and a prominent parabola on the outer whorl. There is
little to separate this species from O. ventricosus, of which it may prove to be a
junior subjective synonym.

Astieria dalpiazi Rodighiero (1919) was compared with O. ventricosus but
has never been figured. It was said to have short, weak, thin primary ribs which
terminate in tubercles on the umbilical shoulder from which arise 2-3 secondary
ribs. The secondaries bifurcate at various levels on the flanks so that 4-6 ribs
cross the venter for each tubercle. Parabolae are present. According to
Rodighiero (1919), ‘A.’ dalpiazi differs from O. ventricosus in being more inflated,
with a narrower, deeper umbilicus. However, Von Koenen’s (1902) species
does not have bifurcating secondaries and is probably specifically distinct.

Olcostephanus rigidus (Baumberger) (Fig. 156) does not appear to be
specifically distinct from this form, and supports its treatment as the inner
whorls of a macroconch. Baumberger’s species was erected for moderately
inflated forms with 23-24 rursiradiate primaries on the outer whorl, each
terminating in umbilical bullae from which arise bundles of 3-4 prorsiradiate
secondaries commonly with 2 intercalated ribs between bundles. There is a
prominent parabola on the outer whorl. The whorl section is semicircular. Thus,
there is no significant feature by which this species can be distinguished from
O. ventricosus and they should be considered conspecific.

Since the outer whorls of this probable macroconch form are unknown,
as is the microconch dimorph, its true status is not known, and must await
study of topotype material.

304 ANNALS OF THE SOUTH AFRICAN MUSEUM

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Fig. 156. Olcostephanus (Olcostephanus) ventricosus (von Koenen). The
lectotype, designated herein, of Astieria rigida Baumberger, from the
Swiss Jura (after Baumberger 1908). x 1.

Occurrence

This species is known from the Swiss Jura, northern Germany, South
Africa, the south of France, and possibly Tanzania.

Olcostephanus (Olcostephanus) uitenhagensis (2) (Kitchin, 1908)
Figs 157, 158C—D, 159, 160C-—D

Holcostephanus uitenhagensis Kitchin, 1908: 206, pl. 11.

Holcostephanus (Astieria) uitenhagensis Kitchin, Wegner, 1909: 89. Kilian, 1910: 214.

Holcostephanus uitenhagensis Kitchin, Hatch & Corstorphine, 1909: 295, fig. 73 (right-hand
specimen only).

Rogersites uitenhagensis (Kitchin) Spath, 1930: 150. Besairie, 1936: 141.

Olcostephanus uitenhagensis (Kitchin) Spath, 1939: 19. Riccardi et al., 1971: 90. Reyment
& Tait, 1972: 60.

Material

Three specimens, all macroconchs (SAM-5069, SAM—PCU1524, SAM-—
PCU1605).

305

REVISION OF LATE VALANGINIAN CEPHALOPODA

® “690S-INVS

‘adAjo[OY 94} JO SMOIA

BSS

0 X ‘prhow jeusojur poarosoid [100d

juoIf pue jeroyeT “(S) (UIYDITY) sisuasoyuajin (snuvydajso21Q) snuvydajso7jO “LS{ “sid

306 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 158. A-B. Olcostephanus (Olcostephanus) uitenhagensis (Kitchin) (9). Lateral and ventral

views of SAM-—PCU 1605, retaining patches of recrystallized test. x 0,44. C—D. Olcostephanus

(Olcostephanus) sp. (2). Ventral and lateral views of SAM-—PCU1531. Note fine, flexuous
ribbing and parabola. x 0,55.

REVISION OF LATE VALANGINIAN CEPHALOPODA 307

Holotype

By monotypy, the original of Holcostephanus uitenhagensis Kitchin (1908:
206, pl. 11) (Fig. 157) from ‘... between milestones 244-243 on the Graaff-
Reinet railway, about three miles from Uitenhage’. The holotype, SAM-—5069
is in the South African Museum.

Diagnosis

Large (about 200 mm diameter), compressed macroconchs which become
markedly evolute on the final whorl. Flanks prominent, steep and slightly
convex. On the penultimate whorl! rursiradiate primaries terminate in 19 bullae
on the umbilical shoulder. On the final whorl the primaries become very weak,
disappearing entirely on the internal mould, and the tubercles become swollen
and rounded, as well as fewer (15) in number. Flexuous prorsiradiate secondaries
arise in bundles of 3—5, with 1-2 intercalated ribs between bundles. Parabolae
lacking, on outer whorls at least. Peristome simple.

Description

Only macroconchs of this species have been described, and until the onto-
genetic variation is known it is highly unlikely that the microconch dimorphs
will be recognized.

The shell is a large, compressed cadicone, involute up to the umbilical
bullae except on the last two-thirds of the body chamber when the umbilical
seam egresses rapidly and the final whorl becomes about 30 per cent evolute at
the peristome. The umbilicus is fairly narrow and rather deep, with vertical
umbilical walls on the inner whorls and an acute umbilical shoulder. On the
body chamber, with the egression of the umbilical seam, the slope of the
umbilical wall decreases markedly and the shoulder becomes rounded.

Also with the egression of the umbilical seam, the prominent rursiradiate
primaries weaken considerably and are almost entirely effaced on the adoral
portion of the final whorl. At the same time, the distinctly bullate umbilical
tubercles of the penultimate whorl become swollen and rounded. There are
15 umbilical tubercles on the body chamber, whereas the penultimate whorl
is ornamented with 19 bullae.

Specimen SAM-—PCU1524 is a well-preserved example of this species which,
unlike the holotype which represents an internal mould, has the recrystallized
test preserved. Unfortunately it has been slightly crushed laterally.

On this example, the secondary ribbing is rather fine and slightly prorsi-
radiate, becoming distinctly coarser and more inclined near the peristome. On
the penultimate whorl the secondaries are slightly flexuous, arising radially
from the umbilical bullae only to curve forward low down on the flanks and
then gently recurve so as to cross the venter transversely. On the penultimate
whorl there are 16 secondaries per 3 bullae, while on the adoral portion of the
body chamber there are 25, with 13 ribs within a 60 mm distance along the

308 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 159. Olcostephanus (Olcostephanus) uitenhagensis

(Kitchin) (2). Whorl section of SAM-PCU1524.

Dotted line represents shape of the final peristome.
x 0,5.

venter. Parabolae are lacking on the outer whorls, although the peristome is
strongly constricted, cutting obliquely across 4 ribs to the posterior, and with a
prominently flared adapical margin.

Measurements
No. D H Wi W/H Uo Ui
SAM-—5069 225 88 c.93 |e Fa 87 67 (30)
a ENDS 69 82 1 b9 49 37 (24)
SAM-PCU1524 188 qa (x7/\ 1,00 74 S27)
Pe [55 67 c.64 0,96 Sy 2) (4)
Discussion

Olcostephanus uitenhagensis is a macroconch characterized by the marked
egression of the umbilical seam on the final whorl, its compressed form, and
the change in the nature of the tuberculation on the body chamber.

Olcostephanus of the astierianus plexus do not show the marked egression

REVISION OF LATE VALANGINIAN CEPHALOPODA 309

Fig. 160. A-B. Olcostephanus (Olcostephanus) riccardii sp. nov. (2). Lateral and ventral

views of the holotype, SAM—PCU1577. Note inflated form, lack of parabolae and rounded

umbilical tubercles on the body chamber. x 0,44. C—D. Olcostephanus (Olcostephanus)

uitenhagensis (Kitchin) (2). Lateral and ventral views of SAM-—PCU1524. {Note compressed

form, egression of the umbilical seam, rounded tubercles on the body chamber and constricted
peristome. x 0,44.

310 ANNALS OF THE SOUTH AFRICAN MUSEUM

“s
>
3.
His
{
e
?
2
s
:
:

Fig. 161. Olcostephanus (Olcostephanus) singularis
(Baumberger). The lectotype, designated herein, from
the Swiss Jura (after Baumberger 1908). x 1.

of the umbilical seam displayed by O. witenhagensis in maturity, as well as
commonly retaining bullate umbilical tubercles to the peristome.

Olcostephanus singularis (Baumberger) (Fig. 161) is the closest species to
O. uitenhagensis. It has a moderately inflated shell with a strongly depressed
whorl section and shows marked egression of the umbilical seam on the outer
(?final) whorl. About 12, rounded, swollen, umbilical tubercles on the outer
whorl give rise to bundles of 5-6 slightly prorsiradiate secondaries, between
which are 2-3 intercalated ribs. Parabolae are lacking. This species differs from
O. uitenhagensis in that the umbilical tubercles are swollen and rounded even
on the inner whorls as witnessed by O. klaatschi (Wegner) which is merely
based upon a juvenile of Baumberger’s (1908) species.

Olcostephanus rabei (Besairie) (Fig. 69A—B), as noted by Besairie (1936),
is indeed very close to the inner whorls of O. uitenhagensis. However, without
knowledge of the final whorls, the true affinities of O. rabei are obscure as it
also closely resembles O. sakalavensis.

Occurrence

This species has been known with certainty only from South Africa,
although Reyment & Tait (1972) have recently reported its occurrence in
Argentina.

——<—<—<

REVISION OF LATE VALANGINIAN CEPHALOPODA Si

Olcostephanus (Olcostephanus) riccardii sp. nov.
Figs 162-163

Material

Two specimens, SAM—PCU1577 and BM-—C47130, of which the former is
designated as the holotype, and the latter is a paratype.

Holotype
SAM-PCU1577 from the Algoa Brick & Tile quarries at Coega.

Diagnosis

A strongly inflated, globose species of Olcostephanus with a very depressed
whorl section and a deep, narrow umbilicus. Inner whorls with about nineteen
umbilical bullae becoming swollen and rounded on the body chamber. Para-
bolae lacking.

Fig. 162. Olcostephanus (Olcostephanus) riccardii sp. nov.
(2). Lateral view of the paratype, BM—C47130. x 0,44.

312 ANNALS OF THE SOUTH AFRICAN MUSEUM

Etymology

For Dr A. C. Riccardi of the Museo de Ciencias Naturales, La Plata,
Buenos Aires, in appreciation of his help in providing literature and constructive
criticism of the original manuscript.

Description

The holotype is a strongly inflated, globose cadicone with a narrow
umbilicus and steep umbilical walls. The whorls, involute up to the umbilical
bullae on the inner whorls, become rather evolute on the adoral portion of the
outer whorl. Distinct primaries begin at the umbilical seam and pass backwards
(rursiradiate) to 19 bullae on the sharp umbilical shoulder. On the outer whorl
the bullae become swollen and distinctly rounded and the primaries are effaced
(on the internal mould). There are about 15 tubercles on the final whorl.

On the penultimate whorl secondary ribbing arises in slightly prorsiradiate
bundles of 3-4 ribs, commonly with an intercalated rib between bundles, so
that there are 17 ribs per 4 bullae. The secondaries recurve slightly so as to
cross the venter transversely. On the adoral portion of the body chamber there
are 28 secondaries per 4 bullae, with 8 ribs in a 40 mm distance along the venter,
whereas on the adapical portion of the outer whorl there are 10 secondaries
within a similar distance. |

The peristome is not preserved, but the inflated shell leaves little doubt
that this form represents a macroconch.

Measurements
No. D H Wi W/H Uo Ui
SAM-PCU1577 150 62 94 Il 52 60 45 (30)
a 110 50 83 1,66 39 26 (24)
BM-—C47130 182 i ? i 80 52 (29)
Discussion

The ornament of this species is almost identical to that of the holotype of
O. uitenhagensis, the only difference being that whereas Kitchin’s type, together
with further topotype material, has a noticeably compressed form, this taxon
is distinctly inflated and globose. To admit extremely inflated and strongly
compressed individuals into the same dimorph would so drastically alter the
taxonomy of the group that present evidence does not justify such a move, and
hence a new species is created for the present material.

The inner whorls of this species may be distinguished from O. atherstoni
() by their much greater inflation, and more depressed whorl section.

Mature O. baini macroconchs attain a much larger size than this species,
while the tubercles remain bullate to the peristome. Moreover, O. riccardii
does not show the decrease in inflation near the peristome which characterizes
O. baini macroconchs. Immature examples of O. baini (Q) differ in having
parabolae.

REVISION OF LATE VALANGINIAN CEPHALOPODA 313

Fig. 163. Olcostephanus (Olcostephanus) riccardii sp. nov. (2). Whorl section of the holotype,
SAM-PCU1577. x 1.

314 ANNALS OF THE SOUTH AFRICAN MUSEUM

Olcostephanus perinflatus (Matheron) is close to the inner whorls of
O. riccardii but has radial secondaries, and retains bullate tubercles on to the
body chamber.

Occurrence

This species is known only from the Sundays River Formation.

Olcostephanus (Olcostephanus) sp.
Fig. 158C-D.

Material
A single specimen, SAM—PCU1531, representing an immature macroconch.

Description

A moderately large, strongly inflated cadicone, involute up to the umbilical
bullae at all stages on the outer whorl which includes the body chamber. The
umbilicus is very narrow and crater-like, with steep convex walls and subangular
shoulders. Rursiradiate primaries begin at, or close to, the umbilical seam and
terminate in 19 small bullae on the umbilical shoulder of the outer whorl. Each
bulla gives rise to bundles of 3-4 fine secondaries, often with an intercalated
rib between bundles. On the adapical portion of the outer whorl, however, the
nature of the secondary ribbing changes significantly. The secondaries become
noticeably finer and distinctly flexuous. From the bullae the secondaries are
directed radially for a short distance, before curving forwards to become
slightly prorsiradiate and finally recurving so as to cross the venter transversely.
There are 17 ribs per 4 bullae on the adoral portion of the outer whorl, with 10
secondaries within a 30 mm distance along the venter. Two indistinct parabolae,
due to the fact that the outer whorl is preserved as an internal mould, are present
on the body chamber. The whorl section is strongly depressed with a broad,
rounded venter.

Measurements

No. D H Wi W/H Uo Ui
SAM-K1531 93 4] 67 1,63 33 19 (20)
Discussion

This example differs from O. baini macroconchs at a similar diameter by
its finer, more flexuous secondary ribbing.

Olcostephanus (Olcostephanus) astieriformis (Bose, 1923) (Q)
Figs 164-169

Astieria astieriformis Bose, 1923: 72, pl. 1 (figs 1-4). Riedel, 1938: 13.
Olcostephanus astieriformis (Bose) Imlay, 1938: 553.

REVISION OF LATE VALANGINIAN CEPHALOPODA Bile

Fig. 164. Olcostephanus (Olcostephanus) astieriformis (Bése) (2). Lateral view of a slightly
crushed specimen, BM—C47129. x 0,44.

316 ANNALS OF THE SOUTH AFRICAN MUSEUM |

Fig. 165. Olcostephanus (Olcostephanus) astieriformis (Bose) (2). Lateral view of BM—C47132.
x 0,44.

REVISION OF LATE VALANGINIAN CEPHALOPODA 3477

Fig. 166. Olcostephanus (Olcostephanus) astieriformis (B6ése)
(2). Ventral view of BM—C47132. x 0,44.

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UBOLIFY YINOG oY} UI USUIDedS B JO SMOIA [V.IJUSA Pue [eIa}eT “(S) (280g) SNUAOf1Aa1ISD (snuBYyda]soI1Q) SnubYydajsoI]Q “LOT *3t4

ANNALS OF THE SOUTH AFRICAN MUSEUM

318

REVISION OF LATE VALANGINIAN CEPHALOPODA 319

Material

Nine crushed and fragmentary specimens, all macroconchs (BM-—C47132,
BM-—C47129, BM—C47125, SAM-—PCU1555, SAM-—PCU1541, SAM-PCU1554
SAM-—PCU1558, SAM-PCU1553, SAM-PCU1557).

Holotype

By monotypy, the original of Astieria astieriformis Bése (Fig. 168), from
Durango-Zacatecas, northern Mexico.

Description

A rather variable collection of compressed macroconchs belong here, as
well as showing affinities to numerous other nominal species (Figs 169-170).

They are all compressed and show little or no egression of the umbilical
seam of the body chamber. The primary ribs are rursiradiate, terminating in
16-25 bullae on the umbilical shoulder from which arise commonly 3-4
prorsiradiate secondaries, usually with intercalatories between bundles. The
secondaries very occasionally bifurcate. Parabolae are lacking on the outer
whorls, at least, while the inner whorls are currently unknown.

The best preserved example from the Sundays River Formation assignable
to this species is BM—C47132. It is an extremely large macroconch (diameter
275 mm) which shows only slight egression of the umbilical seam of the final
whorl. The shell is rather compressed, with a narrow deep umbilicus and vertical
umbilical walls. The latter are ornamented with 18 rursiradiate primaries
terminating in prominent bullae from which bundles of 4-6, fewer at earlier
growth stages, flexuous secondaries arise. There is invariably 1-2 intercalatories
between bundles. On the adoral portion of the body chamber there are 37
secondaries per 5 bullae, whereas on the adapical portion there are only 23.
Parabolae appear to be lacking.

A second example, SAM-—PCUI555, is a moderately large (190 mm
diameter), compressed cadicone which shows virtually no egression of the
umbilical seam on the final whorl. Primary ribs begin at the umbilical seam and
pass backwards (rursiradiate) to 17 bullae on the umbilical shoulder, from which
3-5 slightly flexuous, prorsiradiate secondaries arise. There is commonly an
intercalated rib between bundles. On the adoral portion of the final whorl there
are 18 secondaries per 3 bullae, with a rib spacing of 6 mm. The whorl section
is about as wide as high.

Measurements
PoNo: D H Wi W/H Uo Ui
BM-—C47132 DUIS) 120 125 1,04 94 74 (27)
a 220 100 ? ? 70 54 (25)

SAM-PCUI1555 190 87 ? ? 70 ~—-50.(26)

ANNALS OF THE SOUTH AFRICAN MUSEUM

320

‘LT X “(EZ6I BOG 191Je) ODIXOJY UIOyIIOU Jo
UONCUIIO Sosieley OY} WOIy “UloINYy poyeusIsop ‘edA}0}99] SUL, “(S) (CSO) SIM4Of14aIIsD (snubYydajso21Q) snubYydajsoz/O *89T ‘SIq

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2.
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REVISION OF LATE VALANGINIAN CEPHALOPODA 321

No. of ribs per half whorl

150 300

Diameter (mm) :

Fig. 169. Plot of rib density against diameter for ‘species’ of the O. astierianus plexus. Triangles
represent individuals from the Sundays River Formation. 1 = O. raricostatus (Bose) (after
Bose 1923, pl. 4 (fig. 1)), 2 = O. quadriradiatus Imlay (after Imlay 1938, pl. 5 (fig. 2)), 3 =
O. astieriformis (Bose) (after Bose 1923, pl. 1 (fig. 2)), 4 = O. scissus (Baumberger) (after
Baumberger 1907, fig. 107), 5 = O. discoideus Imlay (after Imlay 1938, pl. 2 (fig. 5)), 6 =
O. symonensis (Bose) (after Bose 1923, pl. 2 (fig. 7)), 7 = O. catulloi (Rodighiero) (after
Rodighiero 1919, pl. 9 (fig. 9)), 8 = O. rabei (Besairie) (after Besairie 1936, pl. 12 (fig. 8)),
9 = O. astierianus (d’Orbigny) (after Baumberger 1910, pl. 32 (fig. 1)), 10 = O. sayni (Kilian)
(after Baumberger 1910, pl. 32 (figs 2-3)), 11 = O. astierianus (d’Orbigny) (in Baumberger
1910, pl. 29 (fig. 3)), 12 = O. sayni (SAM-9270), 13 = O. scissus (Baumberger) (in Baum-
berger 1907, pl. 23 (fig. 2)), 15 = O. elongatus (Tzankov) (after Tzankov 1943, pl. 6 (figs 3-4)),
16 = O. scissus (Baumberger) (in Matheron 1878, pl. B—20 (fig. 8)), 17 = O. catulloi (Rodi-
ghiero) (in Tzankoy 1943, pl. 4 (figs 1-2)), 18 = O. boesei (Riedel) (after Riedel 1938, pl. 3
(fig. 1)), 19 = O. astierianus (d’Orbigny) (in Riedel 1938, pl. 3 (fig. 3)) 20 = O. schafarziki
(Somogyi) (after Somogyi 1916, pl. 13 (fig. 3)), 21 = O. filosus (Baumberger) (in Bayle 1878,
pl. 55 (fig. 2)), 22 = O. astierianus (dW Orbigny) (in Somogyi 1916, pl. 13 (fig. 2)).

Discussion

Amongst the collections from the Sundays River Formation are a rather
large number of compressed macroconchs, unfortunately generally rather
crushed and fragmentary. These forms differ from O. uitenhagensis in lacking
the marked egression of the umbilical seam of the body chamber, and in that
the tubercles remain bullate to the peristome.

These specimens are undoubtedly close to the type of the genus, O. astieri-
anus (d’Orbigny) (Fig. 171). The latter species is rather compressed, with a
moderately wide, deep, umbilicus and sloping umbilical walls ornamented with
16 radial primaries on the outer whorl. These terminate in prominent, somewhat

322 ANNALS OF THE SOUTH AFRICAN MUSEUM

ae)
Lee
17) 10a ane
8 7
21
5 ce ACS WA
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2 A AA 6 A
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20 40 60 80 100 .
No. of ribs per half whorl

Fig. 170. Plot of rib density against umbilical bullae for ‘species’ of the O. astierianus plexus.
Numbers and symbols as for Figure 85.

Fig. 171. Olcostephanus astierianus (d’Orbigny) (2). Ventral ‘and lateral views of the lectotype
from the Lower Hauterivian of Castellane, France. x 1.

REVISION OF LATE VALANGINIAN CEPHALOPODA 323

con AU pry,
1p
LbA “A

‘\

Fig. 172. Olcostephanus (Olcostephanus) astierianus (d’Orbigny) (2). A copy of
D’Orbigny’s (1842a) protograph. x 1.

rounded umbilical tubercles on the outer whorl that give rise to bundles of
5—6, fine, prorsiradiate secondary ribs, commonly with 2-4 intercalatories
between bundles. Contrary to D’Orbigny’s (1840) (Fig. 172) protograph, the
umbilical seam does not egress on the outer whorl, but remains just above the
umbilical tubercles. This species seems to be based upon a macroconch, with a
(?) simple peristome preserved at only 89 mm. According to Baumberger
(1907: 28), the inner whorls bear parabolae, in which case they are to be expected
on the outer whorls of the microconch. Olcostephanus astierianus differs from
the present species in having fewer, rounded tubercles on the outer whorl and,
perhaps, in possessing parabolae on the early whorls. Its small adult size, like
much of the west European material, is probably environmentally controlled
and is not herein considered of specific importance.

Olcostephanus astierianus globulosus (Kilian) (in Roch 1930: 313) was erected
for the example of ‘Astieria sayni’ figured by Baumberger (1908: 1, pl. 25
(fig. 3a—b)). Since, however, the specimen in question is listed as “Astieria
guebhardi’, presumably it is this individual to which Kilian referred. In the
writer’s opinion, it is too inflated and with too many bullate umbilical tubercles
to be conspecific with O. astierianus. Indeed, the original identification, that is
as O. guebhardi, is more likely to be correct.

324 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 173. Olcostephanus (Olcostephanus) boesei (Riedel). The holotype from the Upper
Valanginian of Caqueza, Colombia (after Riedel 1938). x 1.

Fig. 174. Olcostephanus (Olcostephanus) catulloi (Rodig-
hiero). The holotype from Venice (after Rodighiero 1919).
1;

REVISION OF LATE VALANGINIAN CEPHALOPODA 325

Olcostephanus boesei (Riedel) (Fig. 173) is a somewhat compressed species
with a moderately wide umbilicus and steep umbilical walls. The latter are
ornamented with 7-8 relatively pronounced primary ribs per half whorl, each
terminating in a bulla on the umbilical shoulder and giving rise to 4-5 fine,
prorsiradiate secondaries. There is commonly 3-4 intercalated ribs between
bundles. Parabolae seem to be lacking. This is a finely ribbed species of the
astierianus plexus which was compared with O. sayni (Kilian), from which it
was distinguished by its fewer umbilical bullae. It is, therefore, doubtfully
separable from O. scissus (Baumberger).

Olcostephanus catulloi (Rodighiero) (Fig. 174) is a moderately involute
species with a fairly narrow umbilicus. About 25 primary ribs on the outer
whorl terminate in prominent bullae on the umbilical shoulder, from which
arise 3—5 prorsiradiate secondaries with 1-3 intercalated ribs between bundles.
There are 16 secondaries per 3 bullae, with about 125 ribs across the venter of
the outer whorl. This species differs from O. astieriformis in being more finely
and densely ribbed, with more numerous umbilical bullae. It is, therefore,
close to O. sayni.

Olcostephanus destefanii (Rodighiero, 1919) is a finely ribbed species, allied
to O. sayni, which is yet to be figured. Between 14 and 15 primary ribs terminate
in tubercles on the umbilical shoulder and give rise to bundles of fine, slightly
flexuous secondaries, with intercalated ribs between bundles. It differs from
O. astieriformis in its finer, denser, flexuous secondary ribbing, in which respect
it approaches O. sayni gerecseiensis Somogyi.

Fig. 175. Olcostephanus (Olcostephanus) elongatus (Tzankov). The holotype from the Upper
Valanginian of Placovo, Bulgaria (after Tzankov 1943). x 4.

ANNALS OF THE SOUTH AFRICAN MUSEUM

326

‘TX “(L061 J9d19quineg
Joye) ving SsIMg oY} WOdJ ‘UloJOY poyeUsIsap ‘odAjoJOa_ OY (4) (AodIOqUINYY) susojif (snUDYda1s0I1Q) SnuBYdajsOyO “OL ‘3I4

REVISION OF LATE VALANGINIAN CEPHALOPODA 327

Fig. 177. Olcostephanus (Olcostephanus) filosus
(Baumberger) (after Bayle 1878), x 1.

Olcostephanus elongatus (Tzankov) (Fig. 175) is based upon a strongly
compressed, crushed juvenile only 20 mm in diameter. There are about 20
primaries on the outer whorl which terminate in bullae on the umbilical shoulder,
from which arise 4-5 prorsiradiate secondaries, some of which occasionally
bifurcate. Parabolae are lacking and the outer whorl shows a rapid increase in
height (? due to crushing). This species is close to O. astieriformis but seems to
be more densely ribbed and is thus doubtfully separable from O. symonensis
(Bose).

Olcostephanus filosus (Baumberger) (Fig. 176) is a rather compressed
macroconch species with a moderately wide umbilicus and steep umbilical walls.
About 26 slightly rursiradiate primary ribs terminate in bullae on the umbilical
shoulder and give rise to bundles of 9-10, very fine, prorsiradiate secondaries
between which are fine intercalatories. Parabolae apparently lacking. This
species is readily distinguishable from O. astieriformis by its much denser, finer
ribbing. Although O. filosus has somewhat more primary ribs than O. sayni,
individuals such as that figured by Bayle (1878) (Fig. 177 herein) suggest that
this character is somewhat variable and population studies are likely to show
that O. filosus is a junior subjective synonym of O. sayni. Unfortunately the
concept of O. filosus has become somewhat confused by the inclusion of the
specimen figured by Matheron (1878) as Ammonites mittreanus [sic] d’Orbigny
into Baumberger’s (1907) species. Matheron’s (1878) example (Fig. 178) is

ANNALS OF THE SOUTH AFRICAN MUSEUM

328

Fig. 178. Olcostephanus (Olcostephanus) scissus (Baumberger). Matheron’s

oft-quoted figure of O. mittreanus d’Orbigny, based on a specimen from the

south of France, and assigned by most authors to O. filosus (Baumberger)

(after Matheron 1878). x 1.

Let

van
=
eee

er, oy
pas erase! or

ors
ee

Als ee

ens

crs TI

PETS een
a .

Fig. 179. Olcostephanus (Olcostephanus) paronae (Rodighiero). The holotype,

by monotypy, from Venice (after Rodighiero 1919). x 1.

REVISION OF LATE VALANGINIAN CEPHALOPODA 329

easily distinguished, from both O. sayni and O. filosus, in the presence of only
14 umbilical bullae on the outer whorl. This individual should, therefore, be
assigned to O. scissus (Baumberger).

Olcostephanus paronae (Rodighiero) (Fig. 179) is another species which is
closely allied to O. sayni. It shows marked egression of the umbilical seam on
the outer (?final) whorl so that about 40 per cent of the preceding whorl is
visible in the umbilicus. The umbilical wall is steep on the inner whorls. The
final whorl is ornamented with about 20 umbilical tubercles from which arise
bundles of 3-4 prorsiradiate secondary ribs which frequently bifurcate. On the
adoral half of the outer whorl (?the body chamber), nearly every long rib
bifurcates and has an adjacent intercalated rib high up on the flank. The finer,
denser secondaries, and their frequent bifurcation, distinguish this species from
O. astieriformis.

Olcostephanus potosinus Castillo & Aguilera (Fig. 180) was based upon
two individuals, without type designation. Consequently, the original of the
specimen here figured is selected as lectotype. This specimen is a strongly
compressed, crushed, individual which shows frequent umbilical bullae from
which arise 3-6 prorsiradiate secondaries so that there are about 120-130
secondary ribs across the venter of the lectotype. This species is close to
O. astieriformis but seems to be more finely ribbed and, as such, probably
has priority over O. symonensis (Bése) from the same region.

Olcostephanus quadriradiatus Imlay (Fig. 181) has an ovate, compressed
whorl section with a moderately narrow umbilicus and vertical umbilical walls.
Primary ribs terminate in 23 prominent umbilical bullae on the outer whorl
from which arise bundles of 3-5 slightly prorsiradiate, almost radial secondaries
with 1-2 intercalated ribs between bundles. This is a macroconch species in
which parabolae are apparently lacking. This species differs from O. astieri-
formis only in the almost radial direction of its secondary ribs. Population studies
may show that this character is not of specific importance.

Olcostephanus raricostatus (Bose) (Fig. 182) is a compressed species with
a narrow umbilicus and arched venter. Rursiradiate primaries terminate in
18—23 bullae on the umbilical shoulder, from which arise 3-4 slightly flexuous,
prorsiradiate secondary ribs which frequently bifurcate. Parabolae are
apparently lacking. The distant ribbing of this species is distinctive and it seems
likely that O. huizachensis (Cantu Chapa) is based upon either the microconch
or the inner whorls of the macroconch of O. raricostatus.

Olcostephanus sayni (Kilian) (Fig. 183) is a compressed species with a
moderately narrow umbilicus and a subtrigonal whorl section. The umbilical
seam of the lectotype egresses markedly on the outer whorl, suggesting this is
also the final whorl. The steep umbilical walls are ornamented with about 22
radial primaries which terminate in bullae on the umbilical shoulder and give
rise to bundles of 4-5, fine, prorsiradiate secondaries which frequently bifurcate.
There are 1-2 intercalated ribs between bundles. On the adoral portion of the
outer whorl of the lectotype there are 23 ribs across the venter per 4 umbilical

330 ANNALS OF THE SOUTH AFRICAN MUSEUM ;

Fig. 180. Olcostephanus (Olcostephanus) potosinus Castillo & Aguilera (Q). The lectotyde,
designated herein, from northern Mexico (after Castillo & Aquilera 1895). x 1.

REVISION OF LATE VALANGINIAN CEPHALOPODA 35)

4
” ,
So4i3 v

TAs 4 :
Me n,,

ANS ty

”
cedidiedl et 1d
.

Fig. 181. Olcostephanus (Olcostephanus) quadriradiatus Imlay (2). The holotype from the
Taraises Formation of northern Mexico (after Imlay 1938). x 1.

ANNALS OF THE SOUTH AFRICAN MUSEUM

332

he,

wid Ney at,

reeyy

Fig. 182. Olcostephanus (Olcostephanus) raricostatus (Bose) (2). The lectotype, designated
herein, from the Taraises Formation of northern Mexico (after Bose 1923). x 1.

Fig. 183. Olcostephanus (Olcostephanus) sayni (Kilian). The lectotype from the Lower Haute-
rivian of Castellane, France x 1.

REVISION OF LATE VALANGINIAN CEPHALOPODA 333

nee. Pea
4 “1 > SA Pate
L ‘ y. ze ‘ cay
ny cate ~N oi Mt : vf
Ae St A AAS

Fig. 184. Olcostephanus (Olcostephanus) sayni gerec-
seiensis (Somogyi). The holotype (after Somogyi
1916). x 1.

bullae. Parabolae are lacking. This species may be distinguished from the
present material by its more numerous umbilical bullae and finer, denser
secondary ribs. Somogyi (1916) distinguished O. sayni gerecseiensis (Fig. 184)
on the basis of its flexuous secondary ribbing.

Olcostephanus scissus (Baumberger) (Fig. 185) is a species which has long
been confused with O. filosus and O. sayni, from which it is readily distinguish-
able by its fewer umbilical bullae. It is a compressed form with a narrow umbilicus
and sloping umbilical walls ornamented with 14 radial primaries. These termi-
nate in bullae giving rise to generally 4, fine, prorsiradiate secondaries which
frequently bifurcate high up on the flank. There are frequent intercalated ribs
between bundles and parabolae are apparently lacking. This species is very close
to O. astierianus from which it seems to differ only in possessing bullate umbilical
tubercles and in the frequent bifurcation of the secondary ribs in O. scissus.
O. astieriformis has more numerous bullae and does not show bifurcation of the
secondary ribs.

Olcostephanus subfilosus Spath (Fig. 186) is based upon a nucleus which
bears parabolae and displays about 16 umbilical bullae from which arise
numerous fine, prorsiradiate secondaries, with intercalated ribs between bundles.
A topotype example (Fig. 187) in the Oxford University Museum, OUM-K 1207,
shows a rather narrow, deep, crater-like umbilicus with steep, convex walls
and an evenly rounded umbilical shoulder. Primary ribs begin at, or close to,
the umbilical seam and curve backwards (rursiradiate) to about 20 bullae on
the umbilical shoulder of the outer whorl. Each bulla gives rise to 4-6 prorsi-
radiate secondaries with 1-2 intercalated ribs between bundles. There are
about 125 ribs on the outer whorl. The flanks are convex, converging towards
the narrowly arched venter and giving the whorl section a subtrigonal outline.

334 ANNALS OF THE SOUTH AFRICAN MUSEUM

— Le eee” 5 +.

AvintadSs
x pts RO eta tei a's a:
Cr ais, Hedy tld EE

Fig. 185. Olcostephanus (Olcostephanus) scissus (Baumberger).
The holotype, by lectotype designation herein, from the Swiss
Jura (after Baumberger 1907). x 1.

Fig. 186. Olcostephanus (Olcostephanus) sub-

filosus Spath. The holotype from Speeton,

Yorkshire (after Pavlow & Lamplugh 1892).
<035:

There are two distinct, oblique parabolae on the outer whorl. This species
seems to be based upon a juvenile of O. sayni, although the presence of para-
bolae have still to be proven on the inner whorls of the latter species.
Olcostephanus symonensis (B6se) (Fig. 188) was distinguished from
O. astieriformis by its finer, more numerous secondaries with 60 in a space
where O. astieriformis shows only 45. It is, therefore, close to O. potosinus, as
well as resembling O. sayni in the frequent bifurcation of secondary ribs. With
revision, it seems likely that Bése’s (1923) species will fall into synonymy.
There is in the Natural History Museum in Paris a cast of the original of
Paquier’s (1900) Holcostephanus variegatus (Fig. 189). It shows the original to

REVISION OF LATE VALANGINIAN CEPHALOPODA 31305)

Fig. 187. Olcostephanus (Olcostephanus) subfilosus Spath. A topotype example in the
Oxford University Museum. Note the subtrigonal whorl section, numerous bullae and
parabolae. This species may be based on the inner whorls of O. sayni (Kilian). x 2.

be poorly preserved and abraded, so much so that it is difficult to discern many
of the umbilical bullae and ribs on the outer whorl and these could not be
counted. What can be seen of the ribbing shows rather fine secondaries, about
as wide as the interspaces, which frequently bifurcate on the flanks. There is a
prominent parabola near the adoral end of the outer whorl, which displays a
subtrigonal whorl section. This species is close to O. sayni but with distinctly
coarser ribbing. It can be distinguished from O. scissus and O. astieriformis
by the presence of parabolae.

Occurrence
This species is currently known only from northern Mexico and South

Africa.

Olcostephanus (Olcostephanus) coahuilensis Imlay, 1938 (9)
Figs 190A-B, 191
Olcostephanus coahuilensis Imlay, 1938: 553, pl. 1 (figs 1-3).

Material
A single macroconch, SAM—PCU1550, retaining recrystallized test.

Holotype

The original of Olcostephanus coahuilensis Imlay (Fig. 191) from the
Taraises Formation of northern Mexico.

336 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 188. Olcostephanus (Olcostephanus) symonensis (Bose) (2).
The lectotype, designated herein, from the Taraises Formation of
northern Mexico (after Bose 1923). x 1.

Spl

REVISION OF LATE VALANGINIAN CEPHALOPODA

slieg ‘wnesnyy A1O}SIF{ [eINIeN 9Y} Ut odAJO[OY 9} Jo ysvd VY ‘(JoINbeg) Smppsal4vA (snuvydajsoz1Q) snuvydejsoz]O “681 “SIA

338

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 190. A-B. Olcostephanus (Olcostephanus) coahuilensis Imlay (¢). Lateral and front views

of SAM-—PCU1550, showing moderate inflation and fine, radial secondaries. x 0,44. C—D.

Olcostephanus (Olcostephanus) cf. perinflatus (Matheron) (2). Ventral and lateral views of

AM-4292b. Note small umbilicus, extreme inflation, fine secondaries occasionally bifurcating,
and apparent absence of parabolae. x 0,75.

REVISION OF LATE VALANGINIAN CEPHALOPODA 339

Description

The shell is a large, inflated cadicone, involute up to the umbilical bullae,
except on the final whorl when the umbilical seam egresses somewhat. The
umbilicus is narrow with moderately steep, convex umbilical walls, the latter
ornamented with 17 rursiradiate primaries which terminate in bullae on the
umbilical shoulder. From these arise 4-5 radial secondaries, commonly with
1-2 intercalatories between bundles. There are 32 secondaries per 5 bullae on
the outer whorl, with 11 ribs in a 50 mm distance along the venter, and about
50 ribs per half whorl. Maximum inflation is attained about half a whorl
behind the peristome.

Measurements
No. D H Wi W/H Uo Ui
SAM-PCU 1550 195 We) 90 1,20 87 52 (27)

tn
ws,

~ po SP eee
e eee,

gt ee
ater , Meany
“2h oeis

Fig. 191. Olcostephanus (Olcostephanus) coahuilensis (Imlay) (¢). The holotype from the
Taraises Formation of northern Mexico (after Imlay 1938). x 1.

340 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 192. Olcostephanus (Olcostephanus) guebhardi (Kilian). The holotype from Escragnolles,
France (after Kilian 1902). x 1.

Discussion

The Uitenhage example differs from Imlay’s type only in having slightly
fewer umbilical bullae per whorl, a difference which is not considered of specific
importance.

Olcostephanus guebhardi (Kilian) (Fig. 192), of which O. sharpei (Kara-
kasch) (Fig. 193) is a synonym, closely resembles the South African material,
but is much smaller and does not show the inflation seen in the present shell
The differences may well be those between dimorphs, but until the European
material is revised, the writer prefers to use Imlay’s name. Olcostephanus
actinotus (Baumberger) also resembles the present species, but has a very
different, subtrigonal whorl section.

Occurrence

Olcostephanus coahuilensis is currently known only from Mexico and South
Africa.

Olcostephanus (Olcostephanus) cf. perinflatus (Matheron, 1878) (Q)

Figs 151E—-F, 190C—D, 194-195
Compare
Ammonites perinflatus Matheron, 1878: pl. 20B (fig. 8a—b).
Ammonites stephanophorus Matheron, 1878: pl. 20B (fig. 9).

REVISION OF LATE VALANGIiNIAN CEPHALOPODA 34]

Fig. 193. Olcostephanus (Olcostephanus) guebhardi

(Kilian). The holotype of Astieria sharpei Karakasch

from the Upper Valanginian of the Crimea (after
Karakasch 1907). x 1.

Fig. 194. Olcostephanus (Olcostephanus) perinflatus (Matheron) (2). The holotype, by mono-
typy, from the south of France (after Matheron 1878). x 1.

342 ANNALS OF THE SOUTH AFRICAN MUSEUM

Olcostephanus perinflatus (Matheron) Spath, 1939: 23. Fatmi, 1977: 270.
Olcostephanus cf. perinflatus (Matheron) Spath, 1939: 25, pl. 6 (fig. 6).

Material

Two specimens, AM-—4292b, and PEM-1468/89, the latter a mere fragment,
both preserved as internal moulds.

Holotype

The original of Matheron’s (1878) plate 20B (fig. 8a—b) (Fig. 194 herein)
from the south of France.

Description

The shell of AM-4292b is an extremely inflated, globose, cadicone, with a
very narrow, deep, crater-like umbilicus. The umbilical walls are steep, convex,
and slightly overhanging, and ornamented with about 22 rursiradiate primaries
terminating in small bullae on the umbilical shoulder. The latter give rise to
3-4 rather fine, slightly prorsiradiate secondaries, which on the outer whorl
are slightly flexuous, and between which are generally a single intercalated rib.
Very occasionally a secondary rib is seen to bifurcate on the flanks, so that there
are about 50-60 secondaries per half whorl on the outer whorl. Parabolae are
absent on the outer whorl, but it is unknown whether they occur on the inner
whorls.

Discussion

As suggested by Kilian (1892), the writer believes Ammonites stephanophorus
Matheron (Fig. 195) merely to represent the juvenile growth stages of Olco-
stephanus perinflatus. The Uitenhage specimens are only tentatively referred to
Matheron’s species because of uncertainty as to what O. perinflatus really
looks like, since it has never been photographically refigured or redescribed
and there is reason to believe that Matheron’s (1878) illustration may be some-
what idealized.

Fig. 195. Olcostephanus (Olcostephanus) perinflatus

(Matheron) (2). The holotype of Olcostephanus

stephanophorus (Matheron) from the south of France,
a juvenile (after Matheron 1878). x 1.

REVISION OF LATE VALANGINIAN CEPHALOPODA 343

Fig. 196. Olcostephanus (Olcostephanus) balkanicus (Tzankov). The syntypes
from the Lower Hauterivian of Bulgaria (after Tzankov 1943). x 3.

Olcostephanus globosus Spath, of which O. pachycyclus Spath represents
an early growth stage (Fatmi 1977), is a macroconch species which is
undoubtedly close to O. perinflatus. Indeed, it would seem to differ from
Matheron’s (1878) illustration only in its larger adult size, a character which is
possibly environmentally related and hence doubtfully of specific importance.
However, the Pakistan material is retained as distinct until Matheron’s (1878)
species is restudied on the basis of type and topotype material.

Olcostephanus balkanicus (Tzankov) (Fig. 196) is based upon a strongly
inflated nucleus which seems to differ from the inner whorls of O. perinflatus,
that is from O. stephanophorus, in having a much wider umbilicus, somewhat
fewer (15-17) umbilical bullae and distinctly prorsiradiate secondaries. How-
ever, it shows prominent parabolae.

Occurrence

Olcostephanus perinflatus is at present known with certainty only from the
south of France, but it may also be present in South Africa and Pakistan.

344 ANNALS OF THE SOUTH AFRICAN MUSEUM

Family Berriasellidae Spath, 1922
Subfamily Neocomitinae Spath, 1924
Genus Neohoploceras Spath, 1939

Type species Ammonites submartini Mallada, 1882;
by original designation of Spath, 1939

Neohoploceras subanceps (Tate, 1867)

Fig. 197

Ammonites subanceps Tate, 1867: 150, pl. 7 (fig. 3a—b).

Reineckia subanceps (Tate) Newton, 1896: 150.

? Leopoldia depereti Sayn, 1907: 59, pl. 4 (figs 6-7).

Solgeria subanceps (Tate) Spath, 1930: 151, pl. 13 (fig. 4a-c). Du Toit, 1954: 384.
Neohoploceras subanceps (Tate) Klinger & Kennedy, 1979: 18.

Material

The holotype, BM—C32197, from the Sundays River is the only undoubted
specimen known.

Holotype

By monotypy, the original of Ammonites subanceps figured by Tate (1867:
150, pl. 7 (fig. 3a—b)), from the Sundays River, and now in the British Museum,
BM-C32197.

Fig. 197. Neohoploceras subanceps (Tate). The holotype,
BMNH-C32197, from the Sundays River Formation. x 2.
Photo: W. J. Kennedy.

Diagnosis

Small, somewhat inflated, with 12-14 ribs beginning at the umbilical seam
of the outer whorl, each rib with a weak umbilical bulla and terminating in a
swollen tubercle at about mid-flank, from which prorsiradiate ribs bifurcate
or trifurcate. There are occasional single and intercalated ribs. All ribs terminate
in small ventrolateral tubercles and are interrupted across the venter by a smooth
zone. Constrictions are present.

REVISION OF LATE VALANGINIAN CEPHALOPODA 345

Description

The following description is based on a plastercast of the type sent to the
writer by M. K. Howarth: the specimen is small, but appears to have recrystal-
lized test preserved. The shell is somewhat compressed, with a whorl section
about as wide as high. The umbilicus is rather shallow and moderately evolute,
the outer whorl covering slightly more than half of the preceding whorl.
Maximum width is at mid-flank. The umbilical wall is gently sloping, with a
well-rounded umbilical shoulder.

On the outer whorl, about 12-14 ribs begin at the umbilical seam and pass
almost radially outwards to the umbilical shoulder where they develop small
but distinct umbilical bullae. From here the ribbing is slightly prorsiradiate,
and commonly swells at about mid-flank into a lateral tubercle, from which
ribs frequently bifurcate or trifurcate. Occasionally only a single rib arises
from the mid-lateral tubercle, in which case the latter is usually weakly
developed, while there is also the odd intercalated rib. Ribbing is very weakly
developed across the venter on the adoral portion of the outer whorl, and gives
the impression that the smooth zone may have disappeared with age. There
appear to be two constrictions on the outer whorl.

Measurements

No. D H Wi W/H Uo Ui
BM-C32197 16 8 acs 0,94 4. 5G)
Discussion

The small size of the holotype, which represents a juvenile growth stage,
does not allow for proper comparison with other species, especially since
juveniles are rarely figured or described. One of the few exceptions is the work
of G. Sayn (1907). Amongst the examples figured by Sayn, N. depereti and
N. provinciale (Sayn) both bear a close resemblance to Ammonites subanceps.

Neohoploceras depereti (Sayn) is moderately inflated and bears prominent -
constrictions. The ribbing is of two types, simple ribs lacking tubercles and
stronger ribs which bifurcate or trifurcate from a prominent lateral tubercle
and also have umbilical tubercles. All ribs bear weak ventrolateral clavi. The
venter is grooved and smooth. The only difference between N. depereti and
N. subanceps appears to be in the possession of more prominent tubercles by
the former. However, since the larger of Sayn’s syntypes is 33 mm in diameter,
as against only 20 mm for N. subanceps, the differences are probably only
ontogenetic. :

Neohoploceras provinciale (Sayn) is similar to N. subanceps but is even
larger than N. depereti and consequently comparison is still more difficult. It
is, however, more closely ribbed than N. depereti, although its validity will be
resolved only with a revision of the French material.

It is not possible to compare satisfactorily the South African species with

346 ANNALS OF THE SOUTH AFRICAN MUSEUM

the rich Madagascan faunas due to the size differences. Such a comparison
will have to await ontogenetic studies on the Madagascan material.

Occurrence

Neohoploceras subanceps is present in the Sundays River Formation and
perhaps the zone of Saynoceras verrucosum in south-east France.

Genus Distoloceras Hyatt, 1900

Type species Ammonites hystrix Phillips, 1829;
by original designation of Hyatt, 1900

Distoloceras spinosissimum (Hausmann, 1837)

Figs 198-200

Ammonites spinosissinum Hausmann, 1837: 1458.

Crioceras spinosissimum (Hausmann) Holub & Neumayr, 1882: 273, pl. 1 (fig. la—c). Kitchin,
1908: 225.

? Distoloceras cf. spinosissimum (Hausmann) Spath, 1924: 75.

Distoloceras spinosissimum (Hausmann) Spath, 1930: 152, pl. 13 (fig. 1). Besairie, 1932: 44,
pl. 16 (fig. 13). Du Toit, 1954: 384. Collignon, 1962: 51, fig. 887.

Distoloceras hirtzi Collignon, 1962: 32, pl. 185 (figs 846-849).

Material

The holotype, in the Greifswalde Universitats-Museum, and two examples
in the British Museum (BM—C32194, C10819) are the only specimens so far
recorded from the Sundays River Formation.

Holotype

By monotypy, the original of Crioceras spinosissimum (Hausmann) figured
by Holub & Neumayr (1882: 273, pl. 1 (fig. la-c)) from the Sundays River.

Diagnosis

Only uncoiled fragments of this species are known from the Uitenhage
Group. At this stage the whorl section is polygonal, slightly compressed, with
flat flanks. Ornament comprises strong radial ribs ornamented with umbilical,
lateral and ventrolateral tubercles, between which are intercalated varying
numbers of weaker, more flexuous ribs which may or may not have lateral and
ventrolateral tubercles. The umbilical and ventrolateral tubercles frequently
form spines. Ribbing joining the ventrolateral spines across the venter is convex
adorally.

Description

This is a large species of Distoloceras of which only uncoiled fragments
are currently known from the Sundays River Formation.
In the holotype, judging from Holub & Neumayr’s figure, the adapical

REVISION OF LATE VALANGINIAN CEPHALOPODA

347

lver

(Hausmann). The holotype from the Sundays R

Formation (after Holub & Neumayr 1882). x 0,75.

nostisstmum

Fig. 198. Distoloceras sp

348 ANNALS OF THE SOUTH AFRICAN MUSEUM

portion of the outer whorl, which has already lost contact with the previous
whorl, shows simple straight, uniformly developed radial ribbing, with umbilical
and lower ventrolateral swellings (corresponding to the lateral tubercles on the
adoral portion), although Spath (1930: 152) thought this was possibly due to
corrosion.

Beyond a diameter of about 100 mm intercalated ribs start appearing,
always lacking umbilical tubercles, but occasionally with swelling corresponding
to the lateral tubercles. These intercalatories commonly arise at the umbilical
shoulder, although some may be intercalated half-way up the flank, and vary
considerably in strength. |

With the appearance of intercalated ribs, the ventrolateral and umbilical
tubercles frequently form long spines. These ventrolateral spines may also be
developed on intercalated ribs.

Ribbing weakens somewhat across the tabulate venter and is convex
adorally. The suture line is complex.

A cast of the specimen figured by Spath (1930: pl. 13 (fig. 1)) (BM—C32194),
provided by M. K. Howarth, shows the following features:

Whorl compressed, polygonal in costal section, elliptical intercostally.
Dorsum with a weak median furrow, from which at least some of the ribs arise
and pass backwards (rursiradiate) to the dorsal (umbilical) shoulder. On the
adapical portion of this uncoiled fragment, the flank ribs are very slightly
prorsiradiate. Some are more prominent than others and show faint umbilical
bullae, midlateral, and ventrolateral tubercles. Between these prominent ribs
are intercalated finer ribs which arise above the level of the dorsal shoulder
and show weak midlateral bullae and ventrolateral swellings. All ribs bend
sharply forwards at the level of the midlateral tubercles. On the adoral portion
of this specimen the main ribs all possess well-developed umbilical, midlateral,
and ventrolateral spines, and are separated by 3-4 much finer, slightly flexuous
ribs with small midlateral bullae and ventrolateral swellings. The ribbing is
poorly developed across the venter.

Discussion

In describing a fragment of a tightly coiled form from the Upper
Valanginian of Ambiky, Madagascar, Collignon (1962) considered this species
to be characterized by *. . . section subcarrée, ornamentation de cétes a tubercles
ou les externes acquierent la preponderance’.

Comparison of D. spinosissimum with coiled species of Distoloceras is
most unsatisfactory and consequently the validity of this species will have to
await the further collection of topotype material. It should be noted, however,
that both Besairie (1932) and Collignon (1962) consider fragments of this
species, which occurs plentifully in Madagascar, to be easily recognizable and
typical.

Distoloceras hystrix (Phillips) would seem to be very similar, but with a
more compressed whorl section (although it seems highly likely that the whorl

REVISION OF LATE VALANGINIAN CEPHALOPODA 349

= 199. Distoloceras spinosissimum (Hausmann). Ventral and lateral views of BMNH-—C32194,
figured by Spath (1930). x 0,66. Photo: W. J. Kennedy.

350 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 200. Distoloceras spinosissimum (Hausmann). Ventral and lateral views of BMNH-—
X108A, showing well-developed main ribs and thus very close to D. hirtzi Collignon. x 0,66.
Photo: W. J. Kennedy.

section would change with uncoiling), while prior to the appearance of inter-
calated ribs in D. hystrix, the ribbing is seen to arise in pairs from umbilical
tubercles, whereas in D. spinosissimum they are simple.

Distoloceras laticostatum Imlay is known only from tightly coiled, pyritic
specimens. It is more compressed, with closer, more strongly developed ribs
which may bifurcate at either the umbilical shoulder, or at mid-flank. Bifurca-
tion of ribbing has not been recorded from D. spinosissimum. Distoloceras

REVISION OF LATE VALANGINIAN CEPHALOPODA 351

capulinense Imlay is very compressed, with closer, more regular ribs, some of
which branch at the lateral tubercles.

Distoloceras hirtzi Collignon, from the Lower Valanginian of Madagascar,
is very similar to D. spinosissimum, but was said to differ in having a more
elliptical whorl section, prorsiradiate and not radial ribs, and in having more
strongly developed midlateral tubercles. These are the features shown by
Spath’s topotype example, BM—C32194, and any differences are probably due
to the better preservation of the Madagascan material. Consequently, D. hirtzi
is considered a junior subjective synonym of D. spinosissimum.

Occurrence

Distoloceras spinosissimum is known from the Upper Valanginian of
South Africa, Madagascar, and possibly England.

Distoloceras cf. irregulare Imlay, 1938
Figs 201-202

Compare
Distoloceras irregulare Imlay, 1938: 577, pl. 14 (figs 3, 5, 8-11).

Material .
A single specimen, SAM—PCU1613, from an unknown locality.

Holotype

The original of Imlay’s (1938) plate 14, figures 8-11, from the Taraises
Formation of northern Mexico.

Description

In this fragment of an uncoiled example, the whorl section is subtrigonal,
the flanks converging to the narrow, arched venter, the greatest width being at
the dorsolateral shoulder. The dorsum, which was not in contact with the pre-
vious whorl, shows a furrowed median ridge from which initially radial, but
soon becoming strongly rursiradiate, growth lines arise. Also on the dorsum,
and arising from this median ridge, are very weakly developed rursiradiate
ribs which strengthen towards the dorsal shoulders where they form distinct
bullae. On the flanks the ribbing is rather variable, generally being prorsiradiate,
although one rib is radial, and bending forwards near the venter. Between main
ribs are 2—4 intercalated ribs which are prominent only across the venter. The
tuberculation on the ribbing is rather variable and asymmetrical. There may be
umbilical, lateral, ventrolateral and siphonal tubercles. A spine on one flank
may be completely absent on the opposite side, or represented only by a weakly
developed node. Siphonal tubercles are not developed on all ribs and may
occasionally be eccentrically placed. The strength of the ribbing is also highly
variable, fading and swelling at random. Ribs occasionally bifurcate from a
lateral node.

352 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 201. Distoloceras cf. irregulare Imlay. Front, ventral and lateral views of an uncoiled
fragment, SAM-—PCU1613. x 0,66. :

Discussion

The solitary fragment from the Sundays River Formation is very close to
D. irregulare Imlay, the latter differing only in having an elliptical whorl section
and almost radial ribbing. It would seem that the development of siphonal
spines is an ontogenetic feature as evidenced in the holotype of D. irregulare.

This specimen differs from D. spinosissimum (Hausmann) in having a
subtrigonal and not polygonal whorl section, in possessing siphonal tubercles,
and in the lack of regularity in the tuberculation.

Occurrence

Distoloceras irregulare Imlay is recorded from the middle part of the Upper
Member of the Taraises Formation in Mexico, considered by Imlay (1938) to
be of Lower Hauterivian age, and may also be present in the Sundays River
Formation.

EE —————

REVISION OF LATE VALANGINIAN CEPHALOPODA 353

Fig. 202. Distoloceras cf. irregulare Imlay. Whorl
section of SAM-—PCU1613. x 0,66.

Superfamily HOPLITACEAE Douvillé, 1890
Family Desmoceratidae Zittel, 1895
Subfamily Eodesmoceratinae Wright, 1955
Genus Eodesmoceras Spath, 1924

Type species Ammonites celestini Pictet & Campiche, 1858;
by original designation of Spath, 1924

Discussion

Two subgenera are recognized within this genus (Wright in Arkell et al.
1957), viz. E. (Eodesmoceras) and E. (Miodesmoceras). Miodesmoceras is a
Barremian form, distinguished from Eodesmoceras in being more compressed
and in lacking constrictions, whilst the latter is considered of Valanginian—
Lower Hauterivian age. Eodesmoceras haughtoni Spath from the Uitenhage
Group lacks constrictions, and is much more compressed than the type of
E. (Eodesmoceras) celestini (Pictet & Campiche). Consequently, the differences
between these two subgenera blurr and they appear to be of little taxonomic use.

Eodesmoceras haughtoni Spath, 1930
Fig. 203
Eodesmoceras haughtoni Spath, 1930: 141, pl. 13 (fig. 2a-e). Du Toit 1954: 384.

Material
The holotype, SAM-—227, is the only specimen known.

354 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 203. Eodesmoceras haughtoni Spath. Ventral, front and
lateral views of the holotype, SAM-227, preserved as an
internal mould. x 2.

Holotype

By monotypy, the original of the specimen figured by Spath (1930, pl. 13
(fig. 2a-e)) from ‘... Shore of pan, Zoutpan, Uitenhage’, and now in the South
African Museum.

Diagnosis

A compressed, immature Eodesmoceras in which the only ornament is
very faint sigmoidal growth striae. Constrictions are lacking.

Description

The holotype is small, preserved as an internal mould, and rather involute,
the outer whorl covering about two-thirds of the preceding whorl. The shell is
strongly compressed with broad, flat flanks converging slightly to the narrow
evenly rounded venter. At 14 mm diameter the specimen is still entirely septate.
Constrictions are lacking, whilst Spath presumably observed the growth striae
on the inner whorls where the shell material was preserved. The specimen has
since been glued together and this feature could not be verified.

Measurements

No. D H Wi W/H Ui
SAM-227 14 8 5 0,63 3 (21)
Discussion

Despite the fact that Spath (1930: 142) considered this species ‘. . . probably
represents merely the inner whorls of a larger form like Eodesmoceras celestini
(Pictet & Campiche)’, he still described it as a new species.

The very small size of E. haughtoni makes comparison difficult. It is more
compressed than the type of E. celestini, but this is possibly ontogenetic variation
since the latter is 28 mm in diameter. According to Wright’s (in Arkell et al.

REVISION OF LATE VALANGINIAN CEPHALOPODA 355

1957) diagnosis of E. (Eodesmoceras), presumably based on the type species
E. celestini, constrictions are present although they are not visible in the
illustration of the lectotype. Lack of comparative material does not allow for a
definite statement on the validity of Spath’s species.

Occurrence
This species is known only from the Sundays River Formation.

Subclass DIBRANCHIATA Owen, 1832
Order DECAPODA Leach, 1818
Suborder BELEMNOIDEA Naef, 1912
Family Belemnitidae D’Orbigny, 1845
Subfamily Belemnopsinae Naef, 1922
Genus Belemnopsis Bayle, 1878
Type species Belemnites sulcatus Miller;
by subsequent designation of Douvillé, 1879
Belemnopsis africana (Tate, 1867)

Fig. 204

Belemnites africanus Tate, 1867: 151, pl. 7 (fig. 2). Kitchin, 1908: 225.
Belemnopsis africana (Tate) Spath, 1930: 155. Besairie, 1930; pl. 11 (fig. 12). Spath, 1939,

pl. 24 (fig. 15). Stevens, 1965: 164.
Non Belemnopsis africana (Tate) Besairie, 1930, pl. 23 (figs 6-7, 20-21).

Material
A single unnumbered specimen in the South African Museum.

Fig. 204. Belemnopsis africana (Tate).

Lateral and ventral views of an apical

fragment in the South African Museum.
Scnll:

356 ANNALS OF THE SOUTH AFRICAN MUSEUM

Holotype

By monotypy, the original of the specimen figured by Tate (1967, pl. 7
(fig. 2)), from the Sundays River, now in the British Museum.

Diagnosis
Guard non-hastate, expanding slowly adorally. Ventral groove deep and
broad, prominent throughout growth. Cross-section depressed.

Description

The following description, based on the holotype, is taken from Stevens
(1965: 164): ‘...the guard is non-hastate, its diameter gradually increasing
forwards. The ventral groove, broad and deep, is very prominent throughout
the growth stages of the guard. Cross-sections throughout the length of the
guard are depressed (maximum transverse diameter, 19 mm: maximum sagittal
diameter, 18 mm).’

Discussion

The genus Belemnopsis is characteristic of the Kimmeridgian—Tithonian of
the Indo-Pacific, with a relict fauna surviving in South Africa and Madagascar
until the late Valanginian. Spath (1930: 156) remarked that an alveolar fragment
of B. africana could not be distinguished from B. gerardi Oppel (= B. uhligi
Stevens).

The examples referred by Besairie (1936) to this species are considered by
Stevens (1965) as probably distinct. They differ from Tate’s species in being
more elongate and slender, with an elongate sharply pointed apical region,
while the ventral groove is not as deep.

The rarity of this species in the Uitenhage Group led Stevens (1965: 164)
to ‘... suggest derivation from Upper Jurassic strata, since removed by erosion.
However, the holotype is quite well preserved and not corroded so is probably
not derived from older strata.’

Belemnopsis gladiator Willey (1973: 33, fig. 2) from the Berriasian of
Antarctica is very close to Tate’s species. According to Willey, B. africana
differs in being somewhat shorter, comparatively more robust, with a deeper,
flat-bottomed ventral groove with concave sides and sharp margins. The
differences are, however, slight and since B. africana is still known only from a
handful of fragmentary specimens, probably would not stand up were a popu-
lation of individuals known. This is supported by the fact that Willey (1973)
assigned an apical fragment of Belemnopsis from the Sundays River Formation,
in the British Museum, BM-—C6217, to B. gladiator. Belemnopsis africana may
prove, therefore, to be longer ranging than is generally suspected.

Occurrence

Belemnopsis africana is known from the Lower and Upper Valanginian of
Madagascar, the Upper Valanginian of South Africa, and possibly the Berriasian
of Antarctica.

REVISION OF LATE VALANGINIAN CEPHALOPODA 357

Order NAUTILOIDEA Agassiz, 1847
Suborder NAUTILINA Agassiz, 1847
Family Nautilidae de Blainville, 1825
Genus Eutrephoceras Hyatt, 1894

Type species Nautilus dekayi Morton, 1834;
by subsequent designation of Hyatt, 1894.

Eutrephoceras uitenhagense Spath, 1930
Fig. 205
Nautilus sp. Sharpe, 1856: 201. Kitchin, 1908: 225.
Eutrephoceras uitenhagense Spath, 1930: 139.
Nautilus (Eutrephoceras) 2uitenhagense Spath, Besairie, 1936: 145.

Material

The holotype is still the only specimen recorded from the Sundays River
Formation.

Holotype

The specimen recorded by Sharpe (1856: 201) from the Sundays River
now in the British Museum (BM-11034, Geol. Soc. Coll.).

eee

Fig. 205. Eutrephoceras uitenhagense Spath. Whorl section and
suture (after Spath 1930). x 0,66.

358 ANNALS OF THE SOUTH AFRICAN MUSEUM

Diagnosis |

‘Coiling occlusal, with umbilicus nearly closed. Whorl section rounded,
slightly compressed at first, later flaring, with greatest thickness at inner third
and no edge to high umbilical wall. Suture line with slight umbilical saddle and
shallow lateral lobe, straight across venter. Annular lobe strongly developed.
Test entirely smooth, thick’ (Spath 1930: 139).

Measurements

No. D H W . W/H U
BM-11034 135 55 74 1,35 7(5)
Discussion

The excentric position of the ventral siphuncle in this species was thought
not to be significant by Spath (1930). He compared this species with ‘Nautilus’
boissieri Pictet, from which E. uitenhagense was distinguished by its greater
inflation.

Occurrence

This species is known only from the Sundays River Formation and perhaps
Madagascar.

-SUMMARY

The cephalopod fauna from the Sundays River Formation is revised and
shown to comprise the following species:

AMMONOIDEA

Partschiceras rogersi (Kitchin)

Bochianites glaber Kitchin

Bochianites africanus (Tate)

. (Olcostephanus) atherstoni (Sharpe) (¢ and. )

. (Olcostephanus) densicostatus (Wegner) sp. juv.

. (Olcostephanus) rogersi (Kitchin) (¢ and )

. (Olcostephanus) victoris Spath (Q)

. (Olcostephanus) fascigerus Spath (Q)

. (Olcostephanus) aff. durangensis (Cantu Chapa) (3)
. (Olcostephanus) baini baini (Sharpe) (¢ and 9)

. (Olcostephanus) baini var. sphaeroidalis (Spath) (3g and 9)
. (Olcostephanus) ventricosus (Von Koenen) (?9)

. (Olcostephanus) uitenhagensis (Kitchin) (2)

. (Olcostephanus) riccardii sp. nov. (Q)

SS) 97S) S559 7S79) S59

REVISION OF LATE VALANGINIAN CEPHALOPODA 359

O. (Olcostephanus) astieriformis (Bose) (2)

O. (Olcostephanus) coahuilensis Imlay (9)

O. (Olcostephanus) cf. perinflatus (Matheron) (2)
Neohoploceras subanceps (Tate)

Distoloceras spinosissimum (Hausmann)
Distoloceras cf. irregulare Imlay

Eodesmoceras haughtoni Spath

COLEOIDEA

Belemnopsis africana (Tate)
Belemnopsis gladiator Willey

NAUTILOIDEA
Eutrephoceras uitenhagense Spath

The fauna is dominated by species of Olcostephanus, within which sexual
dimorphism is recognized for the first time from these beds. The fact that three
morphologic types—microconch, macroconch, and inner whorls of the macro-
conch—may be distinguished within a single species (dimorphic pair), together
with occasional gerontic individuals, has led to a proliferation of nominal
species. This, together with complete disregard for intraspecific variation, has
led to taxonomic chaos.

A literature review of sexual dimorphism in ammonites reveals that the
inner whorls of many macroconchs (Q) are indistinguishable from their micro-
conch (3g) counterparts. This, together with the occurrence of rare aberrant
mutants (Cope 1967: 53), supports the contention that some sexually dimorphic
ammonites display consecutive hermaphroditism or protandrism.

Many microconch species of Olcostephanus exhibit parabolae, as do the
immature growth stages of the corresponding macroconch. Not only are
parabolae virtually identical, morphologically, to the peristome but they also
have an identical mode of formation. Consequently, parabolae in Olcostephanus
are interpreted as relict peristomes. Accepting this suggestion, it is of interest
to note that the whoris of many extant gastropods are ornamented with varices,
representing the position of relict apertures developed during pauses in growth.
They would appear, therefore, to be homologous to the parabolae occurring
in Olcostephanus. The fact that in extant Gastropoda varices are known to be
of specific importance supports the contention that they were of equal sig-
nificance in Olcostephanus.

A significant feature associated with sexual dimorphism in Olcostephanus
is the striking degree of convergence in the outer whorls of macroconch forms.
Since O. atherstoni (Sharpe) was one of the earliest such macroconchs to be
described, this name consistently crops up in the older literature when, in fact,
the inner whorls show a different species to be involved. It is probably also this
homoeomorphy which has led to the neglect of parabolae as a specific criterion.

360 ANNALS OF THE SOUTH AFRICAN MUSEUM

The importance of a knowledge of the early ontogenetic stages of macroconchs
for their specific determination cannot be over-emphasized, as well as being
imperative for the recognition of the microconch dimorph. A slight, but
apparently consistent, difference between microconch forms and the inner whorls
of their macroconch appears to be a somewhat higher rib density in the latter.

The subfamily Olcostephaninae is reviewed and considered to include the
following genera and subgenera: Saynoceras (Saynoceras), S. (Ceratotuberculus),
Olcostephanus (Olcostephanus), O. (Subastieria), O. (Parastieria), O. (Jean-
noticeras), O. (Mexicanoceras), Jeanthieuloyites, Valanginites, Capeloites, and
Dobrodgeiceras. The genera Holcostephanus, Astieria, Rogersites, Taraisites and
Satoites are considered junior synonyms of Olcostephanus s.s., whilst Maderia
and Lemurostephanus comprise a heterogeneous assemblage of O. (Olcostephanus)
and perhaps O. (Subastieria) nuclei, and are thus superfluous. In order to
incorporate the new subgenera, the diagnosis of Olcostephanus as given by
Wright (in Arkell et al. 1957) is emended. The new genus Jeanthieuloyites is
proposed for Rogersites quinquestriatus Besairie.

ACKNOWLEDGEMENTS

I should like to express my gratitude to Dr J. Grindley, then Director of
the Port Elizabeth Museum, Dr P. J. Roussouw of the Geological Survey,
Pretoria, and Mr C. F. Jacot-Guillarmod, then Director of the Albany Museum,
Grahamstown, for placing the collections of their respective institutions at
my disposal. I am especially grateful to Dr M. K. Howarth and Mr D. Phillips
of the British Museum for the trouble they went to in supplying me with the
collection of Uitenhage ammonites housed in the British Museum, as well as
casts of type material, and to Mr. C. W. Wright for enabling me to examine
other material. I am also indebted to Prof. H. Eales and Messrs W. Gess,
H. Deacon and B. Every, as well as the quarry owners of the Uitenhage district,
for their help and kindness.

Drs A. C. Riccardi, J. P. Thieuloy, W. J. Kennedy, D. F. B. Palframann,
D. van Z. Engelbrecht, B. F. Kensley, and A. J. Tankard provided helpful
suggestions and criticism, for which I am most grateful.

Without the assistance of Drs A. C. Riccardi, W. J. Kennedy, J. Wiedmann,
H. C. Klinger, M. K. Howarth, R. Busnardo, and J. P. Thieuloy who all went
to a great deal of trouble to supply me with literature which was not available
in the country, this work could never have been completed.

I should like to express my thanks to Mrs S. B. Bruins, former librarian
at the South African Museum, for her efforts in obtaining the literature so
necessary to this study, and to my wife for typing the final draft.

The contents of this paper were submitted for the degree of M.Sc. at the
University of Natal, Durban, in 1973, and the fieldwork was undertaken while
the writer was employed at the South African Museum.

REVISION OF LATE VALANGINIAN CEPHALOPODA 361

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REVISION OF LATE VALANGINIAN CEPHALOPODA 365

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366 ANNALS OF THE SOUTH AFRICAN MUSEUM

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mies:
a ai ¢g

6. SYSTEMATIC papers must conform to the /nternational zi i ;
) (particulary et 2) ind Si). code of zoological nomenclature
__ Names of new taxa, combinations, synonyms, etc., when used for the first time. m
_ followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., nal
_*MOV., SyN. NOv., etc. or
| 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, ie. 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.

i Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
_ Nucula largillierti Philippi, 1861: 87.

, Leda bicuspidata: Nickles, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

i Note punctuation in the above example:

; - comma separates author’s name and year

i 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°51’S 25°39’E), collected by A. Smith, 15 January 1973.

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

7. SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. ‘... the Figure depicting C. namacolus ...’; *. . . in C. namacolus (Fig. 10)...’

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene

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.’ hae

Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.

Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.

é\

M. R. COOPER

REVISION OF THE LATE VALANGINIAN
CEPHALOPODA FROM THE SUNDAYS RIVER
FORMATION OF SOUTH AFRICA, WITH
SPECIAL REFERENCE TO THE GENUS
OLCOSTEPHANUS

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Full text of "Annals of the South African Museum = Annale van die Suid-Afrikaanse Museum"

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